AN ECOLOGICAL STUDY OF LAMINARIA SINCLAIRII
AND L. LONGIPES
by
JAMES W. MARKHAM A.B., Stanford University, Stanford, California, 1961 M.S., University of Washington, Seattle, Washington, 1963
A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
in the Department
of
BOTANY
We accept this thesis as conforming to the required standard
THE UNIVERSITY OF BRITISH COLUMBIA
JUNE, 1969 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and Study.
I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission.
Department of Tetany
The University of British Columbia Vancouver 8, Canada
Date June 10, 1969 ABSTRACT
Laminaria sinclairii (Harvey) Farlow, Anderson and Eaton, found from Southern California to Central British Columbia, and L^ longipes Bory, found from Southeast Alaska to the Kurile Islands, differ in several ways from most other kelp plants. Their most distinctive feature is the rhizome-like holdfast, composed of many haptera, from which arise multiple stipes, each bearing a single blade. The two species are very similar to each other and have been distinguished in the past primarily by the presence of mucilage ducts in the stipe of L. sinclairii and the absence of these in the stipe of L. longipes. In order to determine whether the two species are indeed distinct, their distribution, ecology, growth, and reproduction were studied in the laboratory and on beaches in Alaska, British Columbia, and Oregon.
The gross distribution of both species appears to be controlled by temperature. Transplants and laboratory cultures indicate that L. longipes is adapted to lower temperatures than L. sinclairii. Salinity apparently has little influence on distribution, as both species tolerate wide ranges of salinity.
L. sinclairii was studied in_ situ on three beaches in Northern Oregon, where the plants are subjected to heavy surf. The sand level on the beaches rises through the summer so that the plants are partly or wholly buried under sand by late summer. The first heavy storms in the fall remove most of the sand. Maximum growth occurs in early summer, prior to burial. The blades are lost in December and new ones are regenerated in January. Ripe sori are produced on the old blades just before they are lost and on the new blades just after they appear. There is little evidence from either field or laboratory studies to indicate that the gametophytes which develop from the spores in these sori normally produce sporophytes. Sexual reproduction of this type is difficult because of the scouring action of the sand. In March and April there is considerable production of new stipes and blades from the haptera at the margins of the holdfast. This vegetative proliferation is apparently the normal method of reproduction.
L. longipes was observed in situ in Alaska on only five occasions. Growth is greatest in summer and sori are produced in December. Laboratory cultures indicate that sexual reproduction is very rare in this species. L. longipes is rarely associated with sand.
Transplants and laboratory cultures indicate that production of mucilage ducts in the stipes of the two species is not affected by changes in environmental conditions. Comparison of the two species shows they differ in several other points besides mucilage ducts, including length of stipes, width of blades, winter loss of blades, morphology of gametophytes, and habitat. The evidence confirms that they should be retained as two separate species. iv
TABLE OF CONTENTS
PAGE
ABSTRACT ii
LIST OF TABLES vii
LIST OF FIGURES ix
I. INTRODUCTION 1
II. TERMINOLOGY 7
III. GENERAL MATERIALS AND METHODS 8
A. General Field Methods 8 B. Collection, Transport and Transplanting 8 C. Tagging and Growth Measurements 9 D. Laboratory culture 11
IV. DISTRIBUTION AND AUTEOOLOGY OF LAMINARIA SINCLAIRII 13
A. Geographical Distribution 13 B. Habitat and Autocology on Oregon beaches 15 1. Environment 15 2. Occurrence of L. sinclairii 31 3. Associated plant species 33 4. Seasonal cycles 37 5. Discussion 40
V. DISTRIBUTION AND AUTE OOLOGY OF LAMINARIA LONGIPES 42
A. Geographical Distribution 42 B. Habitat and Autecology at Aats Bay, Coronation Island 43 1. Environment 43 2. Occurrence of L. longipes and associated species 46 3. Growth and reproduction of L. longipes at Aats Bay 47 4. Discussion 48 V PAGE
VI. EXPERIMENTAL ECOLOGY 50
A. Field Work 50 1. Transplants 50 a. Alaska beaches 50 b. Oregon beaches 52 c. British Columbia beaches i. Whiffen Spit 54 ii. River Jordan 55 iii. Stanley park 56 d. Discussion 57 2. Rock clearing 59 a. Introduction 59 b. Methods 59 c. Results 59 d. Discussion 62 3. Transition zone experiments 63 a. Introduction 63 b. Procedures and results 63 c. Discussion 65
B. Laboratory Work 66 1. Sporophytes 66 a. Photoperiod 66 b. Salinity 67 c. Temperature 67 do Discussion 69 2. Gametophytes 69 a. introduction 69 b. Materials and Methods 70 c. Results 71 do Discussion 72 3. Haptera 73 a. Introduction 73 b. Materials and Methods 74 c. Results 78 i. Temperature 79 ii. Light intensity 80 iii. intial Size 80 vi
PAGE
VII. TAXONOMY OF L. SINCLAIRII AND L. LONGIPES 84
A. Introduction 84 B. L. sinclairii 85 1. Description 85 2. Distribution 86 3. Habitat 86 4. Comments 86 C. L. longipes 87 1. Description 88 2. Distribution 88 3. Habitat 88 4. Comments 89 D. Comparison of Species 89
VIII. GENERAL DISCUSSION AND CONCLUSIONS 93
IX. SUMMARY 101
X. BIBLIOGRAPHY 104
XI. TABLES I - XXIII 111
XII. APPENDIX I. Summary Description of Field Stations 151 vii
LIST OF TABLES
TABLE PAGE
I Seawater temperature and salinity over distribution ranges of L. sinclairii and L_. longipes. 112
II Distribution of Laminaria sinclairii. 114
III Distribution of Laminaria longipes. 116
IV Analysis of sand grain size on Oregon beaches. 118
V Temperature and precipitation at Alaska
stations. 119
VI Mean temperature on Urup Island. 120
VII Temperature and precipitation at British Columbia stations. 121 VIII Temperature and precipitation at Oregon stations. 123 IX Long term temperature and precipitation at two California stations. 124
X Measured values of seawater temperature and salinity on Oregon beaches. 125
XI Calculated mean monthly seawater temperatures on Oregon beaches. 126
XII Mean values of seawater temperature and salinity at Arch Cape for 1960-1963. 127
XIII Sand level at Indian Beach. 128
XXV Sand level at Short Sand Beach. 129
XV Sand heights of beach at Arch Cape, Oregon. 13 0
XVI Associated plant species. 131
XVII Seasonal distribution of plant species at Indian Beach. 136
XVIII Seasonal distribution of plant species at Short Sand Beach. 139 viii
TABLE PAGE
XIX Seasonal distribution of algal species at Aats Bay, Coronation Island. 142 XX Summary of seasonal cycles on Oregon beaches. 145
XXI Summaries of in situ growth measurements on Oregon beaches. 146
XXII Growth of multi-punched blade of L. sinclairii in situ at Indian Beach. 148 XXIII Dimensions of pressed specimens of L. longipes from 33 Alaskan sites. 149 ix
LIST OF FIGURES FIGURE PAGE 1 Distribution ranges of Laminaria sinclairii and L. longipes. 2 2 Mean seawater temperatures over ranges of L. longipes and L. sinclairii 4 3 Mean air temperatures over ranges of L. longipes and L. sinclairii. 5 4 Emergence and submergence of vertical distribution extremes of L. sinclairii during period from June 1966 to September 1967. 14
5 Sites in Oregon where L_. sinclairii was collected or studied. 18 6 Aerial and general views of Indian Beach and Short Sand Beach. 23 7 Indian Beach study area. 24
8 Short Sand Beach study area. 26
9 Variation in sand levels at Indian Beach in 1967, 27 10 Variation in sand level at study rocks at Indian Beach in 1967. 28
11 Variation in sand and water levels at Short Sand Beach in 1967. 30 12 L. sinclairii and associated species at Indian Beach. 34 13 Submergence in freshwater and association of L. sinclairii with other species at Short Sand Beach. 36
14 Aats Bay study area. 44 15 Transplant stations. 51
16 Cleared rock at Indian Beach. 61
17 Results of growth experiments on haptera of L. sinclairii. 77 18 Habit of L. sinclairii and L. longipes. 91 ACKNOWLEDGEMENTS
I wish to express my gratitude and appreciation to
Dr. R. F. Scagel for his direction, advice, and criticism throughout the course of this study. I also wish to expres my appreciate to Dr. G. C. Hughes, Dr. K. Cole, and
Dr. G.L. pickard for their advice throughout the study; to
Dr. G.H.N. Towers for advice on this manuscript; to
Mr. Stephen Borden for programming the computer analyses; to Mr. J. Thorpe for assistance with various kinds of equipment; to Mr. W.A. Markham for topographic surveys of the Oregon beaches; to Lt. Col. M.R. Simmonds and
Mrs. W.A. Markham for making precipitation data available; to the Oregon State park Department for allowing access to
Indian Beach after hours; to the Department of Botany and
Institute of Oceanography, University of British Columbia for facilities, equipment, and assistance; and finally to the many people who assisted me on various beaches, especially at night in the winter. I wish to acknowledge with thanks the support provided by fellowships from the
University of British Columbia and the National Research
Council of Canada. 1
I. INTRODUCTION
Laminaria Laraouroux, the most common kelp genus of north temperate waters, usually has a very simple morphology. It consists of a holdfast from which arises a single stipe with a flattened lamina, or blade. In three very unusual species the holdfast, composed of many branched haptera, is expanded into a rhizome-like organ from which arise many stipes, each bearing a single blade. One of these species, L. rodriguezii
Bornet, occurs at depths of 100 to 150 m in the Mediterranean and Adriatic Seas (Bornet, 1888). The other two are found in the intertidal and subtidal zones of the north pacific Ocean.
L_. longipes Bory is found from the Kurile Islands through the
Aleutian Islands and the Gulf of Alaska into Southeast Alaska.
L. sinclairii (Harvey) Farlow, Anderson and Eaton occurs from central British Columbia to Southern California (Fig. 1). Since no material of L_. rodriguezii was available, the two pacific species only are the subject of this study.
There are several reasons why a study of L. sinclairii and
L. longipes seemed warranted. The presence of multiple stipes suggested that the growth of these plants might differ somewhat from that of other kelps. It had been observed that the basal portions of L. sinclairii are often buried under sand; as few plants can withstand such burial, this suggested that the ecological adaptations of this plant might be somewhat unusual.
Finally, most taxonomic treatments have separated these two similar species by the presence of mucilage ducts in the stipes b
Figure 1 Distribution ranges of Laminaria Sinclair
and L. longipes.
3 of L. sinclairii and their absence in the stipes of L, longipes. Burrows (1964) has shown that presence or absence of mucilage ducts in the blade of L. saccharina (L.) Lamour. can be controlled by temperature. In her studies, plants grown at 10°C developed mucilage ducts, whereas those grown at 5°C did not. This is of interest because the average temperature of the seawater in Alaska and the Kurile Islands is markedly lower than that of the seawater in the area from British Columbia to California (Fig. 2). The difference in air temperature is even greater (Fig. 3). In view of this, it appeared that the presence or absence of mucilage ducts in the two species might be merely a response to the environment and might be altered by changing the environment. if this were the case, the presence or absence of mucilage ducts would not be a sufficient criterion for separating the two species. It seemed possible, therefore, that the two species might be one.
h.- sinclairii and L. longipes have been described by several authors, most recently by Druehl (1968) as part of a general taxonomic treatment of the genus. However, very little experimental work has been done with them. Setchell (1905) studied growth and regeneration in the blade and stipe of L. sinclairii and Myers (1925) cultured the gametophytes of L. sinclairii. Apparently no experimental work has ever been done on L. longipes.
This investigation consists of three parts. The first is an examination of the distribution of the two species throughout ure 2 Mean seawater temperatures over ranges of
L. longipes and L. sinclairii.
Port Hueneme is in Ventura County, California,
very near the southern limit of distribution of
L. sinclairii. Sitka is the nearest point to
Coronation Island for which there are long range
seawater temperature data. Note lack of overlap in
winter and annual means for the two ranges. 20
COLDEST WARMEST ANNUAL 18 MONTH MONTH MEAN
L. sin c. Port Huenerne
15 L. sine. Port L. long. Hueneme I 4 Sitkqi L. sin c. Short Port Huenemec 13 Sand Beach
I 2 Pescadero Pis Urup
10 [ope Island 9 Sitka ^^Hop 8 IS.
Hope Island1^ 7
5 Attu Urup L. long. Sitka 4
m Urup
Mean Seawater Temperatures over Ranges of L. Ion gipe s & L. sinclairii gure 3 Mean Air temperatures over ranges of L_. longipe
and L. sinclairii.
Oxnard is in Ventura County, California. Bull Harbour is on Hope Island, at the north end of Vancouver Island, British Columbia. Note lack of overlap in winter, summer, and annual means for the two ranges. °C 20 COLDEST WARMEST ANNUAL 19 MONTH MONTH MEAN L. sine. Oxnqrd
-6 u JUrup Is. Mean Air Temperatures over Ranges of L. longipes & L. sinclairii 6 their ranges in relation to oceanographic conditions, together with a more detailed presentation of autecological studies on four beaches. L. longipes was studied at Aats Bay, Coronation Island, Alaska during five visits in 1965 and 1966. L. sinclairii was studied during regular visits in 1965-1967 on three beaches in Northern Oregon: Indian Beach, Arch Cape, and Short Sand Beach.
In the second part, growth and reproduction of both species under experimental conditions are treated. Growth of sporophytes was followed in the field and in tanks of seawater in the laboratory under various conditions. Reciprocal transplant experiments were carried out in an attempt to assess the influence of environment on the morphology and anatomy of the plants. Gametophytes were cultured in a study of sexual reproduction. The growth of isolated pieces of haptera of L. sinclairii was also investigated in laboratory culture.
In the third part of the investigation, the validity of the two species as separate entities is examined in light of the information obtained from field observations and laboratory and field experiments. 7
II. TERMINOLOGY
Each of the two species studied has a holdfast of branched haptera which bears numerous stipes, each stipe in turn bearing a blade, or lamina. Because of the multiple stipes, confusion may arise as to what constitutes an individual plant. In this study, the terms "plant" and "clump" are used to refer to the holdfast with all its stipes and their blades. When an individual stipe or blade is referred to, it is designated "stipe" or "blade". 8
III. GENERAL MATERIALS AND METHODS
A. General Field Methods
The fluctuation in sand height at certain locations on two
Oregon beaches (Indian Beach and Short Sand Beach) relative to rocky reference points was measured each month by means of a pocket rule and an Abney level. A detailed map of each of the
two beaches (Fig. 7, 8) showing the location of all rocks with
L. sinclairii as well as its upper and lower limits, was prepared using a surveyor's transit and stadia rod. An IBM
1130 computer was used to calculate the total time in any given month that plants at various heights were out of water or under water in daylight and in darkness, based on published data for
tides and hours of daylight (Fig. 4)
B. Collection, Transport, and Transplanting
Plants were collected in the field by cutting the holdfasts loose from the rocks with a knife. In most instances, holdfasts bearing more than 40 stipes were separated into two or more parts. The plants were then transported in wet newspaper on ice in a freezer chest to holding tanks at the University of
British Columbia or directly, when possible, to another beach in transplant experiments. The plants were never out of water more than 36 hours and showed no apparent damage from this
treatment. In the case of transplants between Alaska and
Vancouver the plants were kept in large holding tanks in the shade on the deck of the ship en route. The water in these 9 tanks was changed every two to five days and never reached more than 10°C, due to the low ambient air temperature. In the transplant experiments, two methods were employed for attaching the plants in the field. The first, used in areas with little surf and many loose but stable boulders (Volga island, River Jordan, Whiffen Spit, Stanley park) (Fig. 6) consisted of placing two large rubber bands around a boulder and the holdfast. In the second method, employed in areas with greater surf and lacking loose boulders (Aats Bay, Indian Beach, Short Sand Beach), two or three spikes were driven into the bedrock and the holdfasts were attached to these with several rubber bands.
C. Tagging and Growth Measurements
In the field and in laboratory tanks, plants were identified by a piece of plastic flagging tape tied around a portion of the holdfast. Individual stipes were identified by a piece of flagging tape tied around the stipe. For plants growing in situ, each stipe was considered separately. For transplants in the field or in the laboratory, three stipes, designated A, B, and C were selected for growth study on each plant. A was the shortest stipe (and perhaps the youngest), C was the longest, and B was a stipe of intermediate length, usually closer to C than to A in length.
In order to determine the growth more accurately, a small hole was punched 10 cm above the base of each blade measured, using a cork borer and an apparatus like that used by Sundene 10
(1964), consisting of a centimeter rule with a notched piece of metal fastened to the lower end and a hole at the 10 cm mark. The notch was placed over the stipe at the transition between stipe and blade and the blade was punched through the hole in the rule. A new hole was punched in the same manner every time a measurement was made and three figures were then recorded: length of stipe, length of blade, and distance between the last two holes. In laboratory cultures and field transplants, the width of the blade at 5 cm above the base was also recorded. However, in plants growing in situ this measurement was not made after it was observed that a change was rarely recorded.
It was found that tying flagging tape around stipes in the field sometimes injured them, especially those in areas subjected to heavy surf. In later field measurements made in situ no identifying markers were used at all, a punched hole being the only indication that a blade and its stipe had been measured previously. Thus, stipes could not be identified as individuals but only as members of a particular group. How• ever, the distance between punched holes provided an absolute measure of growth of one part of the plant even if the stipe and its blade could not be identified as a particular individual in the previous month's data. At the beginning of each series of measurements 20 to 25 blades were punched within a small area (ca. 20x20 cm) on a rock and in succeeding months only those found to have holes were punched and 11 measured again. This continued until too few (usually- less than 10) previously punched blades were found, at which time new blades were punched. In winter, when no blades are present, a few stipes were tagged to indicate a definite area on the rock and then 2 0 to 25 stipes were measured in this small area.
D. Laboratory Culture
/<
For laboratory culture work,two New Brunswick psycrotherm Incubator Shakers and three walk-in controlled temperature rooms were employed. The Psycrotherms were kept at 8°C. Various light periods were employed using cool white fluorescent tubes with an intensity of 150 ft-c. Cultures were grown with and without shaking. The three controlled temperature rooms were maintained at 5°, 8°, and 10°C. In each of them two light intensities, approximately 20 ft-c (215.2 lux) and 150 ft-c
(1614.0 lux) were employed.
In each of the three controlled temperature rooms three 120 1 tanks of seawater were used for culturing sporophytes. Two of the tanks in each room contained seawater from Juan de Fuca Auto Court near Otter point on the west coast of Vancouver Island, designated West Coast water. The salinity was 30.9%o t 0„5%„ depending on the month of collection. The third tank in each room contained water from Stanley park,
Burrard Inlet (salinity = 27.9%o t 0.4%o) designated Stanley park water. The water was changed at approximately monthly intervals with freshly collected water. Aeration was 12 provided by an aquarium bubbler. The tanks were illuminated by cool white fluorescent tubes which gave an intensity at the surface of the water of 150-200 ft-c (1614-2152 lux). The depth of the water was 35-40 cm. photoperiods ranging from 8 to 16 hours were employed during the course of the study.
Gametophytes and portions of haptera were cultured in 250 ml glass culture dishes, standard 100 mm glass Petri dishes and 60 mm plastic Petri dishes. Four different seawater media were employed, all based on West Coast seawater: raw seawater
(SW), seawater filtered through cotton wadding (SWF), Erd-Schreiber medium (ES) (F0yn, 1934), and enriched Erd-Schreiber
(ES+) in which 1 ml of "ASP 2" medium (Provasoli, McLaughlin, and Droop, 1957) was added to each liter of Erd-Screiber. In addition, straight ASP 2 was employed in a few gametophyte cultures. One mg/1 of Ge02 was added to each medium to retard the growth of diatoms. The media were changed at intervals ranging from one day to one month, depending on the experiment. 13
IV. DISTRIBUTION AND AUTECOLOGY OF LAMINARIA SINCLAIRII
A. Geographic Distribution
L. sinclairii is found from Hope island, British Columbia
(50o56'N, 127°58'W) to Ventura County, California (34°19'N,
119°23.3'w) (Fig. 1). It has not been recorded between Hope
Island and Box Island, near Tofino on the southwest coast of
Vancouver Island.
The mean annual seawater salinity values show an increase
from north to south in the plant's range, with the mean at
Hope Island 31.7%0 and at port Hueneme, Ventura County, 33.6%0
(Table I). The mean annual seawater temperature also increases from north to south, from 8.6°C at Hope Island to
14.3°c at port Hueneme. Mean February temperatures range
from 7.2°C at Hope Island to 13.2°C at port Hueneme. Mean
August temperatures range from 10°C at Hope island to 16.8°C at Port Hueneme. Figure 2 illustrates the difference between
these conditions and those in the range of L. longipes. As many of the plants are out of water more than they are under water (Fig. 4), the air temperature and precipitation are
also important considerations. Tables VII, VIII and IX show
average temperatures and precipitation for selected sites in
British Columbia, Oregon, and California. The annual mean air
temperatures range from 8.7°C at Hope Island to 15.2°C in
Ventura County. Figure 3 illustrates the differences between
these conditions and those in the range of L. longipes. 14
Figure 4 Emergence and submergence of vertical distribution extremes of L. sinclairii during
period from June 1966 to September 1967.
Any vertical line drawn from top to bottom of each graph adds up to 100 percent of the hours in that month. "Dark" is percentage of total hours the point was in darkness. "Light" is percentage of total hours the point was in daylight. Line between "wet" and "dry" divides time under water from time out of water. 1966 1967 LIGHT
6T89 10 II 12 I 23456763 DARK I I I I 1 1 1 I I I I I 1 I
Highe st
• 3.0 ft L. Sinclqirii (•0.92m) DRY Short Sand WET Beach
Highest + 1.5 ft. DRY L. sinclairii (•0.46 m) Indian Beach WET
DRY
0.0 ft. Mean Tide (0.0 nt) WET
DRY Lowest
-1.5 ft. L. sinclairii (-0.46 m) WET Short Sand Beach
DRY
Low est -2.0 ft. L. sinclairii (-0.6lm) yVET
Indian Beach 15
Table II lists the sites at which L. sinclairii has been
collected or observed. The information on distribution was
obtained from collections available in the University of
California Herbarium, Berkeley; the phycological Herbarium of
the University of British Columbia; the author's own
observations and collections; and published records. L.
sinclairii has been found at relatively few places in British
Columbia and Washington, but is much more common in Oregon
and California, with very few exceptions, the plants are
found on beaches which are fully exposed to surf and have a marked seasonal fluctuation in sand level. They are generally
larger when found lower in the intertidal zone and in more
exposed sites. They are also larger near the southern limits
of distribution. It was also observed that plants from
California produce much more mucilage than those from Oregon,
although no quantitative measurements were made.
B. Habitat and Autecology on Oregon Beaches
1. Environment
The autecology of L. sinclairii was studied on three
beaches in Northern Oregon: Indian Beach, Short Sand Beach,
and Arch Cape (Fig. 5). Indian Beach is located in Ecola
State park and Short Sand Beach is in Oswald West State park.
Thus the forest and the land in general are relatively little
disturbed behind the beaches. To the north of Arch Cape there
is an almost continuous row of houses along the land bordering
the beach. South of the Cape there are very few houses in 16 the first kilometer, partly because of the very steep cliffs. The first two beaches were visited at various times over the period 1965-1967 and at least once a month from August 1966 through September 1967. At Arch Cape L. sinclairii plants are found in an area which is very difficult to reach except in the calmest weather and at the lowest tides of the summer. The study site here was visited only in June, July, August and September 1967, although it was observed from a distance on many other occasions.
On all three beaches there is a regular cycle of removal and deposition of sand (Fig.. 9,10,11,16) . The sand level on the beach rises from April or May until the first big storms (usually in September or October) remove most of the sand in a very short time, sometimes in as little as 24 hours. The sand forms offshore bars during the winter and then is transported to the beach again the following summer. The supply is continually augmented by sand which is brought to the sea by nearby streams and rivers.
Sand samples for each beach were analyzed for grain size, using a standard set of Endecott sieves (Table IV). The results are very similar for all beaches. The sand is very clean and well sorted. This, and the size distribution reflect the great wave action on the beaches. Over 97 percent of the sand on each beach is fine grained (0.25 -0.1 mm diam} to very fine grained (0.1 - 0.05 mm diam.). Spectroscopic analysis shows that the sand is primarily composed of quartz. Most of it is derived from metamorphic rocks with a smaller percentage of 17
granitic material. The main source of the sand is very likely the Columbia River (W.R. Danner, Dept. of Geology, U.B.C,
Pers. comm.).
Shepard (1963) noted that the supply of sand on the beach in the summer often varies directly with the amount of runoff the previous winter. This was observed to be true on the
Oregon beaches. The winter of 1966-1967 had much more precipitation than the previous winter (Table VIII) and the sand level on the beach was noticably higher in the summer of 1967 than in 1966.
All three beaches are in fully exposed locations. During winter storms the surf is very heavy and even on the calmest summer days there is some surf. The waves tend to approach from the northwest during calm weather but from the southwest during storms. Indian Beach and Short Sand Beach have larger waves than all other beaches in northern Oregon. Each of the two is a relatively short beach formed in an indentation of a prominent headland (Fig. 5). Each is exposed more to the south than to the north. Arch Cape is a headland with a long stretch of beach both to the north and the south, so that it is fully exposed to waves from all directions. The study site at
Arch Cape is a rock directly west of the Cape. The study sites on the other beaches are situated somewhat back of the points of the headlands. These outermost points were never reached.
At Indian Beach and Short Sand Beach, L. sinclairii occurs at each end of the beach. The areas of intensive study were the north end of Indian Beach and the south end of Short Sand 18
Figure 5 Sites in Oregon where L. sinclairii was collected or studied.
"A" and "B" near Arch Cape are sites where precipitation was measured. 124° 0 0' W 19
Beach. Although all the sites must be considered fully exposed,
the Arch cape site is slightly more exposed, followed by the
Indian Beach Site and then the Short Sand Beach site.
The tides in this area are of the mixed semidiurnal type.
The tidal pattern is the result of two component tides, the semidiurnal tide, with an interval between successive high waters of about 12% hours and the diurnal tide with an interval between successive high waters of about 25 hours. There are usually two high and two low waters each day, and successive highs or lows are generally of different heights. The lowest of the low waters occurs in the daytime in early morning in summer and at night in winter.
There are no published tidal data for any of the beaches studied. The nearest location for which there are data is
Astoria, Oregon. No information other than this was obtained for Arch Cape. From tidal data for the study period (June 1966 to September 1967) the following average heights were obtained: mean higher high water (MHHW), 3.91 ft. (1.19m); mean lower high water (MLHW), 2.37 ft (0.72m); mean higher low water
(MHLW), -1.81 ft. (-0.55m); mean lower low water (MLLW),
-4.47 ft (-1.36m). All heights are in relation to long term mean tide taken as 0.0 ft. Thus the average amplitude is about 8.3 ft. (2.5m). The extreme amplitude is approximately
ll ft. (3.4m). For Indian Beach and Short Sand Beach the actual tides appear to correspond to these calculated heights to within 0.5 ft. 20
This area is characterized by relatively heavy precipitation and moderate temperatures. The nearest place for which there are published records of precipitation and air temperature is Seaside, Oregon.(Pig. 5). Unofficial precipitation records have been kept by two individuals at Arch Cape (Fig. 5), one since 1957 (A), and one since 1965 (B). The rain gauges are situated about one kilometer apart, each atop a sea cliff about 20 m east of the beach, and each near a house. The more northern one (A) is located at a somewhat higher altitude and nearer to a house than the southern one (B). The records for Seaside and Arch Cape (Table VIII) show that the study period (1965-67) had below normal precipitation. The records show more precipitation at station B than at A and more at each than at Seaside. The maximum precipitation usually occurs in December or January and the minimum in July or August. The summer of 1967 was particularly dry. The mean total annual precipitation for Seaside is 79.7 inches.
Seaside is situated north of Tillamook Head, a large promontory, and Indian Beach is in the southern part of Tillamook Head. Thus, there may be significant differences in meteorological conditions at the two areas, particularly in precipitation. However, as no other information is available from a nearer location, the Seaside precipitation records are presented as an approximation of the conditions at Indian Beach.
The records for Station B at Arch Cape are presented as an approximation of precipitation at Arch cape and Short Sand Beach. In the absence of any other records, the Seaside air 21 temperatures are used for all three beaches. The annual mean air temperature is 11.0°c (Table VIII), ranging from a monthly mean of 6.3°C in January to 15.7°C in August. The year 1967 differs from the mean, especially in the higher summer temperatures. There are published records of seawater temperature and salinity for Arch cape (1960-1963) and the Seaside Aquarium (1949-1967). In addition, water samples were taken at Indian Beach and Short Sand Beach once a month from October 1966 through September 1967 (Table X). The samples were collected at low tide in the surf as near to the study rocks as possible and temperature readings were taken immediately. The salinity determinations were made with an inductive salinometer at the Institute of Oceanography, University of British Columbia.
In the Arch cape records there is considerable variation between the different years, probably partly because of the great variation in number of samples. The mean monthly seawater temperature ranged from 7.4°c in March 1962 to 15.8°C in July 1963. The mean monthly salinity ranged from 26.9%„ in
April 1962 to 32.9%c in August 1961. The conditions for 1967 are assumed to be within this range. Table XII gives the monthly means for the period 1960-1963.
The monthly readings obtained at Short Sand Beach and Indian Beach were compared with the records from Seaside. In almost all instances, when readings for specific dates were compared, the Seaside temperature was higher. The mean difference was 2.5C° at Indian Beach and 2.0C° at Short Sand 22
Beach. These differences were subtracted from the monthly means for Seaside for each month over the period studied to obtain an approximation of the temperature conditions at each beach (Table XI). Assuming this to be valid, at Indian Beach the mean monthly seawater temperature ranged from 7.6°C in March to 13.9°C in July, with a yearly mean of 10.5°C, whereas at Short Sand Beach the range was from 7.1°C in March to 13.4°C in July, with a yearly mean of 10.0°C. It is not known how typical this year may have been. Monthly means of seawater temperature for Seaside over the period of years (1949-1967) for which there are records are of no use because the annual means have risen over almost the entire period. This may be attributed in part to the steady widening of the Seaside beach due to increased sand deposition which has occurred since the construction of the Columbia River jetty approximately 25 km north of Seaside.
No interpolations are possible from a comparison of the salinity measurements at the two study beaches and Seaside. Local freshwater runoff obviously affects the readings significantly although irregularly, especially at Short Sand Beach (Table X).
The study site at Indian Beach is an area adjacent to two rocky points at the north end of the beach, one a headland,
the other a young stack (Fig 6, 7). There are numerous rocks which extend above, the level of the beach. Most of the rock is primarily olivine basalt. A freshwater stream flows across the beach just south of the study area. Its course is altered 23
Figure 6 Aerial and general views of Indian Beach
and Short Sand Beach.
a. Aerial view of north end of Indian Beach. Arrow shows approximate line of sight in figure b below. Rectangle indicates area shown in figure "7, b. Indian Beach at high tide in August 1967.
c. Aerial view of south end of Short Sand Beach. Arrow shows approximate line of sight in figure d below. Rectangle indicates area shown in figure 8.
d. Short Sand Beach shortly after low tide in March 1967.
24
Figure 7 Indian Beach study area.
L. sinclairii occurs on all rocks shown in figure
and at several points on the rock forming the
shore. Growth measurements were made on "A" and "B".
"C" was partly cleared (see Fig. 16). Sand height
was measured at point "F" and at "A". SCALE 25 with each tidal cycle, but it does not normally come in contact with any of the plants studied. The sand level was recorded at monthly intervals from August 1966 through September 1967 at four points on the beach (Table XIII). The total fluctuation in sand level is at least 1.2 m and probably more on some areas which were not measured. On most of Indian Beach, the sand level is lowered in winter but a sandy beach still remains. On the area studied however, all the sand is removed in winter, exposing the bedrock (Fig. 9a, 10a).
Most of the area studied at Indian Beach is situated so that it is shielded from direct insolation during most of the time the rocks are exposed at low tide. It was observed that the rocks and the plants growing on them always retained enough water so as to appear wet even if they had been exposed by the receding tide for as long as 3 hours. The only exception to this occurred during an extremely dry, hot period in August 1966 when the relative humidity of the air fell below 20% and the air temperature reached 30°C. The rocks appeared completely dry and some of the plants were curling, almost as if burned.
The study site at Short Sand Beach is a series of rocky outcrops along the narrow margin between the southern end of the sandy beach and the steep cliffs which form the southern boundary of the cove in which the beach is located (Fig. 6, 8). The rocks are primarily sandstone, and much softer than those at Indian Beach. A freshwater stream flows across the beach parallel to and near the outcrops. The sand level was recorded lire 8 Short Sand Beach study area.
L. sinclairii occurs on all rocks shown and at several points on the rock forming the shore. Particular measurements or observations were made on "A" and "B".
27
Figure 9 Variation in sand levels at Indian Beach in 1967.
a. April 1967
b. May 1967
c. June 1967
d. July 1967
In c and d, the left bucket (arrow) is on rock
A, the right bucket (arrow) on rock C. The large
rock is the stack indicated in figure 7.
28
Figure 10 variation in sand level at study rocks at
Indian Beach in 1967.
a. April 1967. Note that surf is around rocks, even
at low tide.
b. May 1967. Sand has advanced but is not yet
around most of the rocks.
c. June 1967
d. July 1967
e. August 1967. Some of the blades of L_. sinclairii
have been scoured off rock A.
f. September 1967.
Arrow in all figures indicates rock B. In c, d,
e, f, bucket in foreground is on rock A, bucket in
background is on rock C (= cleared rock; see Fig. 16).
29 at monthly intervals from August 1966 through September 1967 at two points on the beach (Table XIV). The greatest recorded fluctuations of the sand level at the study sites is 70 cm. The freshwater stream which flows so near the rocks interacts with the sand to produce a somewhat different situation from that on Indian Beach. The course of the stream is shifted slightly during each tidal cycle. However, most of the time the stream flows next to the rocks, at least in the upper part of the beach, separating the rocks from the rest of the beach. As the sand level rises in summer, the level of the fresh water rises next to the rocks, so that by late summer as much as 30 cm of some rocks is submerged in fresh water at low tide (Fig. 11). This includes much of the portion of the rocks which is inhabited by L. sinclairii. The presence of the stream, and its constant erosion of a channel, in turn prevents the rocks from being buried so deeply under sand. During the latter part of the summer of 1967 the level of the sand on all the beach north of the stream was at least 1 m above the level of the tops of the rocks studied (Fig. llf). The rocks had about 35-45 cm exposed above the sand in the bottom of the stream channel and about 5-10 cm of this exposed above the freshwater, depending on the height of the rock. In winter, when sand is completely absent next to the rocks, there is usually a 10-20 cm strip of rock
at the bottom, where the stream flows, which is completely bare of plants. The rock above, which supports L. sinclairii and various other algae, is not touched by the stream water. 30
Figure 11 Variation in sand and water levels at Short
Sand Beachin 1967.
a. Level of sand and freshwater stream in June 1967. b. Level of sand and freshwater stream in July 1967. c. Level of sand and freshwater stream in August 1967. Note that sand is higher than study rocks.
In a, b, and c, bucket in foreground is on rock B, bucket in background is on rock A.
d. Study area from above in April 1967. Rock A is out of picture. Rocks in background are completely buried later in summer.
e. Study area from above in July 1967. f. Study area from above in August 1967. Sand in background is higher than study rocks and freshwater stream has risen also.
In e and f, left bucket is on rock A, right bucket is on rock B.
31
The study site at Arch Cape is a large basaltic outcrop directly west of a large stack west of the Cape. The rock is about 7 m from east to west, 2 m from north to south and about 1 m in height from the lowest winter sand level. It is separated from the stack by a stretch of beach about 20 m wide. Somewhat north and west of it is another, larger rock. With the exception of this larger rock, the study rock receives a greater force of surf than any other point for about 5 km along the beach. A freshwater stream flows across the beach about 30 m north of the rock but does not usually come into contact with it.
No measurements of sand fluctuation have been made near the study site at Arch Cape. Measurements of sand height made at a point approximately 1 km farther north on the same beach show a maximum fluctuation of only a little over 1 m (Table XV). It is probably greater near the rock.
2. Occurrence of L. sinclairii L. sinclairii occurs at Indian Beach on many rocks at the north end of the beach (Fig. 7). It is restricted to the region between the +1.5 and the -2.0 ft. (+0.46 and -0.61m) tide levels on all of the rocks. The size of the plants tends to be larger in the lower parts of the distribution.
At Short Sand Beach, L. sinclairii occurs on nearly all rocks which border the sandy beach at the south end (Fig. 8). It occurs only between the +3.0 and -1.5 ft. (0.92 and -0.46m) tide levels. At the upper limits of its distribution it is 32 very small. In June 1967, during a -6.1 ft (-1.86m) tide when the sand was still very low, the lower limits of L. sinclairii were determined on rocks far out toward the point. Below the lowest L. sinclairii, a patch of about 15 cm of completely bare rock was evident.
At Arch Cape, the distribution of L. sinclairii was not mapped. It occurs on several rocks at the outer part of the Cape. It covers the entire surface of the rock on which it was measured. The vertical distribution was not measured. Based on observations during various tides, all the plants appear to be confined to the area between MLLW (-4.5 ft.) (-1.37 m) and LLLW (ca. -7 ft.) (-2.1 m). This is lower than on the other beaches. However, the distribution does not extend into the subtidal zone.
As is shown in Figure 4, there is a considerable difference in total time of submergence between the upper and lower limits of plant distribution. Plants growing above the 0.0 ft. tide level are actually out of water for a greater percentage of the total time then they are under water. As the total time out of water varies and with it the total time which the plant is exposed to sunlight while out of water, there are many differences in temperature, light quality and intensity, and potential for desiccation at different levels. Thus it is to be expected that growth and size vary at the different levels. The sites where growth measurements were made are all at different heights. The measured plants on rock A at Indian Beach are at the 0.0 ft. tide level. Those on rock B are at 33
-0.5 ft (-0.15m). The total time which the plants on rock A are out of water each month is about 7 percent greater than that for those on rock B. The plants on rock B have a greater size and greater average monthly growth than those on rock A (Table XXI). The measured plants at Short Sand Beach are at about the +2.0 ft. (0.61m) tide level (ca. MLHW). These plants are somewhat smaller than those at Indian Beach.
3. Associated Plant Species Lists of all the plant species collected on the three Oregon beaches together with the heights and months of collection are presented in Tables XVI/ XVII/ XVIII. Only two collections were made at Arch Cape. There are very few species present there. On most of the rocks where L. sinclairii grows, no other species are found with the exception of a few plants of L. setchellii on the outer and most exposed parts of the rocks. At Indian Beach L. sinclairii is the most abundant plant in terms of cover on all the rocks on which it grows (Fig. 11, 12). However, several other species are commonly found with it. Phaeostrophion irregulare occurs at the same height and slightly higher than L. sinclairii. Several species of Bossiella are often found growing with L. sinclairii. Dilsea californica is common, usually at a slightly higher level on the same rocks, various crustos'e coralline algae are found on the same rocks above and below L. sinclairii. A few plants of L. setchellii are often associated with L. sinclairii, although L. setchellii tends to occur lower in 34
Figure 12 L. sinclairii and associated species at Indian
Beach.
a. Association of L. sinclairii with other species at 1.0 ft. below mean tide level. L. setchellii and Phyllospadix scouleri occur with L. sinclairii. Hedophyllum sessile is higher on rock. June 1967.
b. Burial under sand of L. sinclairii in August 1967. Some blades have been eroded away.
c. Plants on rock B in June 1967. Multipunched blade is visible. d. Plants on rock A in June 1967. Some punched blades are visible. Note less complete dominance and less luxuriant growth of L. sinclairii as compared with that on rock B which is lower.
35 the intertidal zone and in slightly more exposed locations than L. sinclairii. Phyllospadix scouleri is common with L. setchellii and the lower parts of the L_. sinclairii population (Fig. 12a). Gymnogongrus linearis and Codium setchellii occur from the lower limits of the L_. sinclairii population down to about 50 cm below these limits. Both are usually buried under sand for most of the summer. They are apparently well adapted to this, as they appear healthy when the sand again recedes. L. sinclairii shows more complete dominance on lower rocks (Fig. 13).
At Short Sand Beach, L. sinclairii is dominant on only a few of the lowest, most exposed rocks on which it occurs. On most of the rocks where it occurs there are several other algae in equal or greater abundance (Fig. 13 d,c). Dilsea californica and Farlowia mollis are very abundant on some of the rocks, usually slightly higher than L. sinclairii. Bossiella plumosa, Microcladia borealis, ptilota asplenoides, P_. filicina, P_. pectinata, Iridaea sp. and Prionitis lyallii are very common at the same height as, and slightly higher than L. sinclairii. On many of these rocks, the dominant cover is composed of Microcladia borealis and ptilota spp. Crustose coralline algae occur on the same rocks,above, below, and with L. sinclairii. Gymnogongrus linearis occurs at the same height and somewhat lower than L. sinclairii. Below the level of most of the L_. sinclairii, phyllospadix scouleri is abundant and several plants of L. setchellii are found. Gymnogongrus linearis and Phyllospadix scouleri are often nearly 36
Figure 13 Submergence in freshwater and association of
L. sinclairii with other species at Short Sand
Beach.
a. Some of L. sinclairii for which growth was measured, hanging into freshwater stream, June 1967. Two punched holes are visible.
b. Late summer submergence, August 1967. Bucket is on rock A. Plants on rock A are partly buried under sand as well as being submerged in freshwater. c. Burial under sand in freshwater stream of L. sinclairii, July 1967.
d. Some of L. sinclairii for which growth was measured, showing small size of clump and relative lack of dominance compared with other species, May 1967.
e. Small patches of L. sinclairii on rock with cover of ptilota spp and Microcladia borealis. June 1967.
37 completely buried under sand. Most of the other species mentioned are partly buried or at least coated with sand for much of the summer.
4. Seasonal Cycles
In the course of a normal year, seasonal cycles of four different phenomena occur in L. sinclairii. The plants grow; they bear ripe sori; they lose their blades; and they are buried under sand. The observations on which the following account is based are summarized in Table XX.
In early January, the plants are normally completely without blades. The rocks on which the plants are attached are completely uncovered and sand may be absent from the entire area. Growth of the plants is very slow. Later in January the ends of the stipes split and from the medullary regions new blades begin to develop. The blades continue growing and the rate of growth accelerates. The stipes grow, but much more slowly than the blades. Table XXI summarizes the growth measurements of in_ situ plants on the three Oregon beaches.
Early in February when the plants are only 2-3 cm in length, fertile sori begin to develop at the blade tips. Ripe sori are produced in March and sometimes as late as April. At the beginning of this reproductive period there is considerable difference in the time of sorus development and size of sori developed. Later in the period, the sorus production becomes more synchronized. In April 1967, at the beginning of a low tide series, nearly all the blades in a 38 population on one rock at Indian Beach had ripe sori on the terminal 2 cm of blade. These remained 4 days until the high tide before the last low tide which would uncover the plants for more than an hour in this tide series. During the last low tide it was observed that all the sori had been dropped, leaving a bite-shaped hole at the end of each blade.
Late in March, and in April and May the maximum initiation of new stipes and blades occurs from the holdfasts. New stipes can be distinguished from the older ones by color. During the first year the stipes are the same light brown color as the blades. By the time the stipes are one year old and have lost their blades once, they are very dark, almost black, and very unlike the blades in color. The growth of the other stipes and blades is greatest in May and June.
In April the sand on the beach begins to build up, but is not yet around the rocks on which L. sinclairii grows. Sometime in May or June, the low places are usually covered with sand, leaving the rocks bearing L. sinclairii still sticking out of the sand (Fig. 10c). In years when the sand level is high, the rocks as well as the holdfasts and stipes of most of the L. sinclairii plants on them are completely buried by mid-July, leaving only the blades of L_. sinclairii exposed (Fig. lOd). Such was the case at Indian Beach and Arch Cape in 1967. It would have occurred at Short Sand Beach also, had the stream not kept the sand away from most of the rocks. In other years, as in 1966, the sand did not rise as high and by September only the holdfasts of the plants 39 were buried. In September or October, the first heavy storms remove most of the sand, exposing all the rocks and plants again until the following summer.
As the plants become buried, growth of the blades decreases (Table XXI). However, it was noted that the decrease in growth is much less marked in plants which are growing lower in the intertidal zone and in more exposed areas. In these areas the period of maximum growth may be one or two months later than in other areas. This can be seen in comparing results at Indian Beach with those at Short Sand Beach, which is slightly less exposed, and at Arch Cape, which is more exposed. It is also evident in comparing the two Indian Beach sites, which differ in exposure. The growth at Short Sand Beach may also have been inhibited by the increasing contact with the freshwater stream which rose and impinged on the plants more as the sand level under it rose. As a consequence of the greater growth, the plants generally have longer stipes and blades in the more exposed area.
As the plants become buried, the scouring action of the sand increases. In some areas, many or all of the blades may be lost, possibly due to the scouring. However, this varies with exposure. In the summer of 1967, many blades were lost at Short Sand Beach, fewer at Indian Beach, and none at Arch Cape. The burial was more.complete at Arch cape than at any of the other study areas. The strong freshwater influence at Short Sand Beach may also have contributed to the blade loss. 40
The plants grow very little from September through December. In October the first ripe sori of the old blades are developed. In November large areas of the blades are covered with oblong patches of sori. Experiments show that these sori produce zoospores which develop into gametophytes, at least under laboratory conditions. However, these gametophytes do not usually produce sporophytes.
Mechanical erosion of the blade tips occurs to some extent at all times. Under normal conditions growth proceeds faster than erosion, and the blade lengthens during the year. Beginning in November when growth has practically ceased, endogenous disintegration of the blade tips begins to occur in addition to erosion. These processes continue until by mid-December most of the blades are completely gone, leaving only the bare stipes.
5. Discussion
L. sinclairii was observed at several beaches in Washington, Oregon, and California (Table II). At all of these beaches the general conditions are very similar to those on the study beaches. The beaches are fully exposed to surf and there is evidence of a marked seasonal fluctuation in sand level which causes the plants to be buried in summer. The associated plant species are also similar to those found on the three study beaches (Table XVI). Thus it appears that the beaches selected for study are typical.
The plants of L. sinclairii bear ripe sori during two periods, once just after the new blades begin to develop and once just 41 before the old blades are lost. It appears evident that sorus initiation is not at all dependent upon the age of the blade tissue. The factors which induce this initiation have not been determined.
Both periods of sorus production occur during the time when the sand is at its lowest. This leaves the maximum amount of bare rock available for the zoospores to attach and produce gametophytes which then could produce sporophytes. However, no evidence has been found that this actually occurs (See also pp 59-63 and 69-73).
L. sinclairii grows better the lower it is in the intertidal zone but Kas not^found in the subtidal zone. It appears that it may require a slight amount of desiccation but cannot tolerate very much desiccation. This hypothesis has not been tested, although a lack of desiccation may partly explain the poor growth in laboratory tanks (See pp 66-69) If the plants are growing in the optimum position with regard to surf and desiccation, burial by sand appears to cause no damage. This indicates that the plant is well adapted to withstand burial but is more sensitive to variations in desiccation and wave exposure. The results, such as loss of blades and cessation of growth in less favorable sites, may not be directly due to sand burial at all. 42
V. DISTRIBUTION AND AUTECOLOGY OF LAMINARIA LONGIPES
A- Geographical Distribution
In the Western pacific, the southern limit of distribution of L. longipes is Urup Island (46°00'N, 150°00'E) in the
Kurile Islands (Table III, Fig. 1). North of this, the plant is found on South Sakhalin, on various other islands in the Kurile Islands, at several points along the east coast of
Kamchatka, and on Bering Island. In the Eastern pacific,
L. longipes is found on St. Paul Island in the Bering Sea, and from Attu Island in the Aleutian Islands through the Gulf of
Alaska to Coronation Island (55°49.6'N, 134°17'W) in Southeast
Alaska, A subtidal population has been reported by Druehl
(1968) at Salmon Bank (48°26'N, 123°01'W) San Juan Island,
Washington. The plants from Salmon Bank have blades up to 2 0 cm broad, but otherwise fit the description of L_. longipes.
Until more extensive subtidal collections are made in the
Northeast pacific, it is not possible to assess the significance of this collection as an extension of the range.
The area in which L. longipes grows is characterized by very low winter seawater temperatures (Table I, Fig. 2) ranging from means of 0.5°C at Urup Island to 4.4°C at Sitka.
Summer temperatures range from a high monthly mean of 9.3°C at pyramid Cove to 14.1°c at Sitka. The yearly means range o o from 5.1 C at Urup Island and Attu Island to 8.5 C at Sitka.
The air temperatures are also very low (Table V, VI, Fig. 3) 43 particularly at Urup Island. The salinity within the plant's
range varies from a yearly mean of 27.7%0 at Sitka to 32.1%0 at Pyramid Cove.
Table III lists the sites at which L.. longipes has been collected or observed. All information on distribution in Russian waters has been obtained from published records. Most of the records of distribution in American waters have been obtained from collections available in the Phycological Herbarium, University of British Columbia. At most of the sites observed by the author, L_. longipes grows on rocky reefs in moderately exposed to moderately sheltered areas. No particular variation in morphology was noted in plants from various parts of the range of distribution.
B. Habitat and Autecology at Aats Bay, Coronation Island,
Alaska
1. Environment The study site at Aats Bay is a rocky reef adjacent to a beach composed of gravel and coarse sand (Fig. 14). The reef is composed of argillite rock and is cut by several deep surge channels. There is no evidence to indicate that the reef is ever buried under sand or that the sand level changes. The beach is in a location which is moderately sheltered to moderately exposed. Although there is frequently surf in the winter, in summer the water is often completely calm. The tides in this areas, as in Oregon, are of the mixed semidiurnal type. The lowest of the low waters occurs in the daytime in 44
Figure 14 Location of Aats Bay study area.
Helm Point is the southern limit of distribution of L. longipes.
45 summer and at night in winter. The maximum tidal amplitude at Coronation Island is 16.8 ft (5.12m) the diurnal amplitude is 10.7 ft. (3.26m), and the mean tide level is 8.7 ft. (2.65m) above MLLW (Anon., 1968b).
This area is characterized by relatively heavy- precipitation and low temperatures. There are no published meteorological records for Coronation Island. However, Cape Decision (56°00'N, 134°08"W) is only 10 km from Coronation island (Fig. 15) and it is assumed that the meteorological conditions are very similar at the two places. The records for cape Decision (Table V) show that the annual mean air temperature is 6.3°C, ranging from a monthly mean of 0.9°c in January to 11.7°C in August. The mean total annual precipitation is 76.12 inches, with the greatest amount occurring in October and the least in June. A comparison of these records with those for Sitka (Table V) shows that the average monthly precipitation is greater at Sitka than at Cape Decision for every month of the year. A comparison of the temperature records shows a greater annual temperature range at Sitka than at Cape Decision. This might be expected, since Sitka is on a large island and is largely surrounded by land, whereas cape Decision is more subject to the moderating influence of surrounding water masses. Maps of the area indicate the harbor at Sitka receives more freshwater runoff than does the sea around coronation Island and cape Decision, although exact figures are lacking. This, together with the rainfall, is important in considerations of salinity in the two areas. 46
There are no published records of salinity or water temperature at Coronation Island. In this study these factors were measured only once, in December 1966. At this time the temperature was 4.5°C, 1C° below the reading obtained at
Volga Island, Sitka, 24 hours later. The salinity was 30.78%o,
which was 0.5%o below that recorded at Volga island. Long term records of the seawater temperature and salinity have been published for Sitka (Anon 1967d; see also Table XXIII). If the difference recorded in December is valid for the whole year, the seawater temperature range at Cororation Island should be from approximately 3.4°C in February to 13.1°c in August, with a yearly mean of 7.5°C. These may be fairly close to the correct values, but adequate information is lacking. Because of the greater precipitation and runoff at Sitka, it seems likely that the average salinity in Sitka Harbor is lower than that at Corontation Island. Inasmuch as the one set of recorded observations show the inverse relationship, the average monthly salinities cannot be estimated from the available data.
2. Occurrence of L. longipes and associated plant species L. longipes occurs on the rocky reef in the lower intertidal and upper subtidal zones. Most of the plants are uncovered by a -10.2 ft. (-3.11m) tide. On the sections of the reef where it is found, it is usually the dominant plant in terms of total cover, although in a few places Alaria marginata is equally abundant. Slightly higher on the same 47 reefs, in the mid-intertidal position, Laminaria groenlandica, Hedophyllum sessile and Alaria tenuifolia are abundant. A complete list of the species found on this beach is presented in Table XIX.
3. Growth and reproduction of L. longipes at Aats Bay Growth of in situ plants was followed for only one month, July 1966. Holes were punched in the blades 10 cm above the base in the usual manner. Based on the holes the average growth of the blades for July was 8 cm. The stipes grew also, but much more slowly. It is assumed that maximum growth occurs in the summer months but direct measurements are lacking.
The blade reaches its maximum length sometime during the summer and then begins to decrease in length due to erosion and disintegration of the distal end. This process continues until the blade reaches a minimum length sometime in the winter. Laboratory cultures indicate that endogenously controlled disintegration is more important than physical abrasion in this process. In laboratory tanks, the same annual cycle occurred in the absence of any water motion.
Druehl (1968) states that L. longipes is "perennial from the stipe and holdfast", indicating that the blade is lost completely down to the stipe every year, as is the case with L_. sinclairii. The present study indicates that although the blades are much reduced in winter, they are never completely lost. There are three lines of evidence for this: (1) In December, when L. sinclairii plants are usually totally devoid 48
of blades, plants of L. longipes at Aats Bay all possessed short blades in 1965 and 1966. (2) in laboratory tanks and in transplants to Oregon L. longipes never lost its blades completely. (3) In herbarium specimens of L_. longipes plants from a large number of locations (Table XXIII) there is very often a remnant of the previous year's blade at the distal end of the current year's blade.
Plants of L. longipes bear ripe sori in December. No information is available for other winter months. Laboratory cultures show that these sori liberate spores which develop into gametophytes. Little evidence was found either in the field or in the laboratory to indicate that these gametophytes normally produce sporophytes (See also pp 69-73).
4. Discussion
Aats Bay was visited only five times in the two year study period: twice in the winter and three times in the summer
(Appendix I). Consequently the observations presented here are somewhat limited. It would be very useful to have more detailed information on seasonal growth of the plants, as well as seasonal changes in seawater temperature and salinity. Also, it would be valuable to know if there is a change in sand level.
L. longipes has been collected at sites ranging in exposure from "moderately sheltered" to "fully exposed" (Table XXIII).
Exposure is understood to mean the extent to which a beach is subject to surf and other water motion. This terminology is very difficult to put in quantitative terms. Each collector has 49 described his collecting sites in terms of his own concept of relative exposure. Most collections in Alaska have been made in summer. In some places in Alaska, particularly the Aleutian Islands, there is apparently a very great difference between summer and winter conditions. If a site was visited only once, it is possible that the conditions were very atypical on that particular day, which could lead to an incorrect evaluation of exposure. For many stations in Alaska not visited by the author, it is impossible to state whether the exposure, as it has been recorded, is representative.
Most of the "fully exposed" sites in Alaska which the author has visited are not exposed to as severe surf, or on such a regular basis, as any of the sites on the coast of Washington, Oregon, or California. There are undoubtedly places in Alaska which are as exposed as most places on the coast from Washington to California. However, most of these have not been visited by collectors, due to the difficulty in landing on such beaches from a ship. Future collections, perhaps made with the aid of a helicopter in otherwise inaccessible places, may alter the present picture of habitat distribution of some species. On the basis of present information, L. longipes is normally found on beaches which are less exposed than those which L. sinclairii inhabits. 50
VI. EXPERIMENTAL ECOLOGY
A. Field Work
1. Transplants
In order to investigate the growth and survival of L. sinclairii and L. longipes in other natural environments, and to determine whether mucilage duct development could be induced or suppressed by different temperatures, several field transplants were carried out. These fall into three main categories: L_. sinclairii from Oregon was transplanted to two beaches in Alaska; L_. longipes was transplanted from Alaska to two beaches in Oregon; and both species were transplanted to three beaches in British Columbia. Figure 15 shows the location of all sites utilized in transplant studies.
a. Alaska beaches
Two beaches in Alaska were utilized for transplant experiments. Aats Bay was chosen because it was the site of in situ studies on L. longipes. Volga Island, Sitka (See Appendix I) was chosen because it was assumed to have a similar water temperature to Aats Bay and is easily reached from Sitka. At Volga island, plants were attached to loose rocks which were placed in tide pools where other loose rocks were present. At Aats Bay the plants were attached to the reef on bare spots among the plants of L_. longipes.
In the first transplant attempt, five plants of L. sinclairii were placed at Volga Island in June 1965. All of these were subsequently lost. In the second attempt, five 51
Figure 15 Transplant stations plants were placed at Volga island and four plants at Aats Bay in December 1965. All of those placed at Aats Bay were lost. Of those placed at Volga Island, two were lost, one remained but died, and one remained alive to July 1966. It showed some growth in length as well as production of new stipes. Sections of these new stipes did not reveal any mucilage ducts. In the third attempt, in July 1966, four plants were placed at Aats Bay and four at Volga island. Aats Bay was revisited one month later. At this time three of the plants remained. Two showed no change. One had grown 8 cm, the same growth shown by L. longipes plants marked and measured in situ. In December 1966, two plants remained at Coronation Island, but in very poor condition. Two transplants were found at Volga island at this time. Both had attached to the rocks on which they were planted and appeared healthy.
b. Oregon beaches
Indian Beach and Short Sand Beach in Oregon were utilized for transplant studies. On both beaches the plants were attached to bedrock in areas where L. sinclairii was already present. In the first attempt, five plants of L. longipes were planted at Indian Beach in August 1965. All of these were lost during the following month. In the second
attempt, five more were planted at Indian Beach in December 1965. Again, all were lost in the next month. In the third attempt, two plants were planted at Indian Beach in May 1966. One of these survived without growing to June 1966 and then died 53
In August 1966, two plants of L. longipes were planted at Indian Beach, and two at Short Sand Beach. In addition, one L. sinclairii from Indian Beach was planted at Short Sand Beach, and one from Short sand Beach was planted at Indian Beach. One of the L. longipes at Indian Beach disappeared the following month. The other showed slight growth during September and then was not found again. The L. sinclairii from Short Sand Beach grew during September and October, lost its blades in December, and disappeared in January. At Short Sand Beach one L. longipes and the L. sinclairii remained through November and were torn loose in December. The other L. longipes was lost in November. When examined in September, all three had grown, both of the L. longipes slightly more than the L_. sinclairii. Thereafter no evidence of growth was found and the blades became progressively shorter until the plants were torn loose.
In December 1966, the final transplants were started. Four plants of L. longipes were placed at Indian Beach and
four at Short Sand Beach. Those at Indian Beach were lost almost immediately. At Short Sand Beach, three remained through March and then disappeared. Of these, one showed some growth during January and February, then deteriorated until it was lost. The others remained unchanged through January, then deteriorated until they washed away. in January, when the L. sinclairii plants had no blades or were just beginning to develop new ones, the L. longipes plants retained the long blades of the previous year. 54
c. British Columbia beaches Three beaches in British Columbia were utilized: Whiffen Spit and River Jordan on the west coast of Vancouver Island, and Stanley Park, Vancouver (see Appendix I). The first two were chosen because L_. sinclairii had been found there occasionally, although they are in a much more sheltered area than the normal habitat of L. sinclairii. The third site, Stanley park, is completely sheltered and has a lower salinity than the open coast habitats. It was chosen in an attempt to determine why h. sinclairii and :L. longipes are not found in such habitats. At the British Columbia stations, especially at Stanley park, the range of temperature for both air and seawater is greater than in Oregon or Alaska. In most experiments, plants were placed at all three beaches at nearly the same time. To avoid confusion, the results for each beach are considered separately.
i. Whiffen Spit - At Whiffen Spit, all transplants were attached to loose boulders which were then placed at the base of the landward side of a large rock. The rock provided an easy means of finding the transplants on subsequent visits. In the first attempt, four plants of L. longipes were planted at Whiffen Spit in July 1965. All of these were lost before further observations were made. In November, two L. sinclairii were planted and these disappeared also. In January 1966, four L_. longipes were planted. All remained in place and showed a large blade growth and production of new stipes through March. When the site was last visited in August 1966, 55 two still remained. They appeared healthy and had grown in the previous month. One had attached to the rock on which it was planted. This latter was collected and sectioned. No evidence of mucilage ducts was found. One additional plant of L. longipes was planted in March 1966. It grew through June and then was lost. In March 1966, four L. sinclairii were planted. One disappeared very soon, one died in the first month, and two showed good growth through May. They survived through June and then were lost.
The growth of both the L. sinclairii transplants and the L. longipes transplants was very great from March through May relative to that at other seasons and on other beaches. However, the L. longipes transplants showed a slightly greater growth. Both species produced wider blades than in their normal habitat. There was no twisting and the blades appeared to have developed normally. All plants which survived at Whiffen Spit were shaded and almost hidden by large plants of Hedophyllum hanging from the vertical face of the large rock above.
ii. River Jordan - At River Jordan the transplants were attached to loose boulders from various parts of the beach which were then re-placed. Most of these boulders were hidden under large plants of Egregia menziesii and Phyllospadix scouleri which were attached nearby and lay across large adjacent areas. In November 1965, two L. sinclairii were planted. These were not found again until March 1966. At this time the blades were shorter or absent and there was no evidence that growth had occurred. When examined again in May 56 there were several new stipes and considerable blade growth had occurred. One of the plants had attached to the rock. In March 1966, two other _L. sinclairii were planted. When examined in May these showed about the same blade growth as transplants set out in November and several new stipes were present. In June one of the plants had attached to the rock, the other was dead. In January 1966, two plants of L_. longipes were transplanted to River Jordan. These remained and grew well through June 1966.
As at Whiffen Spit, both species produced wider blades after being transplanted but otherwise appeared healthy and "normal". All plants at River Jordan which grew were protected from direct insolation by large fronds of Egregia menziesii lying on top of them.
iii Stanley park - The transplants at Stanley park were attached to loose boulders which were then placed near one large rock for ease in finding on subsequent visits. L. saccharina occurs above and below the area used for transplanting. In the first transplant experiment, two L. sinclairii were planted in November 1965. One of these disappeared. The other survived to June 1966. It continued growing the whole time according to measurements between punched holes. It attached to the rock on which it was planted.
The blades were never completely lost. They decreased in length to March and then began increasing in length. The second experiment with L_. sinclairii was started in March 1966 when six plants were transplanted. Of these, one died, one grew 57 through May, and four grew through June. Of the latter group, one had attached to the rock by May. Five plants of L. longipes were transplanted to Stanley Park in August 1965. All of these died very soon. Two more L. longipes were transplanted in December 1965. They grew fairly well through May and then died in June.
In winter the L. longipes appeared to be healthier than the L. sinclairii, but as summer approched, the L_. sinclairii appeared healthier. Both species produced blades considerably broader than in their normal habitats. They became very twisted and overgrown with bryozoans, diatoms, and various other epiphytic algae.
d. Discussion It was observed that no mucilage ducts were present in new stipes produced by L. sinclairii plants at Volga Island. This would seem at first to confirm the original idea that mucilage duct formation requires a temperature higher than that normally encountered in Southeast Alaska. However, the new stipes which were sectioned were very small. Subsequently, equally small stipes from plants taken directly from the field in Oregon were sectioned. These did not show any mucilage ducts either. It is concluded that the stipes of L. sinclairii must attain a certain minimum size before mucilage ducts are produced and this size was not attained by any of the new stipes produced in Alaska.
The Alaskan sites, with one exception at Aats Bay in 1966, were only visited at six month intervals. Most of the plants 58 were lost before any growth:measurement could be made. it is quite possible that if the Alaskan sites had been visited as often as the others a comparable number of growth measurements might have been obtained before loss of the plants.
Attachment of the holdfast to the rocks on which the plants had been placed occurred only at Stanley park (three L. sinclairii) River Jordan (two L. sinclairii); Whiffen Spit (two L_. sinclairii, one L. longipes) ; and Volga Island (one L. sinclairii). None of these places is very exposed, which is probably an important factor. More important, however, is the method of planting employed. On all of these sites, loose boulders are present and the plants were fixed to these by means of rubber bands. This method holds the bottom of the holdfast firmly against the rock. In the more exposed areas, which lack loose boulders, the method of fastening the plants to spikes in the rock only held the outplants from washing away but did not hold them in any one position for any length of time. Several unsuccessful attempts were made to devise a better planting method.
The results from the transplants to Stanley park, Whiffen Spit, and River Jordan indicate that L. sinclairii is less adversely affected by high summer seawater temperatures than is L. longipes. The results from Oregon and Alaska are not as clear cut. However the summer temperatures are not as high at these sites as in Oregon or Alaska, either. 59
2. Rock clearing a. Introduction
At various times over a period of one year, portions of rocks were cleared at Indian Beach and Short Sand Beach. The object of these experiments was to determine whether the adjacent L. sinclairii plants would colonize the cleared areas.
b. Methods In all experiments, portions of the L. sinclairii holdfasts were first removed with a knife. The area of rock thus exposed was then scraped and finally burned to within 5 cm of the remaining plants. Thereafter the clearings were examined at monthly intervals as far as possible.
c. Results In the first experiment, a strip about 35 cm wide was cleared between two large clumps of L. sinclairii at Indian Beach in May 1966. In June a slight growth of haptera tips toward the clearing from both sides was evident. In August some haptera had grown 2 cm onto the cleared area. By October the cleared area had been completely crossed by haptera from both sides and some stipes and blades had been produced from these haptera.
The second clearing at Indian Beach was made in December 1966. In February, there was no evidence of growth of haptera nor was there anything else growing on the cleared area. In March, two plants of Hedophyllum sessile each 15 cm long, one plant of Alaria (marginata ?) 12 cm long and several crustose coralline algae were present. 60
The third clearing experiment at Indian Beach was started in April 1967. A strip was cleared from 10 cm above the sand on one side of a rock, up over the top and down to within 10 cm of the sand on the other side (Fig. 16) . Six species were* removed from the rock. L_j_ sinclairii covered most of the area which was cleared, but there were occasional gaps in the holdfast which provided attachment space for other species. On top of the rock Phaeostrophion irrequlare was the second most abundant species. There were also several plants of Bossiella plumosa and an unidentified crustose coralline alga on top. On both sides Hildenbrandia sp. and Bossiella plumosa were found. On one side, extending down to below the sand level, was an extensive mat of Codium setchellii. In May, the haptera at the edges of the L. sinclairii had started growing toward the cleared area. Several patches of Hildenbrandia and a few tufts of Bossiella were evident (Fig. 16a). For the following three months the rock was completely buried under sand (Fig. 16 b,c,d). In September the sand receded, exposing the rock. At this time the L. sinclairii and Codium setchellii appeared healthy and unchanged from the June condition. Otherwise, only some dying tufts of Bossiella plumosa were noted.
At Short Sand Beach, the first clearing experiment was started in November 1966. By March 1967, several haptera of L. sinclairii had encroached onto the cleared area and were putting up stipes and blades in the area. Several tufts of coralline algae were also present. 61
Figure 16 Cleared rock at Indian Beach
a. Rock C in May 1967, one month after strip was cleared. b. Rock C in June 1967. c. Rock C in July 1967. d. Rock C in August 1967.
62
The second experiment at Short Sand Beach was begun in April 1967. A vertical strip was cleared next to the plants for which growth was being measured in situ. The rock at this location is soft sandstone with many holes produced by various animals. The surface was cleared in the usual manner, but the holes were not touched. One month later numerous tube worms were seen protruding from the holes in the rock. By July several encrusting animals had established themselves on the rock but no plants were present.
d. Discussion Several of the clearing experiments were carried out at times when the nearby L. sinclairii plants bore ripe sori. Under such circumstances it seems likely that gametophytes of L. sinclairii should have established themselves on the cleared areas. Although the gametophytes would not have been visible they should have produced sporophytes which would have been easily recognized. However, on all the cleared areas and several other rocks observed on these beaches over a period of two years, no young sporophytes of L. sinclairii were ever observed which had definitely arisen from gametophytes. In every instance, careful examination showed that the young sporophytes had arisen from haptera of older plants and thus were outgrowths of the older plants. As noted previously, two Hedophyllum plants and an Alaria became established on the rock although there were relatively few of these plants very close. This evidence indicates that production of L. sinclairii sporophytes by sexual means is rare. 63
On the other hand, the experiments show that colonization of new areas by outgrowth of haptera of L. sinclairii takes place readily, at least during the period when the plants are not buried under sand.
3. Transition zone experiments
a. Introduction
In most members of the Laminariales the transition
zone between the stipe and blade is generally stated to be the principal . region of growth. It appeared that this might not be the case in L. sinclairii. Accordingly, two types of experiments were conducted in the field to determine the importance of the transition zone. The first of these consisted of cutting off stipes well below the transition zone and observing the behavior of the remaining holdfasts and stipe stubs. This was done twice at Indian Beach and once at Short Sand Beach. The second experiment consisted of marking off regular intervals on a plant and then observing where the greatest growth occurred. This was done once at Indian Beach.
b. procedures and results In the first experiments in which transition zones were removed, stipes were cut at both Indian Beach and Short Sand Beach in October 1966. At Indian Beach all the stipes in a large clump on top of one rock were cut off at least 5 cm below the transition zone, using a large pair of shears. At Short Sand Beach the same procedure was followed except that the clump was first isolated by clearing a strip of rock around it. 64
In November, the stipe stubs on both beaches appeared to be healing over. In addition, at Short Sand Beach there was considerable growth of haptera at the margins of the clump. Late in December many of the cut stipes on both beaches had produced new blades from the tips in an apparently normal manner. At this time, a few of the normal uncut stipes at Indian Beach had blades with ripe sori, but the majority had no blades at all. At Short Sand Beach all the uncut stipes were without blades. In January no further changes in the cut stipes were noted. In February the blades were slightly longer than before and thereafter no further changes were noted at either beach.
In February 1967, the second cutting experiment was started at Indian Beach. This time all the large stipes in one group were cut off well below the transition zone, but all the present year's stipes and blades were left untouched. (The stipes of the present year are easily recognized, as previously noted.) After one month the cut stipes showed no change. The uncut small blades had grown about 3 cm past the cut ends with a total blade length of 6-7 cm. By the end of April about 20 percent of the cut stipes had produced new blades and the rest were unchanged. In June no more cut stipes had produced blades, but the growth of haptera at the margins of the clump
seemed to be greater than normal.
An experiment to determine the region of greatest growth was started in May 1967 at Indian Beach. In a clump of stipes where several stipes and blades were being measured 65 periodically, four blades were punched with many holes. In two of these, the holes were punched every 2 cm, beginning 4 cm above the base of the blade. In the other two, the holes were punched every 2 cm, beginning at 2 cm above the base. Both stipes and blades were measured, but no convenient way was found to mark off intervals on the stipes. In June 1967, only one of the four plants was found, one of those which had been punched from 4 cm above the base (Fig. 12c). The blade was eroded at the end, undoubtedly weakened by the holes, and the total length was less than at the beginning. The total growth of the blade as calculated from the total growth between all the holes was at least 6.3 cm (Table XXII). The stipe increased 1 cm in the same period. The results show that the greatest growth is in the transition zone. The total blade growth is less than 50 percent of the average for other blades in the same place at the same time (Table XXI), which is probably due to the injurious effects of the large number of holes.
C. Discussion
Setchell (1905), studying growth and regeneration in L. sinclairii, noted that a blade was produced from a wound on the side of a stipe. From his studies he postulated that L. sinclairii could regenerate from the stump of a stipe. The present experiments show that this is correct. The results show that although growth is greatest in the transition zone, it is not limited to this zone and the zone is not necessary for the production of new blades. 66
Under normal conditions the blades are lost in December and regenerated in January. On the stipes which were cut in October, new blades appeared in December, about one month earlier than normal. However, after this early beginning the blades showed little further development, at least in that year. Removal of the transition zone also seemed to stimulate the growth of haptera. It is possible that these results indicate the presence of an auxin or other substance in the old blade which inhibits new blade formation and growth of haptera. However, such a possibility has not been investigated, and at present no adequate explanation of these phenomena is available.
B. Laboratory Work
1. Sporophytes
Sporophytes of L. sinclairii and L. longipes were cultured for varying periods of time in the 120 iT'tanks of seawater previously described. The cultures were planned to determine the effects of different temperatures, salinities, and photoperiod on the growth and reproduction of the plants. New plants were collected and introduced into the tanks several times during a two-year period, plants were removed whenever they showed no change for a period of three months or were obviously deteriorating, a. photoperiod
Plants of both species were grown at photoperiods of 16 hours light/8 hours dark, 12 hours light/12 hours dark, 67
and 8 hours light/16 hour dark, plants were grown in each photoperiod for up to three months under each temperature regime (5°, 8°, and 10°C). The photoperiods were usually six months out of phase with those found in nature, i.e., short days in summer. The various photoperiods did not have any noticeable effect on either growth or reproduction of plants of the two species. After conclusion of the photoperiod experiments all cultures were run at 16 hours light/8 hours dark.
b. Salinity,
The Stanley park seawater was used as low salinity water and the West Coast seawater was used as high salinity water, plants were grown in each type of water at 5°, 8° and
10°C. No difference was evident in growth of the plants in the different salinities at the same temperature.
c. Temperature
Various plants were grown at 5°, 8°, and 10°C over a period of two years. During the first nine months the photoperiod was changed every three months. Thereafter it was maintained at 16 hourslight/Bhours dark. Maximum growth of both species occurred somewhat earlier in the year at 5°C than at 8° or 10°C. The maximum growth of L. sinclairii was in March at 5°C and in June at 8° and 10°c. The maximum growth of L. longipes occurred in January at 5°C and in
February at 8° and 10°C. With these exceptions, the different temperatures appeared to have very little effect on the growth of the plants. 68
In all plants cultured in the tanks, certain common features were evident. No variations in temperature, salinity, or photoperiod had any effect on production of sori or loss and regeneration of blades. Sori were produced on L. sinclairii plants collected in late September after one week in culture. However, other plants of the same population became ripe at the same time in nature. Blade loss always occurred at the same time as in nature unless the loss was due to the
plant's dying. Likewise, blade regeneration occurred at the same time as in nature at all temperatures, although the rate of growth of the new blades varied with temperature. After new blades were produced, the "collars" (Setchell, 1896) formed by the frayed ends of the split stipe ends were much more obvious than in the field, presumably because there was no wave action to remove the excess tissue.
Generally, the plants which grew in the laboratory tanks grew well and showed normal behaviour for two to four months and then ceased showing blade growth. At this time some of
them then began to deteriorate; but in most, growth of haptera and production of new stipes and blades from the holdfasts continued for up to 15 months after blade growth had ceased. Many of these haptera were removed and used in the experiments on haptera described below. Although new stipes were produced from the haptera, no growth was recorded on any of the stipes which were already present on plants when brought into the laboratory. 69
d. Discussion The tank results are valuable in that they indicate that production of sori and blade loss and regeneration are not controlled by temperature, salinity, or photoperiod. However, the results in the tanks cannot give a very good picture of the growth of the plants. The plants apparently behaved normally for a short time after being placed in the tanks. However, plants were not placed in the tanks often enough to give a true picture of growth over a whole year. The tanks contained water very similar to that in which the plants normally grow, but this water was only changed once a month. This allowed a much greater buildup of waste products than occurred in nature and probably a considerable depletion of various nutrients. This, and the lack of tides and wave action, make the tanks very poor substitutes for the natural conditions.
2. Gametophytes
a. Introduction At the beginning of this study it appeared that L. sinclairii and L. longipes were possibly ecotypes of the same species. In order to test this hypothesis, it was planned to cross the two species. To accomplish this, it was necessary to obtain sexually mature gametophytes in culture. Gametophytes of L. sinclairii were cultured by Myers (1925). In her cultures, the gametophytes produced sporophytes, thus indicating a typical kelp life history. However, she noted that 70
the gametophytes continued to grow vegetatively and, after six months, were larger than any of the sporophytes produced in culture. So far as is known, the gametophytes of L. longipes have never been cultured previously.
b. Materials and Methods Sporophytes bearing ripe sori were collected at various times during the winters of 1965, 1966 and 1967 (L. longipes was only collected in December) and brought into the laboratory. They were then kept in the dark in a moist plastic bag at 5°, 8°, or 10° C for four to 36 hours, then in a paper towel for two to eight hours at the same temperature. After this period of partial drying the sori were wiped carefully to remove epiphytic contaminants and then cut into small pieces with a razor blade. These pieces were placed in SWF or ES in 250 ml glass culture dishes or standard 100 mm Petri dishes. The dishes were placed in 5°, 8°, or 10°C under 20 ft-c or 150 ft-c. After intervals which rangedfrom six hours to six days in various experiments (apparently depending upon the stage of development of the sorus at the time of collection) zoospores could be seen swarming from the pieces of sorus. Several milliliters of the spore suspension were pipetted into each culture dish. A large number of different combinations of culture conditions were tested.
Five different media were employed at various times: SW, SWF, ES, ES+, and ASP 2. Large glass culture dishes (250 ml), standard glass Petri dishes (100mm) and small (60mm) Petri dishes of both glass and plastic were used. In various 71 experiments the medium was changed at intervals of 1, 2, 4, 7, 10,' 14, or 28 days. Most cultures were kept at least one year. After a culture had run for two to three months the medium was usually changed only once a month. The cultures were maintained at 5°, 8°, 10°, and 15°C under light intensities of 20 ft-c or 150 ft-c provided by cool white fluorescent lights. Three photoperiods were tested: 8 hours light/16 hours dark, 12 hours light/12 hour dark, and 16 hours light/8 hours dark. In one experiment gametophyes of various ages were maintained for periods of one day to two weeks in total darkness. Cultures were also kept in an east-facing window where the temperature varied from 10° to 25°C. Some cultures were run on the shaker in the psycrotherm incubator. In several instances cultures were also subjected to changes of temperature, media, frequency of media change, light intensity, photoperiod and various combinations of these factors.
It was not possible to carry out any crossing experiments, "for v-easot\s hottd below, c. Results
Under several different conditions sporophytes were produced by L. sinclairii gametophytes, but never more than a
few in any one culture. L. longipes gametophytes only produced sporophytes twice: once in 1966 after 10 weeks in ES at 10°C under 20 ft-c (five sporophytes) and once in 1967 after 24 weeks in ES+ at 5°C under 150 ft-c (three sporophytes). The various differences in culture conditions seemed to have little effect on the production of sporophytes. In the ES+ medium the gametophytes had larger, more deeply pigmented cells than in the other media. Longer and less deeply pigmented 72 cells were produced under higher light intensities. At 5°c the gametophytes had fewer and smaller cells. However, none of these differences appeared to affect the production of oogonia, antheridia, and ultimately, sporophytes. Under most conditions the gametophytes continued growing vegetatively and appeared healthy after more than one year in culture.
In most instances the male and female gametophytes of L. sinclairii were easily distinguished. As is typical of kelp gametophytes, the males have smaller, more numerous cells and many branches, whereas the females consist of a few larger cells, and few branches. In several instances, oogonia and antheridia were noted. In L. longipes, however, a different situation prevails. The gametophytes are apparently all of one morphological type and are typical of neither male nor female kelp gametophytes. The cells are of intermediate size and the branches, although numerous, are not as numerous as in male gametophytes of L. sinclairii. No antheridia or oogonia were ever noted. In L. longipes it was thus impossible to distinguish males from females.
d. Discussion It is possible that the gametophytes of L. longipes are bisexual but until antheridia and oogonia arefound, this will remain in question. With the exception of Chorda tomentosa (Sundene, 1963), the gametophytes of all kelps reported are unisexual.
The crossing experiments were not carried out because two essential conditions were lacking. The gametophytes of 73
11- longipes could not be distinguished from each other and thus males and females could not be isolated. Secondly, no conditions were found under which either species would produce sporophytes regularly, which would have served as a control.
In previous experiments the author has cultured gametophytes of Laminaria saccharina (L.) Lamour., Hedophyllum sessile (C. Ag.) Setch, and Nereocystis lluetkeana (Mert.) P. and R. under the same conditions of temperature, light, and media employed in the present study. In many instances abundant sporophytes were produced. Robinson (1967) cultured several species of Alaria under the same conditions in the same culture rooms at the same time that L. sinclairii and L. longipes were being cultured. One species (Alaria marginata p. and R.) was obtained on the same Oregon beaches as L. sinclairii. In most of Robinson1s cultures sporophytes were produced abundantly. The present study indicates that, relative to other members of the Laminariales, the sexual production of sporophytes by gametophytes of L. sinclairii and L. longipes is very limited.
3. Haptera
a. Introduction Field and laboratory observations show that there is vegetative production of new blades and stipes from the haptera at the edges of the holdfasts of L. sinclairii. This phenomenon was studied further using laboratory cultures of isolated pieces of haptera (Markham, 1968). 74
b. Materials and Methods Sporophytes of L. sinclairii growing in the large culture tanks provided a source of haptera for use in the experiments. After two to three months in the tanks, many haptera had grown out from the bottoms and edges of the holdfasts, especially at 8° and 10°C. These haptera were removed from the parent plants, washed in filtered seawater, and cut to a uniform length of 20 mm. In the experiments testing the effect of initial size, pieces 2.5, 5, 10, and 15 mm long were used. For all sizes the haptera either had one growing tip or were cut off at both ends; in the latter instance, the most distal end was at least 20 mm back of the original apex of the hapteron. The pieces with one cut end and one growing tip were designated "E haptera", whereas those with both ends cut off were designated "1J haptera".
Preliminary experiments were run to test the effects of various media and culture dish sizes. All of these experiments were run at 8° and 10°C and 150 ft-c. Four media were employed: ES+, ES, SWF, and SW. Two types of culture dishes were tested: 250 ml glass culture dishes containing 200 ml of media and 20 mm plastic Petri dishes containing 15 ml of media. The media were changed at 2-week intervals. These experiments showed that the maximum percentage of haptera produced outgrowths in 250 ml dishes containing 200 ml of ES+.
After the preliminary results were obtained, experiments were run to test the effects of shaking and sand scouring. In 75 these experiments 20 mm pieces of haptera were secured to small rocks with rubber bands around the middle portions of the haptera. These rocks were then placed in 250 ml culture dishes. Generally the rocks were large enough so that only one could fit in each culture dish. For the shaking experiments, the dishes were then filled with just enough ES+ to cover the rock and its attached haptera. These were then placed in the Psycrotherm incubator shakers and the speed of shaking adjusted so that the medium reached the rim of the culture dishes but did not spill over. For the scouring experiments sterilized beach sand from Oregon was added to the culture dishes until it covered the rocks but not the haptera on top of the rocks. The dishes were then filled with ES+ until the haptera were just covered and the dishes were placed on the shakers with those in the shaking experiments. The water motion caused the sand to wash back and forth over the haptera. Two pieces of haptera of L. longipes were also cultured on the shaker, each on a rock without sand. In the incubator shakers the temperature was 8°C, and the light intensity was 150 ft-c, with a photoperiod of 16 hours light/ 8 hours dark. Control experiments were run in a second Psycrotherm incubator without shaking. The medium was changed every two weeks.
A 3-cm piece of stipe was cut out of each of two plants of L. sinclairii, halfway between the holdfast and blade (approximately 10 cm of stipe was cut off each end). These pieces were then fastened to rocks and placed in culture 76 dishes in the same manner as the haptera. One dish contained sand and ES+, the other only ES+. These were cultured without shaking in the control psycrotherm. All haptera were cultured for 14 weeks and stipes for 20 weeks.
These experiments were preliminary in that they were simply designed to determine whether it was possible for haptera to produce blades and stipes at all under such conditions. For this reason results were recorded only at the end of the experiments and then only as presence or abs_ence of blades.
The final series of experiments tested the effects of temperature, light intensity, and initial size of haptera. All these culture experiments were carried out in 250 ml glass culture dishes containing 200 ml of ES+ medium. The medium was changed every two weeks except in the experiments testing very small sizes. In the latter instance it was changed every four weeks. Experiments on the effect of temperature were run at 5°, 8°, 10°, 15°, and 20°C. All other experiments were run only at 8° and/or 10°C. Light intensities of 2 ft-c (21.5 lux), 150 ft-c (1614 lux), 250 ft-c (2690 lux), and 500 ft-c (5380 lux) were employed in light intensity experiments. All other experiments were run under 150 ft-c only. The photoperiod for all experiments was 16 hours light/8 hours dark. All these experiments were run for ten weeks. The resulting data were sorted with an IBM 113 0 computer. In the final analysis, all 20 mm haptera were analyzed for their responses to light and temperature (Fig. 17a) whereas the effects of initial size were analyzed separately. 77
Figure 17 Results of growth experiments on haptera of
L. sinclairii
a. percentage of E and N haptera producing blades, haptera, and growth in length at various temperatures and various light intensities.
b. 2.5 mm pieces of N. haptera, cultured at 10°c under 150 ft-c, showing lateral blades.
c. 20 mm pieces of E and N haptera, cultured at 10°C, under 150 ft-c, showing lateral and terminal blades and one lateral hapteron.
d. 20 mm pieces of E and N haptera, cultured at 10°C, under 2 ft-c, showing lateral and terminal blades and lateral haptera.
e. Nonmedian section of the stipe of a blade arising as
a lateral outgrowth from a hapteron, x 70 (Diameter of hapteron is 1 mm). f. An enlarged portion of e, x 210.
All results at 10 weeks, b, blade; co, cortex,
h, original piece of hapteron; lh, lateral hapteron arising
from original hapteron; me, meristoderm; st, stipe.
78
c. Results The results of the culture experiments were varied. The haptera either showed no change; grew in length; produced lateral haptera, i.e., they branched (Fig. 17c,d); or they produced lateral or terminal blades (Fig. 17b,f,c,). Normally a blade outgrowth appeared first as a small blade, and two to three weeks later a small stipe developed between the blade and the parent hapteron. Sections indicate that only the outer cortex is involved in production of lateral outgrowths from haptera (Fig. 17e,f).
In the shaking experiments blades were produced in both the shaking and control cultures and the shaking appeared to have little effect. The haptera of L. longipes produced blades in the same manner as the haptera of L. sinclairii.
The sand scouring experiments showed that both branches and blades can be produced under conditions of constant scouring, although the number and size of outgrowths are less than in the control experiments. In two instances in the scouring conditions, a hapteron produced one branch which bent downward and attached firmly to a rock.
The two pieces of stipe each produced outgrowths from both ends. Although somewhat deformed, the outgrowths were recognizable as blades. The stipes had been held horizontally in the dishes, and the blades all bent upward. There is a strong indication that the stipe lacks any internal polarity. However, further experiments on polarity were not conducted. 79
In most of the experiments, different results were obtained for E haptera and N haptera; for blade outgrowths and haptera outgrowths; and for lateral blade outgrowths and terminal blade outgrowths. A few generalizations can be made. The N haptera never grew in length under any conditions. When E haptera grew, the growth always occurred at the uncut end. A greater percentage of E haptera produced outgrowths than did N haptera under any given set of conditions. The N haptera never produced both haptera and blade outgrowths on the same hapteron, whereas E haptera occasionally did.
For original haptera lengths of less than 15 mm, lateral blades were produced by N haptera, whereas terminal blades were produced by E_ haptera. For the standard 20 mm haptera, N haptera generally produced lateral blades whereas E haptera produced lateral and/or terminal blades. Terminal blades generally appeared earlier than lateral blades. In the final analyses of the results, outgrowths were considered merely as blades or haptera, with no regard to whether they were lateral or terminal.
i. Temperature - At 20°C there was a considerable growth of various contaminants. No positive growth response was shown by any of the haptera and several began to disintegrate. It is concluded that 2 0°C is a lethal temperature. Within the limits of 5° to 15°C, various results were obtained.
The greatest percentage of E haptera grew in length and produced lateral haptera or branches at 10°C (Fig. 17a). No growth occurred at 15°C. The greatest percentage of N haptera 80 produced lateral haptera at 8 C. The percentage of E haptera producing blades rose with increasing temperature, with the maximum of 100 percent at 15°C. The maximum percentage of N haptera produced blades at 5°C. The average number of blades per E hapteron which produced blades was highest at 8°c, or intermediate temperature, whereas for N haptera it was highest at the extremes of temperature, 5° and 15°C (Fig. 17a).
ii. Light Intensity - The greatest percentage of
E haptera grew in length and produced lateral haptera at 150 ft-c (Fig. 17a). No growth occurred at 500 ft-c. The greatest percentage of N haptera produced lateral haptera at 2 ft-c. The percentage of E_ haptera producing blades rose with increasing light intensity to a maximum of 100 percent at 500 ft-c. The maximum percentage of N haptera produced blades at 250 and 500 ft-c. The average number of blades per E_ hapteron which produced blades rose steadily with increasing light intensity to a maximum at 500 ft-c.(Fig. 17a). For N haptera the number of blades was highest at 2 ft-c, but the next highest occurred at the other extreme, 500 ft-c.
Haptera tended to arise at lower light intensities and in instances where both haptera and blades appeared at low light intensities, the haptera generally appeared first.
iii. Initial Size - The size experiments showed that even pieces of haptera only 2.5 mm long can produce outgrowths of normal appearance (Fig. 17b). However, two effects of the small size were noted, pieces of haptera shorter than 15 mm 81 produced blades, but never haptera. Secondly, the size of the outgrowth appears to be correlated with the size of the original hapteron; larger pieces produce larger outgrowths.
d. Discussion The differences between the E haptera with one growing tip and the N haptera with no intact tips can be interpreted in a number of ways. The effect may be chiefly due to injury caused in removing the ends, as indicated by the scarcity of terminal outgrowths from the N haptera. It may be due to the relative age of the pieces of haptera, since N haptera are farther from the growing tip and thus older. It could be expected from this that the N haptera would have less potential for redifferentiation and would thus produce fewer outgrowths. Such was observed to be the case in this study. However, field studies also showed that an entire plant can be cut in half and new haptera will grow out from the center, or oldest, haptera. Yet another possibility, which has not been investigated, is that some substance such as an auxin, which stimulates haptera growth and blade production is produced or stored in the apex. Several authors have reported the presence of auxins in various'members of the Laminariales (Van Overbeek, 1940a, b; Williams, 1949; Mowat, 1965) and one of them (van Overbeek, 1940b) reported auxin specifically in the haptera of Costaria costata.
Several authors, most recently Dawson (1966), have noted that L. sinclairii reproduces vegetatively from a rhizome-like holdfast. The present study has shown that the haptera need 82
not be attached to the parent plant for such reproduction to occur. Even very small pieces of haptera can produce blades and stipes and thus ultimately a whole new plant. Assuming that this could take place in nature after a portion of a hapteron is accidentally cut loose from the parent plant, the E type of hapteron, that is, one which has only been cut once, seems much more likely.
The conditions under which the maximum number of blades are produced per hapteron are not the same as those under which the maximum number of haptera produce blades. The latter would seem to be the more important consideration, if this production is regarded as a means of vegetative reproduction. Each piece of hapteron, no matter how many attached stipes and blades it has, is still only part of one plant.
The experiments indicate that 20°C is a lethal temperature, but the highest light intensity tested, 500 ft-c, is not harmful. This is to be expected from observed natural conditions. A water temperature of 20°C is very unlikely on the Oregon Coast, whereas light intensities greater than 500 ft-c do occur. Separate temperature and light intensity experiments show a maximum blade production at 15°C and 500 ft-c. However, all temperature experiments were run at 150 ft-c and all light intensity experiments at 10°C, so that it may be incorrect to assume that a combination of high temperature and high light intensity would result in high blade production. 83
Nevertheless, if one makes this assumption, a maximum blade production should occur in early summer when the water temperature is rising to 15°C and the light intensity is still high because the plants have not yet been buried under sand. Field studies indicate this is indeed the case. The scouring experiments indicate that such production can occur even if the haptera are partly buried in sand, provided there is sufficient light. 84
VII. TAXONOMY OF L. SINCLAIRII AND L. LONGIPES
A. Introduction
L. sinclairii and L. longipes are very distinct from all other species of the genus. The most similar of the other species may be the Mediterranean plant, L. rodriguezii Bornet, because of its multiple stipes. However, illustrations of L. rodriguezii indicate that although it has multiple stipes, they are very few and widely separated, unlike the situation in L. sinclairii and L. longipes. Furthermore, unlike all other species of Laminaria, it is restricted to very great depths (100-150 m) (Bornet, 1888).
L_. sinclairii and L. longipes are much more similar to each other than to any other species in the genus. Nevertheless, they are distinct from each other. Previously published reports have emphasized the presence or absence of mucilage ducts in the stipe for distinguishing the two species. The present study has revealed at least five differences between the species which have not been previously reported. These are blade width, seasonal loss of blades, morphology of gametophytes, and temperature tolerances. On the basis of these and other differences reported below, it is clear that they are two separate taxa. 85
B. Laminaria sinclairii (Harvey ex Hooker f. and Harvey)
Farlow, Anderson and Eaton, 1877-1889.
Harvey, 1852, p. 87 (as Lessonia sinclairii) Farlow, Anderson and Eaton, 1878, fasc. 3, p. 118 Areschoug, 1883, p. 6 (as Hafqyqia sinclairii) Anderson, 1891, p. 220 Howe, 1893, p. 67 De Toni, 1895, p. 343 Collins, Holden and Setchell, 1895-1919, fasc. 7 Setchell, 1896, pp. 44-46; 1905, pp. 139-169; 1912 pp. 131, 134, 137, 140, 141, 148, 150 Myers, 1925, pp. 114-116 Setchell and Gardner, 1925, p. 598 Okamura, 1932, p. 73 Smith, 1944, p. 135, pi. 31 Doty, 1947, p. 40 Sanborn and Doty, 1947, pp. 9, 13, 21, 30 Shchapova, 1948, pp. 99, 100, ll7, 120 Scagel, 1957, p. 98 Silva, 1957, pp. 43,44 Dawson, 1958a, p. 66; 1958b, pp. 186, 188, 201, 204; 1958c, pp. 235, 238, 242, 260; 1959, pp. 144, 161, 162; 1961, p. 396 Hollenberg and Abbott, 1966, p. 25 Druehl, 1968, p. 541 Markham, 1968, p. 125-131 1. Description Sporophytes up to 3 m long, perennial from the holdfast and stipe, regenerating new blades after complete loss of old ones. Holdfast an extensive rhizome-like system of branched haptera, sometimes covering an area of 5 m^ or more. The haptera giving rise to many stipes, more than 100 in some instances. Stipes flexible, cylindrical, l-3mm in diameter, up to 60 cm long or greater, mucilage ducts present. Blades narrowly linear, usually less than 3.0 cm wide, of variable length up to 2.5 m, entire, without bullae, mucilage ducts present. Blades generally lost in December, regenerated in
January, plants tending to be larger near the southern limits 86 of distribution. Sori in patches, oblong to irregular in outline, one to many per blade, plants produce sori in October, November, January, February, and March. Gametophytes filamentous, capable of prolonged vegetative growth, unisexual, with males and females distinctly different, rarely producing sporophytes.
2. Distribution Hope Island (50°55'N, 127°58'W) British Columbia to Ventura County, (34°19'N, 119°23.3'W) California.
3. Habitat Growing on rocks in the lower intertidal region, usually partly buried under sand in summer, usually in fully exposed areas, but occasionally in moderately sheltered to moderately exposed areas.
4. Comments
The mean length of the longest stipes from each of 135 plants collected in Oregon is 26.3 cm. The longest measured stipe in this group was 52 cm and the shortest was 11 cm. Plants lower in the intertidal region are generally longer but were rarely collected because of difficult surf conditions. Consequently, the mean length of all stipes on the Oregon beaches is probably greater than 27 cm. plants of L_. sinclairii in California are known to be generally larger than those in Oregon but no comprehensive data are available. The mean width of the blades on the largest stipes of 54 87 pressed specimens from Oregon is 1.75 cm. From measurements of both fresh and pressed specimens from Indian Beach, the shrinkage after pressing is calculated to be 20%. Applying this correction factor, the mean blade width.for all specimens examined is approximately 2.2 cm.
C. Laminaria longipes Bory, 1826
Agardh, C, 1820, p0 133
Bory, 1826, vol. 9, p0 189 Postels and Ruprecht, 1840, p. 10 (as L. saccharina f. angustifolia) Kiitzing, 1849, p. 574 Ruprecht, 1851, p. 232, 350 (as Lessonia repens) Le Jolis, 1855, p. 307-308,311 (as L. ruprechtiana) Agardh, J., 1867, p. 26 (as Arthrothamnus ? longipes) Areschoug, 1883, p. 15 Kjellman, 1889, pp. 7, 9, 17, 43 De Toni, 1895, p. 370 (as Ar thro thamnus ? longipes) Setchell, 1899, pp. 591, 592, pi. 95 Setchell and Gardner, 1903, p. 260 Yendo, 1909, p. 215; 1910, p. 295 Setchell, 1912, pp. 136, 148, 150 Okamura, 1916, p. 172 Setchell and Gardner, 1925, p. 597 Okamura, 1928, p. 53, pis. 13-15 Arwidsson, 1932, p. 153 Miyabe and Nagai, 1932, pp. 196, 197; 1933, pp. 86, 87 Okamura, 1932, p. 72; 1933, pp. 88, 95 Zinova, E., 1933, pp. 24, 25, figs. 8-10 Okada, 1934, p. 46, pi. 43 Yamada, 1935, pp. 1, 6, 7, 18, pi. 6 (as L. longipes var. latifolia) Okamura, 1936, p. 251, fig. 139 Nagai, 1940, pp. 67-70; 1941, p. 262 (as L. longipes f. angustifolia, f. linearis, & f. latifolia) Shchapova, 1948, pp. 93, 98, 100, 120, 127, 130, 131 Tokida, 1954, pp. 30, 114, 115 (incl. L. longipes f. typi Zinova, E., 1954, p. 378 Zinova, A., 1959, p. 153 Dawson, 1961, p. 396 Vozzhinskaya, 1964, p. 425 Druehl, 1968, p. 541 88
1. Description Sporophytes up to 1 m long, perennial from the holdfast and stipe, regenerating new blades while remnants of the old ones remain. Holdfast an extensive rhizome-like system of branched haptera, sometimes covering an area up to 5 m2. The haptera giving rise to many stipes, more than 100 in some instances. Stipes flexible, cylindrical, 1-3 mm in diameter, up to 40 cm long, usually not exceeding 20 cm in length, mucilage ducts absent. Blades narrowly linear, usually less than 5.0 cm wide, of variable length up to 80 cm, entire, without bullae, mucilage ducts present. Blades generally shortest in December or January, but not lost completely. Sori in patches, round to irregular in outline, one to many per blade, plants produce sori in December; no information is available for other winter months. Gametophytes filamentous, capable of prolonged vegetative growth, apparently all of one type, very rarely producing sporophytes.
2. Distribution Urup Island (46°00'N, 150°00'E) Kurile Islands around o o pacific Rim to Coronation Island (55 49.6'N, 134 17'W) Alaska.
3. Habitat Growing on rocks in the lower intertidal and upper subtidal regions, usually in moderately exposed or moderately sheltered areas, but occasionally in fully exposed areas. 89
4. Comments
, The mean length of the stipes measured on herbarium specimens from 33 locations is 8.4 cm. The average stipe length is greater on plants from more exposed sites (Table XXIII). However, the longest stipe recorded, 3 5 cm, is on a plant from an area which is only "moderately exposed". The mean width of the blades of pressed specimens from 33 locations is 1.97 cm. From measurements on both fresh and pressed specimens from Aats Bay, the shrinkage after pressing is calculated to be 37%. This is greater than that for L. sinclairii. However, examination of other species from Alaska indicates a greater shrinkage in them also. It is believed that the concentration of formalin used to preserve the plants was not the same for Oregon and Alaska. Applying this correction factor, the mean blade width for all specimens examined is approximately 3.1 cm. According to some authors, (e.g., yamada, 1935) some plants in the Kurile Islands have blades considerably broader than this.
D. Comparisons of Species
L. sinclairii and_L_. longipes are two distinct species. The present study has shown that there are at least 10 differences between the two species, as follows: 90
L. sinclarii L. longipes
1. Mucilage ducts present in 1, Mucilage ducts absent from stipe. stipe. 2. Stipe in mature plant 2, Stipe in mature plant usually usually over 20 cm long. less than 20 cm long.
3. Blade of mature plant 3, Blade of mature plant usually less than 3 cm wide. usually more then 3 cm wide.
4. Entire blade lost and then 4, Proximal remnant of blade new one regenerated. retained and new one produced while remnant still present. 5. Male and female gameto- 5. Gametophyes in culture phytes in culture morphologically indisting• morphologically different. uishable as to sex. 6. Occurs from Southern 6. Occurs from Southeast Alaska California to Central throughout the Aleutian British Columbia. Islands to the Kurile Islands. 7. Optimal growth at mean 7. Optimal growth at mean temperature higher than temperature lower than 8 c 8°C. 8. Grows in lower intertidal 8. Grows in lower intertidal region. and upper subtidal regions. 9. Usually occurs on fully 9. Usually occurs on moderately exposed sites. exposed or moderately sheltered sites.
10. Usually associated with 10. Rarely associated with sand. sand and buried for part of year.
Figure 18 illustrates the differences in blade width and
stipe length. Extreme examples were chosen to illustrate
the point. The difference is usually not as great as in
figure 18. As noted above, points 3, 4, 5, 7, and 9 have not
been reported before; the others have been mentioned, usually 91
Figure 18 Habit of L. sinclairii and L. longipes
a. L. sinclairii from Pescadero Point, California, x 1/3.
b. L. longipes from cape Spencer, Alaska, x 1/3.
92 without emphasis, by various authors. Except for differences in gametophytes and mucilage ducts, none of these points alone would seem sufficient as a basis for separating the two species. However, all of these differences collectively confirm that L. sinclairii and L. longipes should definitely be retained as two separate species. 93
VIII. GENERAL DISCUSSION AND CONCLUSIONS
The two species studied, L. sinclairii and L. longipes, are very distinct from all other species of the genus with the possible exception of L. rodiguezii as noted. They are also different from most other members of the order Laminariales. There are three features of these two species which are very distinctive. There is an apparent suppression of sexual reproduction. They have a great potential for dedifferentiation of supposedly mature tissues and related to this, a more diffuse meristematic area which allows considerable growth in regions other than the transition zone, especially in the haptera.
A few other kelps exhibit considerable growth in regions other than the transition zone. Egregia spp. develops a very complex thallus primarily by lateral outgrowths from the flattened stipe, with very little growth occurring in the transition zone. In Dictyoneurum californicum Rupr. the stipe becomes prostrate and forms a rhizome-like structure which attaches to the substrate by lateral haptera. Repeated splits in the blade cause segmentation and the formation of many blades, each with a stipe that becomes prostrate so that a clump is formed (Setchell and Gardner, 1925). In Arthrothamnus bifidus (Gmel.) J. Ag. the basal margins of the blades become meristematic and numerous secondary blades arise. The process is repeated many times, forming a rhizome-like structure bearing many blades (Yamada, 1938). The most 94 similar development to that of L. sinclairii is found in the Japanese species Ecklonia stolonifera Okam. The haptera grow into stoloniferous structures and produce new blades at the tips. The blades are deciduous (Okamura, 1915). Okamura states also that with this vegetative reproduction, the formation of zoosporangial sori seems to be suppressed. However, Ecklonia differs from Laminaria in having lateral outgrowths from the blades.
With the possible exception of Ecklonia stolonifera, the suppression of sexual reproduction in the two species studied appears to be unique in the kelps. Sorus production is not suppressed in these two species but there is little evidence that the resulting gametophytes normally produce sporophytes in any number.
In view of the differences between L. sinclairii and L. longipes and other species of the genus, it is possible that they should be removed from the genus Laminaria. One of the chief reasons for removing them from the genus is the meristematic activity of the .haptera which results in multiple stipes. On this basis, however, the two species would have to be removed not only from the genus Laminaria, but also from the family Laminariaceae, as the families of the order Laminariales are based on the type of growth in the meristematic area. Thus, before such a taxonomic revision could be made, a comprehensive review of the entire order would have to be carried out. For this reason, no changes of taxonomy are proposed at the present time. 95
Despite the almost identical external morphology of L. sinclairii and L. longipes, and the many common features by which they differ from other kelps, the two species are also distinct from each other, as has been demonstrated. Many of these differences may be the result of adaptation to different habitats. The most striking difference between the habitats of the two species is in the temperature of the seawater and the air. The mean temperatures for the coldest winter months and the annual means show no overlapping values at all (Table I, Fig. 2, 3). In summer the difference is not as great, but there is only a slight overlap for seawater temperature and none for air temperature. The evidence indicates that sexual reproduction is of very little importance to these species. Nevertheless, it may be significant that the most marked difference in temperature between the areas where the two species are found occurs at the time of year when both species bear ripe sori.
It was observed that plants of L. sinclairii collected in California containedmuch more mucilage and became very slimy soon after collection. The plants collected in Oregon did not become slimy at all. Several authors have stated that kelps growing in the intertidal zone are protected from rapid desiccation by the presence of mucilage. The air temperature and degree of insolation is much greater on California beaches than on Oregon beaches. Hence the danger of desiccation is much greater in California. The greater production of mucilage in the California plants is probably an adaptation in response 96 to this. In Alaska, where L. longipes is found, the average air temperature and the danger of desiccation are considerably less than in Oregon. L. longipes does not produce mucilage to any noticeable degree. Furthermore, it lacks mucilage ducts in the stipe. No definite correlation has been demonstrated between the presence of mucilage ducts and the production of mucilage. Nevertheless, considering the value of mucilage in protection against the effects of high temperatures and Burrows' (1964) demonstration that mucilage ducts tend to be produced only at higher temperatures, it is quite possible that such a correlation may exist. Thus the difference in mucilage ducts in the two species may have arisen originally as a response to different temperatures, even though it can apparently no longer be altered by changing the temperature.
Temperature may be the factor which limits the distribution of L. sinclairii south of Ventura County, California. At Santa Monica, just south of the southern limits of L. sinclairii, both the summer mean temperatures and the yearly mean temperatures are more than 1C° greater than in Ventura County.
At Short Sand Beach the plants of L. sinclairii were regularly immersed for several hours in almost fresh water during the latter part of the summer. Many of the red algae growing on the same rocks died in large numbers during this time. L. sinclairii showed reduced growth and considerable loss of blades but was apparently not irreparably damaged. 97
These observations, together with the results in laboratory- culture and the great variation in salinity within the ranges of both species indicate that salinity is of little importance in controlling the distribution of either species. L. longipes is not normally found associated with sand, although there is no evidence from this study to indicate that it should be any less well adapted to sand than is L. sinclairii. The chief reason may be simply that within the area to which
L. longipes is adapted by its temperature requirements there are very few sandy beaches of the type found in Washington, Oregon, and California.
The chief factor controlling the distribution of L. sinclairii within its temperature range appears to be the presence of sand. It is nearly always restricted to beaches which have a large seasonal fluctuation of sand and where it is periodically buried. It is well adapted to this very harsh environment. In such an area, sexual reproduction by gametophytes is difficult because of the danger of sand scouring. Further, in such a well-adapted plant a nonsexual means of reproduction would ensure successive generations of genetically similar, well-adapted plants. Three other species which are similarly well-adapted to this type of environment apparently lack sexual reproduction altogether: Gymnogongrus linearis (Smith, 1944), Ahnf eltda concinna (Smith, 1944), and
Phaeostrophion irregulare (Mathieson, 1967). Laboratory experiments show that sexual reproduction in L. sinclairii is suppressed even in environments where sand scouring is absent. 98
The need for an alternative, or accessory, nonsexual means of reproduction in this plant is therefore apparent. This study has shown that such a means exists in the production of new stipes and blades from haptera and can operate even if the haptera are detached from the parent plant. The generalized meristematic activity and the considerable regeneration in this plant serve a further adaptive function in enabling the plant to recover after various parts have been accidentailyremoved. This is a great advantage in an environment where sand scouring and the force of the surf are often very great. As L_. sinclairii is rarely found where it does not undergo periodic burial under sand, it would appear that this burial confers some advantage on the plant. its unique methods of growth and reproduction enable it to survive in this very hostile environment which excludes most other algae. Colonization by outgrowths from haptera is very successful in areas which are periodically buried under sand because spores or gametophytes would be scoured away. in areas which are not buried under sand, colonization by gametophytes or spores is more successful because a much larger area can be colonized in a single season. Since L. sinclairii apparently reproduces by gametophytes very rarely, it is excluded from areas where such reproduction is an advantage.
Colonization by haptera can occur easily only on areas of rock adjacent to the area where the plant is already growing. Colonization of new and isolated rocks is a more 99 difficult problem. It is possible that new plants of L. sinclairii are initiated from gametophytes on such areas. The release of spores at a time when little other algal growth is occurring and sand is mostly absent, and a subsequent rapid growth of gametophytes and production of young sporophytes, might enable the plants to become established before competition and/or sand scouring eliminated them. It is assumed that young sporophytes are much more resistant to sand scouring than are gametophytes.
It was noted that ;L. sinclairii appears to be adapted to exposure to severe surf. in the areas of greatest exposure, the plants are larger and continue growing longer after burial by sand. Also, in these areas L. sinclairii shows more complete dominance on the rocks on which it grows. Thus, at Arch Cape, on the entire rock where L. sinclairii was studied, only one other species was found. L. sinclairii can withstand burial better than many smaller plants because it is long enough by late summer so that a portion of the blades usually protrudes from the sand. It is not known to what extent, if any, conduction occurs from the exposed to the buried parts of the plant. When the plant is buried, greater exposure to surf might be an advantage in that greater surf action would cause greater stirring of the sand and at the same time allow greater water motion and aeration around the buried parts of the plant. 100
L. sinclairii has been shown to be uniquely adapted to a very harsh environment. The environment of L. longipes generally lacks sand. However, it too is a very harsh environment, in part because of the low seawater temperatures and even lower air temperatures to which the plant is frequently subjected. Further studies of the type carried out on L. sinclairii are needed to determine exactly what environmental factors are most significant in controlling L. longipes and the ways in which the plant is adapted to its environment. Such information should also clarify further the relationship of L. longipes to L. sinclairii and the relationship of these two unique species to other kelps. 101
IX. SUMMARY
The distribution, ecology, growth, and reproduction of Laminaria sinclairii and L. longipes were studied in the laboratory and on beaches in Alaska, British Columbia, and Oregon over a two-year period. The two species differ from most other kelps in having multiple stipes arising from an enlarged rhizome-like holdfast, composed of many haptera. Each of the stipes bears a blade. The two species are almost identical in external morphology, previous workers have distinguished the two species primarily on the internal anatomy of the stipe: the stipe of L. sinclairii has mucilage ducts whereas that of L. longipes does not. L. longipes occurs from the Kurile Islands through the Aleutian Islands and into Southeast Alaska. L. sinclairii occurs from Northern British Columbia to Southern California. The gross distribution of both species appears to be controlled by seawater temperature. Salinity appears to have little influence on distribution, as both species occur throughout wide ranges of salinity and L. sinclairii is able to withstand great fluctuations in salinity at a single site.
Seasonal cycles of growth and reproduction in L. sinclairii were studied on three beaches in Northern Oregon where the plants are subjected to heavy surf. The greatest growth occurs in early summer. As summer advances, the plants are gradually buried under sand until only the ends of the blades protrude. Growth declines during this period and is very slow 102 in late summer. The first heavy storms in fall remove the sand and expose the plants. In November and early December the plants bear ripe sori. Later in December the blades are lost, leaving only the bare stipes. In January new blades are regenerated and when they are only 2-3 cm in length new sori are produced. Sori are produced into March and thereafter only vegetative growth occurs. In March and April there is considerable production of new stipes and blades from the haptera at the margins of the holdfast.
The distribution of L. sinclairii within its temperature range is primarily controlled by the presence of sand. It is adapted to this harsh environment by its growth and reproduction. It possesses great powers of regeneration and is potentially meristematic in almost any region. Although it regularly produces sori, there is little evidence from either field or laboratory studies to indicate that the gametophytes which develop from the spores in these sori normally produce sporophytes. Sexual reproduction of this type is difficult because of the scouring action of the sand. The normal method of reproduction is apparently vegetative proliferation from the haptera at the margins of the holdfasts.
L. longipes was observed in the field in Alaska on only three occasions in summer and twice in winter. Growth is greatest during the summer months. The plants bear ripe sori in December. Laboratory cultures indicate that sexual reproduction is very rare in this species. The blades are reduced in winter but are not completely lost. L_. longipes is 103
not normally associated with sand. Transplant experiments showed that each of the two species can survive for a time in the habitat of the other but cannot survive a whole year. Transplants to sites in British Columbia where the temperature range is greater than in Oregon or Alaska showed that L.. sinclairii is more adversely affected by low winter temperatures, whereas L. longipes is more adversely affected by high summer temperatures. Transplanting to higher or lower temperatures did not affect the production of mucilage ducts.
Comparison of the two species shows they differ in several points besides mucilage ducts, including length of stipes, width of blades, winter loss of blades, morphology of gametophytes and habitat. The evidence confirms that they should be retained as two separate species. 104
X. BIBLIOGRAPHY
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Anderson, CL. 1891. List of California marine algae, with notes. zoe 2:217-225.
Anon. 1965a. Ambient seawater temperature and salinity. Biologist's and Engineer's Daily Report. Vancouver Public Aquarium.
1965b. Meteorological Observations in Canada. Monthly Record. Can. Dept. Trans. Meteor. Br.
1965c. Tide Tables. High and Low water predictions. West Coast, North and South America including the Hawaiian Islands. U.S.C. & G.S.
1966a. Ambient seawater temperature and salinity. Biologist's and Engineer's Daily Report. Vancouver public Aquarium.
1966b. Climatological Data. Annual Summary 1965. U.S. Dept. Comm.
. 1966c. Meteorological Observations in Canada. Monthly Record. 1966. Can. Dept. Trans. Meteor. Br.
1966d. Tide Tables. High and Low Water predictions. West Coast, North and South America including the Hawaiian Islands. U.S.C. & G.S.
1967a. Ambient seawater temperature and salinity. Biologist's and Engineer's Daily Report. Vancouver Public Aquarium.
1967b. Climatological Data. Annual Summary. 1966. U.S. Dept. Comm.
1967c. Meteorological Observations in Canada. Monthly Record. 1967. Can. Dept. Trans. Meteor. Br.
1967d. Surface Water Temperature and Density, pacific Coast, North and South America and pacific Ocean Islands. U.S.C. & G.S. Publ. 31-3 (2nd Ed.) 85 pp. 105
Anon. 1967e. Tide Tables. High and Low Water Predictions. West Coast, North and South America including the Hawaiian islands. U.S.C. & G.S. 1968a. Climatological Data. Annual Summary. 1967. U.S. Dept. Comm. 1968b. Tide Tables. High and Low Water predictions, West Coast, North and South America including the Hawaiian islands. U.S.C. & G.S. Areschoug, J.E. 1883. Observationes phycologicae. pt 4. De Laminariaceis nonnullis. Nova Acta R. Soc. Sci. Upsal. ser. 3, vol. 12.
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Bory de Saint Vincent, J.B. 1826. In Dictionnaire classique d'histoire naturelle. Paris Vol. 9, p. 189. Burrows, Elsie M. 1964. An experimental assessment of some of the characters used for specific delimitation in the genus Laminaria. j. Mar. Biol. Ass. U.K. 44:137-143.
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Dawson, E.Y. 1958a. Notes on Pacific Coast Marine Algae VII. S. Calif. Acad. Sci. Bull. 57(2):65-80.
1958b. A primary report on the benthic marine flora of Southern California. 169-218. In: Oceanographic Survey of the Continental Shelf Area of Southern California. State (Calif.) water Pollution Control Board. Multilith, 560 pp.
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1959. Third Annual Report Benthic Marine Vegetation. 125-169. In : Oceanographic Survey of the Continental Shelf Area of Southern California (2). State (Calif.) Water Pollution Control Board. Multilith. 169 pp. 106 Dawson, E.Y. 1961. A guide to the literature and distributions of Pacific benthic algae from Alaska to the Galapagos Islands. pacif. Sci. 15:370-461.
1966. Marine Botany. An Introduction. Holt. Rinehart and Winston, Inc. San Francisco xii + 371 pp. De Toni, J. 1895. Sylloge algarum omnium hucusque cognitarium. Vol. 3. Fucoideae. padua. xvi + 638 pp.
Doty, M.S. 1947. The marine algae of Oregon. Pt. I. Chlorophyta and phaeophyta. Farlowia 3(1): 1-65. Druehl, L.D. 1965. On the taxonomy, distribution, and ecology of the brown algal genus Laminaria in the Northeast Pacific. Univ. British Columbia Ph.D. Thesis xv + 133 pp.
1968. Taxonomy and distribution of northeast Pacific species of Laminaria. Can. J. Bot. 46:539-547, pis. I-VII. Eber, L., Saur, J. and Sette, 0. 1968. Monthly mean charts, sea surface temperature. North pacific Ocean. U.S. Dept. Interior. Bur. Comm. Fish. Circ. 258.
Farlow, W.G., Anderson, CL. and Eaton, D.C. 1878. Algae exsiccatae Americae Borealis, Fasc. 3, p 118.
1 F0yn, B. 1934. Lebenszyklus, Cytologie, und Sexualitat der Chlorophycee Cladophora suhriana Kiitzing. Arch, protistenk. 83:1-56.
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Hollister, H.J. 1966. Seawater temperature and salinity observations at British Columbia coastal stations in 1965. Fish. Res. Bd. Can. Man. Rept. Ser. 215. Howe, M.A. 1893. A month on the shores of Monterey Bay. Erythea 1:63-68.
Kjeldsen, c.K. 1967. Effects of variations in salinity and temperature on some estuarine macro-algae. Ore. St. Univ. Ph.D. Thesis. 153 pp.
Kjellman, F.R. 1889. Om Beringhafvets algflora. K. svenska Vetensk.-Akad. Handl. 23 (8). 58 pp. 7 pis. 107
Kujala, No and Wyatt, B. 1961. Surface temperature and salinity observations at shore stations on the Oregon Coast. Ore. St. Univ. Dept. Ocean. ONR Data Rept 6. Kiitzing, F.T. 1849. Species algarum .... Leipzig, vi + 922
Le Jolis, A. 1855. Examen des especes confondues sous le nom de Laminaria digitata Auct., suivi de quelques observations sur le genre Laminaria. Mem. Soc. Sci. Nat. Cherbourg 3:241-312. Markham, J.C. 1967. Sand heights of beach at Arch Cape, Oregon (Unpubl. MS.) Markham, J.W. 1968. Studies on the haptera of Laminaria sinclairii (Harvey) Farlow, Anderson et Eaton. Syesis 1:125-131.
Mathieson, A.C. 1967. Morphology and life history of Phaestrophion irregulare S. & G. Nova Hedwigia 13(3/4): 293-318, pis 64-78. Miyabe, K. and Nagai, M. 1932. On Hedophyllum bongardianum (Post, et Rupr.) Yendo and five species of Laminaria from the North Kuriles. Trans. Sapporo Nat. Hist. Soc. 12:194-205. 1933. Laminariaceae of the Kurile islands. Trans. Sapporo Nat. Hist. Soc., 13 (2):85-102. Mowat, J.A. 1965. The occurrence of auxins and gibberellins in algae. Bot. Mar. 8:149-155. Myers, Margret E. 1925. Contributions toward a knowledge of the life-histories of the Melanophyceae. A preliminary
report. Univ. Cal. Publ. Bot. 13(4):109-124. pis 8-10.
Nagai, M. 1940. Marine algae of the Kurile Islands. I. J. Fac. Agric. Hokkaido Univ. 46(1):1-137, pis I-III. 1941. Marine algae of the Kurile Islands. II. J. Fac. Agri. Hokkaido Univ. 46(2):139-310, pis. IV-VII. Okada, Y. 1934. Kaiso-zuhu (Illustrations of Marine Algae) (Jap. Explan.) (Not seen). Okamura, K. 1915. Icones of Japanese Algae III. [3] + 3 pis + 218 pp + [8] .
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Okamura, K. 1933. On the algae from Alaska collected by Y. Kobayashi. Rec. Oceanogr. Wks. japan 5(l):85-97, 2 pis. 1936. Nihon Kaisosi (Marine Algae of japan) (in Japanese). Oliphant, M., Wyatt, B. and Kujala, N. 1962. Surface temperature and salinity observations at shore stations on the Oregon Coast for 1961. Ore. St. Univ. Dept. Ocean. ONR Data Rept. 8. Postels, A. and Ruprecht, F. 1840. Illustrationes algarum . . . . iv + 22 pp + [2], 40 pis. St. Petersburg.
Provasoli, L., McLaughlin, J.J.A., and Droop, M.R. 1957. The development of artificial culture media for marine algae. Arch. Mikrobiol. 25:392-428. Robinson,, G.G.C. 1967. Cytological investigations of the genus Alaria Greville, as it occurs on the West coast of North America. Ph.D. Thesis, Univ. British Columbia, xi + 136 pp.
Ruprecht, F.J. 1851. Tange des ochotskischen Meeres. In Middendorff, A.T. von, Reise in den alissersten Norden und Osten Sibiriens wahrend der Jahre 1843 und 1844. Botanik 1(2): 191-435, pis. 9-18. St. Petersburg. Sanborn and Doty. 1947. The marine algae of the Coos Bay-Cape Arago Region of Oregon. Ore. St. Monogr. Stud. Bot. No. 8, 66 pp, 4 pi, 1 map.
Scagel, R.F. 1957. An annotated list of the marine algae of British Columbia and northern Washington. Nat. Mus. can. Bull. 150.vi + 289 pp. Setchell, W.A. 1896. Notes on kelps. Erythea 4:41-48, pl.l.
1899. Algae of the Pribilof Islands. In D. S. Jordan, Fur Seals and Fur Seal Islands of the North Pacific Ocean, vol 3:589-596, Washington. 1905. Regeneration among kelps. Univ. Cal. Publ. Bot. 2:139-169, pis 15-17. 1912. The kelps of the United States and Alaska. Appendix K. In Cameron, F.K. A preliminary report on the fertilizer resources of the United States. Appendix K. U.S. Senate Document No. 190, pp. 130-178. Setchell, W.A. and Gardner, N.L. 1903. Algae of Northwestern
America. Univ. Cal. publ. Bot. 1(3):165-418. pis 17-27. 109
Setchell, W.A. and Gardner, N.L. 1925. The marine algae of the pacific Coast of North America. Pt. III. Melanophyceae. Univ. Cal. publ. Bot. 8(3):383-898. Shchapova, T.F. 1948. Geografiicheskoye rasprostaneniye predstaviteley poryadka Laminariales v severnoy Chastitikhogo Okeana (Geographical distribution of representatives of the Order of Laminariales in the Northern part of the pacific Ocean.)(In Russian) Trudy Inst. Okeanol. 2:89-138. Shepard, F.P. 1963. Submarine Geology (2nd Ed.) xviii + 557 pp. 1 map. Harper and Row, New York.
Silva, p.c. 1957. Notes on pacific marine algae. Madrono 14(2):41-51. Smith, G.M. 1944. Marine Algae of the Monterey peninsula, California. Stanford Univ. press, ix + 622 pp. Still, R., Wyatt, B. and Kujala, N. 1963. Surface temperature and salinity observations at shore stations on the Oregon Coast for 1962. Ore. St. Univ. Dept. Ocean. ONR Data Rept. 11. Sundene, O. 1963. Reproduction and ecology of Chorda tomentosa. Nytt Mag. Bot. 10:159-167. 1964. The ecology of Laminaria digitata in Norway in view of transplant experiments. Nytt Mag. Bot. 11:83-107. Tatewaki, M. 1931. The primary survey of the vegetation of the Middle Kuriles. J. Fac. Agric. Hokkaido Univ.
29(4):127-190, pis I-X. Tokida, J. 1954. The marine algae of southern Saghalien. Mem. Fac. Fish. Hokkaido Univ. 2(l):l-264, pis. I-XV. Van Overbeek, J. 1940a. Auxin in marine algae. Pi. Physiol. 15:291-299. 1940b. Auxin in marine plants. II. Bot. Gaz. 101:940-947. Vozzhinskaya, V.B. 1964. The bottom flora of Sakhalin (In Russian) Trudy Inst. Okeanol. 69:33 0-440.
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Wyatt, B. and Gilbert/ W. 1967. Surface temperature and salinity observations at pacific Northwest shore stations for 1965 and 1966. Ore. St. Univ. Dept. Ocean. ONR Data Rept. 25. Wyatt, B., Still R., and Haag, C. 1965. Surface temperature and salinity observations at pacific Northwest shore stations for 1963 and 1964. Ore. St. Univ. Dept. Ocean. ONR Data Rept. 21. Yamada, Y. 1935. The marine algae from Urup, the middle Kuriles, especially from the vicinity of Iema Bay. Sci. pap. Inst. Algol. Res. Hokkaido Univ. l(l):l-26, pis. I-X. 1938. Observations on Arthrothamnus bifidus J. Ag. Sci. Pap. Inst. Algol. Res. Hokkaido Univ. 2:113-118. Yendo, K. 1909. Notes on algae new to Japan. Bot. Mag., Tokyo. 23(270):117-133.
1910. Kaisan Shokubutsu-gaku. (Marine Botany) (Not seen) Zinova, A.D. 1959. (List of marine algae of Southern Sakhalin and the Southern Kurile Islands.)(In Russian) Issled. Dal'Nevost. Morea SSSR. 6:146-161. Zinova, E.S. 1933. Les algues de Kamtschatka (in Russian with French summary). Inst. Hydrol. Explor. d. Mers d'URSS. Fasc. 17:7-42. 1954. (Marine algae of eastern Kamschatka). (in Russian) Komarovskie Chteniya Bot. Inst. Akad. Nauk. SSSR. 2(9):365-400. Ill
XI. TABLES I - XXIII. Table I Seawater Temperature and salinity over Distribution ranges of L_. sinclairii and L. longipes.
LOCATION YEARS MEAN FEB. MEAN AUG„ YEARLY MEAN SOURCE COVERED (or coldest (or warmest month) month)
T°C S%0 T°C S% T°C S%, [~~ Urup Island 1949-1962 0o5 11.6 5.1 Eber, Saur (Mar.) & Sette, 1968 Attu Island: Murder Point 1959 1.6 31. 9 11. 3 31. 0 5. 4 31o8 Anon. 1967d Pyramid Cove 1946- 1964 2.1 32. 9 9. 3 31. 5 5. 1 32.1 Anon. 1967d *Sitka 1943- 1964 4.4 30o 4 14. 1 25. 5 8. 5 27.7 Anon. 1967d
Coronation Island In Dec. 1966, 1 C° below Sitka, 0.5%0 below Sitka. Hope Island, B.C» Hollister (Pine Island 1940-1965 7.2 31.3 10.0 31.8 8.6 31.7 records) 1966 1957 8.0 30o7 15.0 30.5 Widdowson River Jordan 1965
1957 8.0 30.7 15.0 30.5 Widdowson Whiffen Spit 1965 1965-1967 7.1 27.9 12.1 26.8 9.4 27.6 Anon. 1965a, •Stanley park 1966a,1967a La push
7.4 30o3 11.6 32.3 9.8 31.2 Anon. 1967c (Neah Bay records) 1936-1964
Indian Beach (Mar.) (Jul.) Gilbert & (Seaside records . . 7.6 27.7 13.9 32.3 10.5 ?30.9? Wyatt, 1968 converted for T°) 1966"1967 Table I Continued i—i H Arch Cape 1960-1963 9.7 31.4 13.2 32.2 11.3 30.9 Kujala, (Mar.) (Jul.) Wyatt, et al, 1961-1965 Short Sand Beach (Mar.) (Jul.) (Seaside records 1966-1967 7.1 21.5 13.4 33.5 10.0 Gilbert & converted for T°) Wyatt, 1968
pescadero Point 1920-1964 11.9 33.2 14.2 33.7 12.9 33.5 . Anon. 1967d (pacific Grove (Sep.) records)
^| port Hueneme 1920-1963 13.2 33„3 16.8 33.6 14.8 33.6 Anon. 1967d **Santa Monica 1946-1964 13o6 33.5 19.7 33.7 16.2 33.7 Anon. 1967d
* Transplant station only. No L. sinclairii or L. longipes present ** Located just south of southern limites of distribution of L. sinclairii (Fig.l). 114
Table II Distribution of Laminaria sinclairii
SITE POSITION SOURCE ACC. NO. British Columbia * Plover is., Hope Island 50°56 'N, 127°58'W UBC 411 Box Island, W. Coast Vancouver Is. 49°04 'N, 125°47'W UBC 17012 **River Jordan, W.C. Vancouver Is. 48°25 .4'N, 124°04'W JWM+ **Whiffen Spit, W.C. Vancouver Is. 48°21 •N, 123°43'W JWM
Washington Salmon Bank, San Juan Is. 48°26'N, 123°01'W Druehi, San Juan Co. 1965 Cape Flattery, Clallam Co. 48°23'N, 124°43.5'W UC++ partridge Bank, San Juan Co. 48 16'N, 122 51'W Druehl, 1965 Agate Beach, Clallam Co. 48 °10«N, 124°43.8'W UBC 17023 **La push, Clallam Co. 47 53.9'N, 124 37.6' W UBC 24574
Oregon **Indian Point, Indian Beach, Clatsop Co. 45°55.9'N, 123°58.8'W UBC 24673 **Bald Point, Indian Beach, Clatsop Co. 45^55.6'N, 123 58.6 'W JWM **Ecola point, Clatsop Co. 45 55.2'N, 123°58.5 'W JWM **Arch Cape, Clatsop Co. 45°48.2'N, 12.3°58.0'W JWM **Cape Falcon, Tillamook Co. 45 46.4'N, 123 58.5'W UBC 30535 **Short Sand Beach, N. end, Tillamook Co. 45°45.8'N, 123°58.2'W JWM **Short Sand Beach, S. end, Tillamook Co. 45";45.5,N, 123 58'W UBC 24933 Cape Kiawanda, Tillamook Co. 45 13.2'N, 123°9.6'W UC Yaquina Bay Mouth, Lincoln Co. 44°36.8'N, 124°4.1'W Kjeldsen, 1967 Coos Bay-Cape Arago Region, Coos Co. 43°20'N, 124°23'W Sanborn (Bassendorf Beach, & Doty, Lighthouse Beach, Squaw 1947 Island, North Bay, Coos Bay, cape Arago)
California Trinidad, Humboldt Co. 41°03.8«N, 124 09'W UC Shell Beach, Sonoma Co. approx. 38 30'N, 123 25'W UC Second Sled Road, Dillon Beach, Marin Co. 38°15.3'N, 122°25.2'W UBC Duxbury Reef, Marin Co. 37°53.3'N, 122°42'W UC San Francisco, San Francisco Co. 37°47'N, 122°30.8'W UC **Pescadero Point, San Mateo Co. 37°14.5'N, 122°25.2'W UBC 24464 115
Table II - Continued
SITE POSITION SOURCE
approx. Cruz Co. 37°10' N, 122°20*W UC Santa Cruz, Santa Cruz Co. 36°57' N, 122°01.8' W UC Asilomar point, Monterey Co. 36°37. 5' N, 121°56. 5'w Smith,1944 Point Lobos, Monterey Co. 36°31. 2' N, 121°57. 21 w Smith,1944 Lucia, Monterey Co. 36°01. 31 N, 121°331 W UC piedras Blancas Point, San Luis Obispo Co. 35°40' N, 121u17'W UC Estero Bay, San Luis Obispo Co. 35°27« N, 120°57 «W UC Morro Bay, San Luis Obispo Co. 35°22' N, 120°51.3'W UC Pismo Beach, San Luis Obispo Co. 35°08. 9 N, 120°38. 8 w UC point Sal, Santa Barbara Co. 34°54. 2 N, 120°40. 4' vr UC Point pedernales, Santa Barbara Co. 34°36. 1 N, 120°38. 5' w UC Gaviota, Santa Barbara Co. 34°28. 3 N, 120°12. 3 'w UC Point Conception, Santa Barbara Co. 34°26. 8 N, 120°28. 2 'w UC Carpinteria Beach State Park, Dawson, Ventura Co. 34°23. 1 N, 119°30' W 19 58a Two Mi. N.W. of Ventura, Dawson Ventura Co. 34°22. 5 N, 119°28' W 1958a Mussel Shoals, Ventura Co. 34°19'N , 119°23.3'w Dawson 1958a
* UBC = Specimen is in Phycological Herbarium, U.B.C. ** Seen by author at this site + JWM = Seen by author at this site but no collections made. ++ UC = Specimen is in Herbarium, Univ. of California, Berkeley. 11.6 Table III Distribution of Laminaria longipes
ACC. SITE POSITION SOURCE NO. UoS.S.R. Kurile Islands* Urup island (Uruppu) 46°00'N, 150°00'E Okamura, 1928; Nagai, 1940 Shimushir is. (Simusiru) 47°00»N, 152°uO'E Okamura, 1928 Nagai, 1940 Ketoy Is. (Ketoi) 47°20»N, 152°28'E Nagai, 1940 Yankicha Isls. (Usisiru) 47°31'N, 152°49'E Nagai, 1940 Matua is. (Matuwa) 48°05«N, 154°31'E Nagai, 1940 Lovushki Is. (Musisiru) 48°32'N, 153°51'E Nagai, 1940 Kharimkotan Is. (Harumkotan) 49 °07 'N/ 154°31'E Nagai, 1940 Onekotan Is. (Onnekotan) 49°25'N, 154°45 «E Nagai, 1940 paramushir Is. (paramusiru) 50°25'N, 155°50'E~ Miyabe & Shumushu Is. (Simusiyu) 50°45'N, 156°20'E -Nagai, 1932; Atlasova Is. (Alaid, Araid) 50°53'N, 157°27*E Nagai, 1940 S. Sakhalin 48°00«N, 143°00'E Miyabe & Nagai,1932; Vozzhinskaya, 1964 Bering Island 55 00 'N, 166 15'E Miyabe & Nagai Kamschatka 1932 Avachinskaya Bay 52°56 'N, 158°36'E Zinova, 1933 Kuimska Bay p p Zinova, 1933 Morzhovaya Bay 53°16 'N, 159°57"E Zinova, 1954 Kronotsky Bay 54°12 •N, 160°36'E Zinova, 19 54 pankara 58°37 'N, 162°24'E Okamura, 1928 Drankinsky p p Okamura, 1928 Barankorfa p p Okamura, 1928 Zavodsk Cape p p Zinova, 1954 Alaska St. Paul Island 57°10 'N, 170°15'W Setchell & / Gardner, 1903 Aleutian Islands Murder point, Attu Is. 52°48 •N, 173°09'E UBC+ 8145 Casco Bay, Attu Is. 52°48 'N, 173°10'E UBC 8309 Chichagof Pt, Attu Is. 52°57 'N, 173°15'E UBC 8381 Agattu is. 52°55 'N, 173°10'E Setchell & Gardner,1903 Kiska Is. 52°00 *N, 177°30'E Setchell & Gardner,1903 Trapper's Cove, Adak Is. 51°48 'N, 176°50'W UBC 9631 North Is, Bay of Islands Adak Is. 51°50 •N, 176°48'W UBC 8391 Cape Agagdak, Adak Is. 52°00 'N, 176°37"W UBC 8150 Zeto Point, Adak Is. 51°55 'N, 176°34'W UBC 8361 Bugle Point, Great 52°02 .3' N, Sitkin Is. 175°58.8'W UBC 13593 **Ram Pt. Beach, Chernofski 53°24 .6' N. Hbr, Unalaska is. 167 31.6'W UBC 27796 117
Table III - Continued
SITE POSITION SOURCE ACC.NO. **Ram Pt., Chernofski Hbr, Unalaska Is. 53°25 'N, 167 31 .5'W UBC 27797 **Cape Aiak-Lance Pt., Unalaska Is. 53°19..1 'N , 167o 25. 9 •w UBC 27724 **Staraya Bay, Unalaska Is. 53°37..4 -N, 165° 30. 6 W UBC 27793 Cape Sarichef I, Unimak Is. 54 35 'N, 164 56 •w UBC 8338 Cape Sarichef II, Unimak Is. 54°35 'N, 164^56 •w UBC 8214 **Raven Point, Cape Sarichef, Unimak Is. 54°38 'N, 164^51 'W UBC 27790 **E. Anchor Cove, Ikatan Pen* Unimak Is. 54°41..6 'N , 163° 03. 2 W UBC 27736 Gulf of Alaska **Eagle Rock, N.E. Harbor Sanak Is. 54o26 .6'N , 162o 35. 4 •w UBC 13620 **Nagai Island 55°12..6 'N , 159° 55. 2 'W UBC 27715 **paul is. 55°48 .7 'N , 159o 21 W UBC 26878 **Chignik Bay, Nakchamik Is. 56°21 .3 • N, 157o 48. 2 •w UBC 27836 **Aghiyuk Is., Semidi isls 56°13..7 'N , 156o 47 W UBC 27831 **Chirikof Is. 55 49..5 'N , 155° 33. 5 •w UBC 27672 **Gurney Bay, Cape Ikolik Kodiak Is. 56°T7.. 7 'N , 154° 44. 9 •w UBC 27071 pasagshak point, Kodiak Is.5 7 25 'N, 152 29 'W UBC 8162 **Cape Chiniak, Kodiak Is. 57°37..3 -N, 152o 09. 5 •w UBC 8161 **Chiniak Is. 57°36..6 " N, 152° 09.. 6 •w UBC 27682 **Peril Cape, Afognak Is. 58°07..5 -N, 152° 16 W UBC 25644 **Kayak Is. 59°51 »N, 144 33 •w UBC 20935 **Wingham Is. 60 03 'N, 144°24 •w UBC 22765 Southeast Alaska **Cape Spencer 58°14 'N, 136 35 •w UBC 20865 **Cape Ommaney, Baranof Is. 56°10 'N, 134 40 •w UBC 20384 **Aats Bay, Coronation Is. 55"53..7 'N , 134o 16 W UBC 20336 **Helm point. Coronation Is. 55 49..6 'N , 134° 17 W UBC 20319 Washington ++Salmon Bank, San Juan Is. 48° 2 6 'N, 123 01 •w Druehi, 1968
* Ceded by japan to U.S.S.R. in 1945; names in parentheses are former Japanese names. ** L. longipes seen by author at this site + UBC = Specimen is in Phycological Herbarium, U.B.C. ++ Subtidal Only 118
Table IV Analysis of Sand Grain Size on Oregon Beaches
INDIAN BEACH ARCH CAPE SHORT SAND BEACH SIZE % % %
2 nun + 0.01 0.04 0.01 Very coarse
1 - 2 mm 0.06 0.15 0.02 Coarse
0.5 - 1 mm 0.61 2.14 0.27 Medium
0.25 - 0.5 mm 67.85 74.99 75.22 Fine
0.1 - 0.25 mm 31.41 22.64 24.43 Very fine
0.05 - 0.1 mm 0.05 0.01 0.04
Less than 0.05 mm 0.01 0.03 0.01
100.00% 100.00% 100.00%
Total sieved: 2575.5 g 2427.2 g 2415.0 g 119
Table V. Temperature and precipitation at Alaska stations
1965 1966 1967 30-Year Mean
T°C T°C Ppt T°C Ppt T°C Ppt. (xn.) Ltka (Sitka Magnetic) (Anon. , 1966b, 1967b, 1968a) Jan. -1.8 12.81 -3.9 5.61 -0.8 7.91 3.2 7.77 Feb. 0.1 7.31 1.2 6.99 1.3 8.97 4.7 6.38 Mar. 2.4 2.81 2.1 7.68 -0.7 3.44 5.9 6.95 Apr. 4.6 5.42 4.1 5.49 4.2 2.87 9.0 5.35 May 5.8 6.55 6.4 9.78 8.1 4.46 12.2 4.66 Jun. 8.9 8.60 10.8 1.11 10.9 2.60 14.9 3.46 Jul. 12.3 2.42 12.7 4.59 12.1 4.73 16.3 5.20 Aug. 12.4 6.90 12.1 8.63 13.9 7.43 16.7 7.86 Sep. 11.0 6.02 10.8 11.13 10.9 16.03 14.9 11.49 Oct. 7.1 18.66 5.0 20.49 7.2 14.07 10.8 15.27 Nov. 1.8 5.05 0.4 8.72 2.5 12.16 6.9 12.01 Dec. -0.3 11.51 0.9 4.96 0.7 6.26 3.8 10.17
m. 5.3 93.88 5.2 95.18 5.9 90.93 9.9 96.57
Coronation Island (Cape Decision)(Anon., 1966b, 1967b, 1968a ) Jan. 0.5 12.09 -1.5 5.66 1.1 6.84 0.9 6.19 Feb. 2.2 8.64 2.8 7.23 3.1 9.55 1.9 5.54 Mar. 3.9 1.66 3.5 10.48 1.0 1.08 2.6 5.35 Apr. 6.0 6.07 4.9 3.03 5.2 1.77 4.8 4.85 May 6.1 5.30 6.4 7.65 8.7 4.07 7.3 4.20 Jun. 9.4 3.83 10.3 1.48 9.5 2.19 9.7 2.73 Jul. 11.1 2.58 11.6 2.67 11.2 3.94 11.4 3.64 Aug. 11.6 1.53 11.4 4.42 13.6 6.07 11.7 5.05 Sep. 10.3 2.63 10.5 10.65 11.8 10.65 10.4 7.56 Oct. 8.6 18.43 6.4 11.17 8.4 11.30 7.9 12.25 Nov. 4.4 4.22 2.7 3.86 5.2 10.97 4.7 9.68 Dec. 1.8 8.81 2.5 7.39 2.8 9.53 2.9 9.08
Ann. 6.3 75.77 6.0 75.69 6.8 77.96 6.3 76.12 120
Table V - Continued
1965 1966 1967 15-Year Mean T°C Ppt. T°C Ppt. T°C Ppt. T°C Ppt. Attu Island (Anon., 1966b, 1967b, 1968a)
Jan. -0.7 5.47 -0.8 5.14 -1.2 6.71 -0.6 4.18 Feb. 0.4 15.45 -2.1 4.90 -1.7 5.87 -0.7 4.14 Mar. 0.5 17.80 -0.5 2.62 0.3 0.65 -0.1 3.86 Apr. 2.7 16.60 0.9 3.48 1.4 0.02 1.7 4.68 May 4.8 2.59 3.8 7.81 4.9 0.01 4.0 3.85 Jun. 6.7 1.26 5.8 2.18 7.2 0.04 6.6 3.19 Jul. 9.1 5.36 8.6 2.90 9.0 1.20 9.0 4.75 Aug. 10.4 3.96 10.6 4.39 10.9 3„74 10.5 5.64 Sep. 8.5 3.93 9.7 11.52 8.9 5.87 8.9 6.04 Oct. 5.0 6.72 5.4 5.02 5.8 7.81 5.3 6.68 Nov. 1.9 3.93 3.4 9.50 1.9 1.95 2.1 4.79 Dec. 0.4 5.33 0.4 3„75 0.4 0.78 0.7 4.35
Ann. 4.3 88.40 3.8 63.21 4.0 34.65 4.0 56.15
Table VI Mean Temperature on Urup Island (Tatewaki, 1931)
T°C Jan. -1 to -8 Feb. -3 to -8.5 Mar. -4 to -8 Apr. 1 to 2 May 2.5 to 4 Jun. 5 to 10 Jul. 11 Aug. 10 to 15 Sep. 9.5 to 11 Oct. 2 to 3 Nov. No Data Dec. 0 to -1.5
Ann. 2.5 to 4.0 121
Table VII Temperature and precipitation at
British Columbia Stations
Bull Harbour (for Hope Island) Long term Mean T° Ppt Jan. 3.8 7.64 Feb. 4.4 5.71 Mar. 5.9 5.96 Apr. 7.6 4.27 May 10.0 3.06 jun. 12.0 1.95 Jul. 13.5 2.59 Aug. 13.7 2.94 Sep. 12.2 4.73 Oct. 9.7 9.12 Nov. 6.6 9.72 Dec. 4.9 10.37 ANNUAL 8.7 68.06 Sooke (E. Sooke - Anderson Cove) (Anon., 1965b, 1966c, 1967c)
1965 1966 1967 Long Term Mean T°c «?«fc\ T°c pPfc T°c pPfc T°c pPfc- Jan. * * * * 5.0 13.88 * * Feb. * * * * 5.7 8.18 * * Mar. * * * * 5.1 3.76 * * Apr. * * * * 7.0 2.62 * * May * * 10.7 1.02 11.4 1.07 * * Jun. * * 13.2 0.80 15.4 0.25 * * Jul. * * 15.0 1.07 * * * * Aug. * * 15.8 0.55 18.0 0.00 * * Sep. * * 14.7 1.43 15.4 2.91 * * Oct. * * 9.4 6.03 10.7 11.58 * * Nov. * * 6.9 5.98 7.0 4.32 * * Dec. * * 6.1 9.96 4.1 7.46 * * 122
Table VII Continued
River Jordan (Anon., 1965b, 1966c, 1967c) 1965 1966 1967 Long Term Mean T°C Ppt. T°C Ppt. T°C Ppt. T°C Ppt. (in.) Jan. 4.0 12.57 4.9 11.16 * 22.28 4.0 10.37 Feb. 5.3 12.69 5.8 6.63 5.8 17.24 4.1 8.59 Mar. 6.3 1.58 6.0 9.14 5.4 8.58 6.5 6.80 Apr. 8.2 4.06 8.2 2.65 7.0 3.48 9.0 4.52 May 9.6 3.80 9.8 3.01 10.4 1.50 11.2 2.74
Jun. 12.6 0o66 12.5 2.81 14.3 0.29 13.0 1.90
Jul. * 0.84 14.1 1.49 * 0.80 15.0 1015 Aug. * 2.28 14.8 1.99 15.9 0.28 15.1 1.58 Sep. 12.2 3.47 13.8 2.97 * 5.67 14.3 3.47 Oct. * 8.26 * 10.55 * 22.71 11.0 8.50 Nov. 8.5 11.26 7.4 12.55 7.8 8.53 8.4 10.61 Dec. 9.9 10.35 * 18.83 4.1 15.71 5.3 13.19
ANNUAL * 71.85 * 83.98 * 107.07 9.7 73.42
Vancouver (Kitsilano) (Anon. , 1965b, 1966c, 1967c) 1965 1966 1967 Long Term Mean (HMCS Discovery) T C Ppt. T°C Ppt. T°C Ppt. T°C Ppt. (in.) Jan. 2.8 7.35 3.7 8.85 4.8 11.87 5.7 8.59 Feb. 4.6 9.63 4.8 3.51 5.5 5.53 5.9 6.41 Mar. 5.7 2.26 6.9 4.20 5.7 5.72 * * Apr. 9.9 2.45 9.2 1.24 8.0 3.21 * * May 11.6 2.15 12.7 2.43 13.1 2.15 12.6 2.67 Jun. 16.1 0.57 15.2 2.54 18.3 0.55 17.4 2.67 Jul. 18.8 0.35 17.1 2.84 18.8 1.07 * 1.70 Aug. 18.0 3.11 17.8 1.67 20.3 0.23 19.4 1.64 Sep. 13.8 0.58 15.8 2.85 * * 16.4 3.38 Oct. 11.7 8.20 10.0 7.09 * * 11.0 6.84 Nov. 7.8 5.68 7.1 9.45 * * 7.3 8.14 Dec. 3.8 7.78 6.1 14.62 * * 4.2 9.95
10.38 50.11 10.53 61.29 * * * 61.75
* = No Data 123
Table VIII Temperature and Precipitation at Oregon Stations
Seaside (Anon., 1966b, 1967b, 1968a)
~ 1965 „ 1966 1967 Long Term Mean T C T"C Ppt. T C Ppt. T°C Ppt. Un.) Jan. 6.4 19.11 6.8 10.42 7.7 16.85 6.3 11.90 Feb. 7.4 6.42 7.1 6.81 7.4 6.86 7.2 9.84 Mar. 9.1 1.22 7.3 9.82 6.9 8.75 7.9 9.27 Apr. 9.8 4.49 10.4 2.92 7.5 5.48 9.7 5.41 May 10.6 2.91 10.5 1.95 11.3 1.00 11.8 3.31 Jun. 12.8 1.18 14.1 1.64 14.4 1.33 14.0 3.12 Jul. 15.1 0.41 15.2 0.81 16.0 0.23 15.3 1.23 Aug. 16.4 1.85 15.2 0.76 16.4 0.09 15.7 1.53 Sep. 14.6 0.49 15.8 1.98 16.8 2.46 14.9 2.94 Oct. 13.6 3.75 12.2 6.18 13.4 10.55 12.5 7.57 Nov. 10.8 12.53 9.9 9.17 10.3 7.47 9.1 10.40 Dec. 6.1 12.89 8.7 15.63 6.3 11.59 7.4 13.18
Ann. 11.1 67.25 11.1 68.09 11.2 72.66 11.0 79.70
Arch Cape Stations A & B Precipitation only (in.) 1965 1966 1967 A B A B A B Ten-yea(Statiorn MeaA n Jan. 17.99 22.60 8.98 11.14 17.14 21.79 12.71 Feb. 6.34 7.62 6.06 6.68 5.42 6.73 9.77 Mar. 1.68 1.50 9.97 12.37 10.78 13.05 8.82 Apr. 6.41 7.59 2.13 2.71 5.16 5.75 5.60 May 2.41 2.94 2.39 2.69 2.12 2.10 3.29 Jun. 1.54 1.57 1.90 2.15 1.46 1.65 3.11 Jul. 1.26 1.24 1.36 1.46 0.47 0.51 1.26
Aug. 2.37 2062 1.36 1.85 0.20 0.20 2.19 Sep. 1.02 1.17 3.34 3.43 4.16 4.51 3.32 Oct. 3„78 4.49 6.76 8.05 9.00 12.59 7.26 Nov. 11.42 11.83 10.96 9.27 6.16 7.20 12.09 Dec. 14.53 16.76 14.70 20.81 12„49 15.74 11.46
Ann. 70o35 81.93 69.89 82.61 74.54 91.82 81.45 124
Table IX Long Term Temperature and Precipitation at two California Stations.
Santa Cruz, Santa Cruz Co. Oxnard, Ventura Co. (Anon., 1968a) (Anon., 1968a)
T°C Ppt.(in.) T°C Ppt. Jan. 9.4 6.84 11.9 3.33 Feb. 10.4 5.81 12.2 2o99 Mar. 11.7 4.15 12.9 2.27 Apr. 13.0 2.11 14.0 1.13 May 14.7 1.01 15.2 0.13 Jun. 16.9 0.21 16.4 0.05 Jul. 17.2 0.03 18.2 0.00 Aug. 17.2 0.06 18.4 0.03 Sep. 17.4 0.27 18.1 0.08 Oct. 15.4 1.38 16.8 0.40 Nov. 12.6 9.58 14.8 1.14 Dec. 10.3 7.12 12.9 3.20
Ann 13.9 31.25 15.2 14.75 Table X Measured Values of Seawater Temperature and Salinity on Oregon Beaches
MONTHLY MEAN INDIAN BEACH SHORT SAND BEACH SEASIDE
Date S%o T°C Date T°C Date T°C
17 Aug. •66 33.843 13 Sep. •66 32.058 8.0 14 Sep. •66 30.833 7.8 Sep. '66 31.6 14.8 15 Oct. •66 30.898 9.0 16 Oct. •66 - 10.0 Oct. '66 31.8 13.4 11 Nov. •66 30.840 10o0 12 Nov. '66 8.876 10.0 Nov. '66 31.6 11.8
27 Dec. •66 25.210 902 28 Dec. •66 23.974 9.0 Dec. •66 30.1 11.4 24 Jan. '67 29.689 8.5 25 Jan. •67 16.222 8.5 Jan. '67 27.9 10.6 23 Feb. '67 27.735 9.0 24 Feb. '67 21.479 8.5 Feb. '67 26.3 9.9 29 Mar. '67 28.608 7o2 28 Mar. •67 17.797 7.2 Mar. •67 27.4 9.6 24 Apr. '67 29.252 9.0 26 Apr. •67 21.103 9.5 Apr. •67 26.2 11.0 23 May •67 25.821 9.5 22 May •67 24.194 10.0 May •67 27.5 12.8 20 Jun. •67 28.988 13.0 22 Jun. •67 8.521 11.0 Jun. •67 27.9 13.9 19 Jul. •67 23.827 15.7 20 JUlo '67 5.357 14.5 Jul. '67 29.8 15.9 17 Aug. '67 32.295 12.0 18 Aug. •67 33.523 9.0 Aug. •67 29.7 14.5 2 Sep. •67 32.190 14.0 3 Sep. '67 31.115 13.0 Sep. •67 29.0 15.8
* Wyatt and Gilbert, 1967; Gilbert and Wyatt, 1968. 126
Table XI Calculated Mean Monthly Seawater Temperatures * on Oregon Beaches .
INDIAN BEACH SHORT SAND BEACH (Seaside - 2.0 C°) (Seaside - 2.5 C°)
1966 Oct. 11.4^ 10.9 Nov. 9.8 9.3 Dec. 9.4 8.9
1967 Jan. 8.6 8.1 Feb. 7.9 7.4 Mar. 7.6 7.1 Apr. 9.0 8.5 May 10.8 10.3 Jun. 11.9 11.4 Jul. 13.9 13.4 Aug. 12.5 12.0 Sep, 13.8 13.3
Mean: 10.55°C 10.05°C
* Calculated by subtracting mean difference from Seaside monthly means (Wyatt and Gilbert, 1967; Gilbert and Wyatt, 1968) for October 1966 to September 1967. 127
Table XII Mean values of Seawater Temperature and „ * Salinity at Arch Cape for 1960-1963.
MONTH T°C S%» January 9.03 30.84 February 9.70 31.36 March 9.01 31.03 April 10.88 28.66
May 12.36 29.53
June 12.61 30.91
July 13.13 31.62 August 13.22 32o24
September 12.46 32.12
October 12.41 32.00
November 11.15 30.96 December 9.80 30.45
MEAN: 11.31 30.97
* Kujala and Wyatt, 1961; Oliphant, Wyatt
and Kujala, 1962; Still, Wyatt and Kujala,
1963; Wyatt, Still, and Haag, 1965. 128
Table XIII Sand Level at Indian Beach. Location of points shown on Figure 7. Allheight s are
cm down from standard reference point.
SAND by A: SAND by F: DATE ROCK A X Y PT. F E D (Inner) (Outer) (Inner) (Outei
•66:17/8 146 208 183 193 257 213 13/9 146 157 163 193 246 *
15/10 146 246 218 193 320 254 11/11 146 246 218 193 234 *
27/12 146 246 218 193 259 236
'67:24/1 146 246 218 193 259 236
23/2 146 246 218 193 259 236 29/3 146 246 218 193 259 236
25/4 146 246 218 193 259 236 25/5 146 246 218 193 J 191 185
21/6 146 218 * 193 198 *
19/7 146 i 112 112 193 .' 137 137
17/8 146 J 130 130 193 I 137 137
2/9 146 246 218 193 208 *
* := No Data
1 - • - Rock is Buried
Number underlined = under water Table XIV Sand Level at Short Sand Beach. Location of points shown on Figure 8. All heights are
cm down from standard reference point.
BEACH NORTH WATER-LEVEL DOWN SAND SAND OF STREAM FROM TOP OF DATE ROCK A by A ROCK B by B N. of A N. of B ROCK A ROCK •66:18/8 211 269 213 267 * * * * 14/9 211 239 213 252 * * * * 16/10 211 269 213 282 * * 81 * 11/11 211 277 213 279 * * * * 28/12 211 320 213 287 * * * 94 •67:25/1 211 320 213 287 * * * * 24/2 211 320 213 287 * * * * 28/3 211 320 213 287 * * * * 26/4 211 297 213 * * * * 135 24/5 211 320 213 287 * * * * 22/6 211 274 213 259 198 * 64 64 20/7 ^ 211 302 213 287 198 207 33 25 18/8 2ll * 213 * 168 * 18 13 3/9 211 249 213 254 226 * 28 *
* = No Data 130
Table XV Sand Heights of Beach at Arch Cape, Oregon
(Markham, 1967 unpubl.)
Date 31 Jul 5 Sep 2 Oct 2 Nov 27 Nov 31 Dec 26 Jan Distance 66 66 66 66 66 66 67
0+00 50.00 50.00 50.00 50.00 50.00 50.00 50o00 +50 40.00 40.53 40.18 40.30 40.31 39.85 39.60 1+00 37.18 39.72 38.13 38.20 37.98 37.82 37.42 +50 36.03 38.86 36.50 36.66 36.03 36.22 35.72 2+00 35.60 38.16 35.16 34.89 35.31 34.77 34.25 +50 33.56 37.06 33.71 33.34 32.93 33.42 33.14 3+00 31.83 35.98 32.21 31.95 31.63 32.22 31.78 +50 31.10 34.28 31.01 31.39 30.44 31.12 30.78 4+00 29.99 32.72 30.01 30.09 29.41 30.32 29.81 +50 31.70 29.16 28.58 29.57 28.85 5+00 Deep 31.06 28.31 27.80 28.62 28.14 +50 pool 30.48 27.18 27.60 6+00 26.68 26.74 +50 26.28 Edge of cobbles 0+50.4 0+48.4
Date 24 Feb Mar 30 Apr 24 May 1 Jul 6 Aug 67 67 67 67 67 67 Distance 0+00 50.00 50.00 50.00 50.00 50.00 50.00 +50 39.83 39.96 39.93 40.07 40.09 40.10 1+00 37.93 37.26 37.52 37.57 37.63 37.60 +50 35.93 35.39 35.57 35.84 36.49 36.50 2+00 34.37 33.82 3.3.65 34.08 34.58 35.00 +50 33.05 32.29 32.11 32.72 32.84 32.66 3+00 31.87 31.02 30.88 31.35 31.45 30.59 +50 30.77 29.89 29.98 30.26 30.63 29.80 4+00 29.82 28.94 28.42 29.51 29.59 29.23 +50 28.87 28.16 28.62 29.02 29.59 28.40 5+00 28.02 27.59 28.42 28.45 26.87 28.00 +50 27.12 26.84 28.17 29.02 27.27 27.84 6+00 26.27 27.72 27.16 27.07 26.64 +50 27.27 26.12 7+00 28.82 25.67 +50 25.87 8+00 25.07 Edge of Cobbles 0+47.0 0+47.0 0+44 0+40 Notes: Distance 0+00 is a fixed point from which all other points are measured and established at 50-foot intervals; it has arbitrarily been assigned a height of +50 feet and all other heights have been corrected to this reference. Edge of cobbles is the distance west of 0+00 at which cobbles and sand meet. All heights and distances are in feet. Table XVI Associated Plant Species ro H Aats La Indian Arch Cape Short Pesc. Bay Push Beach Cape Falcon Sand Point Heights:S L M H L M H L M H L M H L M H L M H L M H RHODOPHYTA * * * Ahnfeltia concinna J.Ag. X X * X X Ahnfeltia plicata (Huds.) Fries X X X Antithamnion sp. X Bonnemaisonia nootkana (Esp.) Silva X Bossiella corymbifera (Manza) Silva * X
Bossiella dichotoma (Manza) Silva x x X Bossiella plumosa (Manza) Silva * X X X X Bossiella sp. X X X X X Calliarthron sp. x Callithamnion pikeanum Harv. X X X Callophyllis sp. X Ceramium washingtoniense Kyiin X Constantinea simplex Setch X X X Constantinea subulifera Setch. X Corallina officinalis L. x x Corallina Vancouveriensis Yendo X X X X X X X Corallina sp. X X X X coralline (crustose) X X X X X X coralline X X X X Cryptopleura sp. X X X X X X X Cryptosiphonia woodii J. Ag. X Cumagloia andersonii (Farl.) S. & G. X X X X X Dermatolithon dispar (Fosl.) Fosl. X X * X Dilsea californica (J. Ag) 0. Kuntze * X Endocladia muricata (Harv.) J. Ag. X X X X X X X X X X X Erythrophyllum delesseriodes J. Ag. X X X X X X X X Euthora cristata (L.) J. Ag. X X Farlowia mollis (Harv. & Bail.) Farl & Setch. Fauchea sp. X X X X X X Gigartina sp. X X X X X X X X XXX X Con tin ued Aats La Indian Arch Cape Short Bay Push Beach Cape Falcon Sand S L M H L M H L M H L M H L M H L M H Gloiopeltis furcata (p. & R.) J. Ag. x Gloiosiphonia verticillaris Farl. x x Gracilaria verrucosa (Huds.) papenf. Grateloupia sp. * x X X * * X Gymnocfongrus linearis (Turn,) J.Ag. X X Halosaccion glandiforme (Gmel.) Rupr. X X x x Halymenia californica Smith & Hollenb. x X X Hildenbrandia sp. * x X Iridaea sp. X X * X X X X XXX XXX Kallymenia sp. X X X Laurencia spectabalis P. & R. X Lithothamnion sp. X Lithothrix aspergillum J. E. Gray X Melobesia sp. X Membranoptera sp. X X * Microcladia borealis Rupir. X X * XXX Microcladia sp. X X * X X Odonthalia kamtschatika (Rupr.) J. Ag. X X Odonthalia sp. XXX X X X X X Opuntiella californica (Farl.) Kylin X Petrocelis franciscana S & G. X X X X Petrocelis middendorfii (Rupt.) Kjell. X Peyssonelia pacifica Kylin X X Phycodrys sp. X * * * Plocamium oregonum Doty X X X * * X X Plocamium pacificum Kylin * X X Plocamium violaceum Farl. X X Plocamium sp. X X Polyneura latissima (Harv.) Kylin X Polyporolithon sp. X Polysiphonia sp. X X X X X x * Porphyra sp„ X X XXX XXX X X X X X Table Continued oo ro H Aats La Indian Arch Cape Short Bay Push Beach Cape Falcon Sand Heights: S L M H L M H L M H L M H L M H L M H Porphyrella gardneri Smith & Hollenb. x Prionitis lanceolata Harvey Prionitis linearis Kylin x x x prionitis lyallii Harv. X * x Prionitis sp. x x X X X xxx x * xx* x Pterosiphonia bipinnata (p. & R.) Falkenb. X X X X Pterosiphonia sp. x x X X X Ptilota asplenoides (Esp.) c Ag. X X Ptilota filicina (Farl.) J.Ag. Ptilota pectinata (Gunn.) Kjell. Ptilota tenuis Kylin X Ptilota sp. X X X X * * X X Rhodoglossum sp. X X Rhodomela larix (Turn.) C. Ag. X X X X xxx Rhodymenia palmata (L.) Grev. X Rhodymenia sp. X X PHAEOPHYTA Alaria fistulosa P. & R. X Alaria marginata p. & R. X X X * X X Alaria nana Schrader X Alaria tenuifolia Setch. X X Costaria costata (Burn.) Saund. X Cymathere triplicata (p. & R.) J. Ag. X X Cystoseira geminata c. Ag. X Cystoseira osmundaceae (Menz.) c. Ag. Desmarestia munda S. & G. * X X X Desmarestia viridis (Mull.) Lamour. X X Demarestia sp. X X X X Ectocarpus sp. X X Egregia menziesii (Turn.) Aresch. Table XVI Cont d Aats La Indian Arch Cape Short Bay Push Beach Cape Falcon Sand Heights: S L M L M H L M H L M H L M H L M H Fucus sp. x x Haplogloia andersonii (Farl.) Lev. X x x * Hedophyllum sessile (C. Ag.) Setch. X X X X x Heterochordaria abietina (Rupr.) S. & G. X x x Laminaria groenlandica Rosenv. xxx Laminaria longipes Bory X Laminaria setchellii Silva X X X X X Laminaria sinclairii (Harv.) Far., And. & Eat. * * * Leathesia difformis (L.) Aresch. X X X X X Lessoniopsis littoralis (Farl. & Setch.) Reinke X X X X Macrocystis ihtegrifolia Bory Myelophycus intestinale Saund. Nereocystxs luetkeana (Mert.) P. & R. X Pelvetiopsis limitata (Setch.) Gard. X X * X X Phaeostrophion irregulare S. & G. X Pleurophycus gardneri Setch. & Saund. X Postelsia palmaeformis Rupr. Punctaria sp. X Pylaiella sp. X Ralfsi a sp. X X Sargassum muticum (Yendo) Fens. X Scytosiphon lomentaria (Lyng.) J. Ag. X X X Soranthera ulvoidea p. & R. X X X X
CHLOROPHYTA Cladophora sp. * * * * X X X X Codium setchellii Gard. X X Enteromorpha intestinalis (L.) Link X X Enteromorpha linza (L.) J. Ag. X X Enteromorpha sp. X X X X X X X X Table XVI Continued in n rH Aats La Indian Arch Cape Short Pesc. Bay Push Beach Cape Falcon Sand Point L M H L M H L M H i L M H L M H L M H L M H
Rhizoclonium sp. X X X Spongomorpha coalita (Rupr.) Coll. X Spongomorpha sp. X X X Ulothrix sp. X * Ulva lactuca L. X X X Ulva sp. X X XXX X
CHRYSOPHATA * * Colonial diatoms X X X X X
ANTHOPHYTA * * * * X Phyllospadix scouleri Hook. X X X X
S# subtidal; L, lower intertidal; M, mid intertidal; H, high intertidal
* ab^ove = associated with sand. Table XVII Seasonal Distribution of Plant Species at Indian Beach cn Jun 6 6 Apr 6 7 May67 Jun67 Jul67 Aug67 Sep67 L M H L M H L M H L M H L M H L M H L M H H Height:
RHODOPHTYA * * * x x Ahnfeltiaconcinna J. Ag. x x Ahnfeltia plicata (Huds.) Fries x Bonnemaisonia nootkana (Esp.) Silva * X X Bossiella dichotoma (Manza) Siva X X X * X x X Bossiella plumosa (Manza) Silva X X X X x X X Bossiella sp. X Callithamnion pikeanum Harv. X X Constantinea simplex Setch. X X X X * Corallina officinalis L. X Corallina Vancouveriensis Yendo X X Corallina sp. X X coralline (crustose X X X Cryptopleura sp. X X Cryptosiphonia woodii J. Ag. X X X X X X X X X Dilsea californica (J. Ag.) 0 Kuntze X X X X Endocladia muricata (Harv.) J. Ag. X * X Gigartina sp. X X X X * * Gymnogongrus linearis (Turn.) J. Ag. X X Hildenbrandia sp. X X X X X X X X * Iridaea sp. X X Kallymenia sp. X Laurencia spectabalis P. & R. Melobesia sp. * Microcladia borealis Rupr. X X Odonthalia sp. X X Petrocelis franciscana S. & G. X * Peyssonelia pacifica Kylin * X X X X Plocamium - oregonum Doty X * X Plocamium pacificum Kylin X Polocamium violaceum Farl. X X Plocamium sp. Table XVII Cont inued
Jun 6 6 Apr 6 7 May 6 7 Jun67 Jul 6 7 Aug 6 7
H Height: L M H L M H L M H L M H L M H L M H Polysiphonia sp. X X Porphyra sp. X X X X X Porphyrella gardneri Smith & Hollenb. X Prionitis lyallii Harv. * X X X X Prionitis sp. X X X * Pterosiphonia bipinnata (pc & R.) Falkenb. X X X X X X Pterosiphonia sp. X Ptilota tenuis Kylin Ptilota sp. X X X X X X X X Rhodomela larix (Turn.) C. Ag.
PHAEOPHYTA X X X X X Alaria marginata P. & R. X * X Alaria nana Schrad. X X X Desmarestia munda S. & G. X Desmarestia sp. X Haplogloia andersonii (Farl.) Lev. X X X X X X X Hedophyllum sessile (C. Ag.) Setch. X X X X Laminaria setchellii Silva X X X X x X Laminaria sinclairii (Harv.) Farl. And. & Eat. X X X X X X X Lessoniopsis littoralis (Farl. & Setch.) Reink X X X X Buried ... Phaeostrophion irregulare S. & G. X X X Ralfsia sp. CHLOROPHYTA Codium setchellii Gard. * X X X X Buried Enteromorpha linza (L.) J. Ag. X Enteromorpha sp. X X X Ulva sp. Table XVII Continued
Jun66 Apr 6 7 May 6 7 Jun 6 7 JU167 Aug 6 7 Sep67 Height: L M H L M H L M H L M H L M H L M H L M H
CHRYS OP HYTA * * Colonial diatoms X X
ANTHOPHYTA * * * * phyllospadix scouleri Hook. X X X X X X X
* above x = associated with sand. Table XVIII Seasonal Distribution of Plant Species at S hort Sand Beach
Jun. Apr. May Jun. Jul. Aug. 66 67 67 67 67 67 Height: L M H L M H L M H L M H L M H L M H RHODOP HYTA Ahnfeltia concinna j. Ag. * * * * Ahnfeltia plicata (Huds.) Fries x x x x Bossiella corymbifera (Manza) Silva x Bossiella plumosa (Manza) Silva x Bossiella sp. X Callithamnion pikeanum Harv. X Constantinea simplex Setch. X Corallina Vancouveriensis Yendo X X X X X coralline (crustose) X coralline X X X Cryptopleura sp. X X Cryptosiphonia woodii J. Ag. X X X x Cumagloia andersonii (Farl.) S. & G. X Dermatolithon dispar (Fosl.) Fosl. * Dilsea californica (J. Ag.) 0. Kvintze X X X Endocladia muricata (Harv.) J. Ag. X X Ervthrophyllum delesseriodes J. Ag. X X Farlowia mollis (Harv. & Bail0) Farl. & Setch. X X Gigartina sp. X X X X X X x Gloisiphonia verticillaris Farl. X X Grateloupia sp. X
Gymnogonqrus linearis (Turn.) j0 Ag. X * * * * X Halvmenia californica Smith & Hollenb. X X X X Hi1denbrandia sp. X X
Iridaea sp. X X X X X X X X Kallymenia sp. X Microcladia borealis Rupr. X X X * X X X Odonthalia sp. X X Petrocelis franciscana S. & G. X X Plocamium oregonum Doty * X X X Table I Continued o r-i Jun. Apr. May Jun. Jul. Aug. Sep. 66 67 67 67 67 67 67 Height: L M H L M H L M H L M H L M H L M H L M H polyneura latissima (Harv.) Kylin Polysiphonia sp. x Porphyra sp. X X x x porphyrella gardneri Smith & Hollenb. X prionitis lanceolata Harv. X x prionitis linearis Kylin X prionitis lyallii Harv. X x x X Prionitis sp. XXX Pterosiphonia bipinnata (p. & R.) Falkenb. * x X Pterosiphonia sp. X Ptilota asplenoides (Esp.) C. Ag. * * X X ptilota filicina (Earl.) J. Ag. x X Ptilota pectinata (Gunn.) .Kjell. * X Ptilota sp. X X * * X X x X X X X X Rhodomela larix (Turn.) C. Ag. x
PHAEOPHYTA X Alaria marginata p. & R. X X X X * X X X X Alaria sp. X Desmarestia munda S. & G. X Desmarestia sp. X Ectocarpus sp. X X Fucus sp. X X X X X * X X X Hedophyllum sessile (C. Ag.) Setch. X X Laminaria setchellii Silva X * * * * Laminaria sinclairii (Harv.) Farl. And. & Eat. X X X X Leathesia difformis (L.) Aresch. X X Lessoniopsis littoralis (Farl. & Setch.) Reinke X Pelvetiopsis limitata (Setch.) Gard. X Table XVIII Continued
Jun. Apr. May Jun. Jul. Aug. Sep. 66 67 67 67 67 67 67 Height: L M H L M H L M H L M H L M H L M H L M H
Phaeostrophion irregulare S. & G. X pilayella sp. X Soranthera ulvoidea P. & R. X X
CHLOROPHYTA * * Cladophora sp. X X X X X Enteromorpha intestinalis (L.) Link X X X X Enteromorpha sp. X X X Rhizoclonium sp. X Spongomorpha coalita (Rupr.) Coll. X Ulva lactuca L. X Ulva sp. X X X X X X
CHRYSOPHYTA Colonial diatoms X X
ANTHOPHYTA * * * * * * Phyllospadix scouleri Hook. X X X X X X
* above x = associated with sand Table XIX Seasonal Distribution of Algal Spe es at Aats Bay, Coronation I sland
Jun. 67 Dec. 65 Jun. 66 Dec. 66 Height: S L M H S L M H S L M H S L M H RHODOPHYTA Antithamnion sp. x Bossiella sp. x X X Calliarthron sp. x Callophyllis sp. X Constantinea subulifera Setch. X Corallina Vancouveriensis Yendo X Corallina sp. X X coralline (crustose) X X X X Cryptopleura sp. X Cryptosiphonia woodii J. Ag. X X Dilsea californica (jTAg.) 0. Kuntze X Endocladia muricata (Harv.) J. Ag. X X Euthora cristata (L.) J. Ag. X Farlowia mollis (Harv. & Bail.) Farl. & Setch. X X X Fauchea sp. X Gigartina sp. X X Gloiopeltis furcata (p. & R.) J„ Ag. Grateloupia sp. X X Halosaccion glandiforme (Gmel.) Rupr. X X Iridaea sp. Kallymenia sp. X Lithothamnion sp. X Lithothrix aspergillum J„E. Gray X Membr anop ter a sp. X X Microcladia sp. X X X Odonthalia kamtschatika (Rupr.) J. Ag. X X Odonathalia sp. X xxx Opuntiella californica (Farl.) Kylin X Petrocelis middendorfii (Rupr0) Kjell. Plocamium sp. X Polyporolithon sp. X Jun. 65 Dec. 65 Jun. 66 Dec. 66 S L M H S L M H S L M H S L M H
Polysiphonia sp. X X Porphyra sp. X X Pterosiphonia sp. X X X x x Rhodoglossum sp. X X X Rhodomela larix (Turn.) c. Ag. X X X X X Rhodvmenia palmata (L.) Grev. X Rhodymenia sp. X X X X PHAEOPHYTA Alaria fistulosa p. & R. x X X Alaria marginata P. & R. X X Alaria tenuifolia Setch. X X X X Costaria costata (Turn.) Saund. X X Cymathere triplicata (p. & R.) j.Ag. X X X CVstoseira osmundaceae (Menz.) C. Ag. X X X Desmarestia viridis (Mull.) Lamour. X X X Desmarestia sp. X X Fucus sp. X X Hedophvllum sessile (C. Ag.) Setch. X X X Heterochordaria abietina (Rupr.) S. & X Laminaria groenlandica Rosenv. X X X X X Laminaria longipes Bory X X X X Leathesia difformis (L.) Aresch. X Mvelophvcus intestinale Saund. X Nereocvstis luetkeana (Mert.) p. &.R. X Pleurophycus gardneri Setch. & Saund.
Planetaria sp. X Sargassum muticum (Yendo) Fens. Scvtosinhon lomentaria (Lyngb.) J. Ag. X Soranthera ulvoidea p. & R. X X Table XIX Continued
Jun. 65 Dec. 65 Jun. 66 Dec. 66 S L M H S L M H S L M H S L M H CHLOROP HYTA Cladophora sp. x Codium setchellii Gard. x Enteromorpha sp. x X Monostroma sp. x X RhizocIonium sp. X X X Spongomorpha sp. x XXX Ulothrix X Ulva X X 145
Table XX Summary of Seasonal Cycles on Oregon Beaches
INDIAN BEACH SHORT SAND BEACH ARCH CAPE 8/66 Sand high. Sand high. Many blades missing Fresh water touches manyplants 9/66 Sand higher. Sand higher. Holdfasts buried. 10/66 Sand all gone. Sand lower Most plants ripe. Many plants ripe. 11/66 Plants ripe. Sand lower yet. plants ripe.
12/66 Some plants ripe, Sand all gone. most lack blades. All blades missing. 1/67 New blades present New blades present (1 cm) (1 cm)
2/67 New blades (3 cm) New blades (3 cm)
3/67 Small terminal sori. Small terminal sori.
4/67 Sand coming back. Sand coming back. Small terminal Many new stipes sori - all dropped and blades. at last tide. Many new stipes and blades.
5/67 Sand higher - not Sand higher - rocks yet to rocks still exposed. bearing plants.
6/67 Sand around rock Sand up, stream First plants bases. flooding measured.
7/67 Sand covers all Sand higher, has Most plants buried, rocks, holdfasts, raised stream level, stipes; only blades plants partly protruding. immersed in fresh• water 8/67 Sand down slightly. Sand very high Most plants buried. Inshore, many blades Many blades All blades present. missing; farther out, missing, plants are intact. 9/67 Sand all gone. Sand unchanged near Sand has receded 50 Most plants rocks. Few blades cm. Plants appear healthy. present except on healthy. rock tops untouched by freshwater. 146
Table XXI Summaries of in situ growth measurements on Oregon Beaches.
DATE NUMBER NUMBER MEAN MEAN DISTANCE MEASURED PREVIOUS STIPE BLADE BETWEEN HOLES LENGTH LENGTH HOLES (cm) (cm) (cm) Indian Beach (A Rock)
5 5 0 28.1 41.9 6 5 5 30.4 46.5 -7.4 7 0 * * * * 8 0 * * * * 9 0 * * * * 10 25 0 11.7 17.0 11 7 0 14.0 13.2 - 12 7 0 17.9 2.7 67/ 1 Est. * * 1.0 - 2 15 0 18.3 6.1 - 3 Est. * * 13.0 - 4 23 0 16.3 18.8 - 5 29 6 19.8 29.9 10.- 8 6 26 26 21.8 40.1 9.2 7 9 9 Buried 42.0 4.6 8 0 * Buried Buried *
Indian Beach (B Rock) (more exposed)
'67/ 5 20 0 35.9 52.1 6 10 10 37.8 66.4 15.7 147
Table XXI Continued
DATE NUMBER NUMBER MEAN MEAN DISTANCE MEASURED PREVIOUS STIPE BLADE BETWEEN HOLES LENGTH LENGTH HOLES (cm) (cm) (cm)
Short Sand Beach •66/ 8 2 0 14.5. 16.8 - 9 2 2 14.5 17.7 1.5 10 25 0 13.1 19.9 - 11 22 5 12.6 17.6 1.1 12 0 * * * * '67/ 1 Est. * * 1.0 2 Est. * * 3.0 - 3 Est. * * 5.0 - 4 27 0 9,. 0 12.7 - 5 25 8 12,. 9 20.2 10.6, 6 24 24 15,. 1 30.2 10.8 7 20 19 14.0 33.3 3.5 8 0 0 * No blades left 9 0 0 * in measurement area
Arch Cape
•67/ 6 23 0 17.8 26.5 7 8 7 Buried 36.1 6.0 8 7 5 Buried 34.3 6.3 9 8 6 22.6 36.3 1.6
* = No Data N.B. In some instances where number measured is greater than number of holes, holes were present but obviously older than one month. 148
Table XXII Growth of Multi-punched Blade of L. sinclairii
in situ at Indian Beach.
Distance (cm) Distance (cm) Growth at time of after one per punching Month Month holes
23 May '67 20 Jun »67 Blade base to A 4.0 8.0 4.0 (2 cm/2 cm) A to B 2.0 3.5 1.5 B to C 2.0 2.5 0.5 C to D 2.0 2.2 0.2 D to E 2.0 2.1 0.1 E to F 2.0 2.0 0.0 F to G 2.0 2.0 0.0 G to H 2.0 2.0 0.0 H to I 2.0 2.0 0.0 149
Table XXIII Dimensions of pressed Specimens of
L. longipes from 33 Alaskan sites.
SITE: BLADE WIDTH (cm) STIPE LENGTH(cm) Min. Max. Mean Min. Max. Mean Moderately sheltered Chichagof Pt., Attu Is. 0. 5 2. 0 1. 2 2. 0 11. 0 7. 2 Trapper's Cove, Adak Is. 1. 5 3. 5 2. 6 3. 0 8. 0 4. 8 Eagle Rock, NE Hbr., Sanak Is. 3. 5 3. 5 3. 5 4. 0 7. 0 5. 5 Gurney Bay, Kodiak Is. 0. 5 4. 5 1. 2 5. 0 15. 0 6. 7 Aats Bay, Coronation Is. 1. 0 4. 0 1. 7 4. 0 9. 0 7. 3
TOTALS 0. 5 4. 5 2. 0 2. 0 15. 0 6. 3
Moderately exposed Casco Bay, Attu Is. 2. 0 3. 0 2. 3 3. 0 12. 0 7. 5 Cape Agagdak, Adak is. 0. 5 1. 5 1. 0 7. 0 20. 0 11. 1 Ram Pt. Beach, Unalaska Is. 1. 5 3. 0 2. 0 1. 0 10. 0 8. 5 Staraya Bay, Unalaska Is. 1. 0 1. 0 1. 0 llo 0 11. 0 11. 0 Cape Sarichef I, Unalaska Is. 1. 5 2. 0 1. 6 7. 0 15. 0 11. 2 Cape Sarichef II, Unalaska Is. 2. 0 4. 0 3. 0 3. 0 5. 0 4. 0 E. Anchor Cove, Unimak Is. 2. 0 2. 5 2. 2 9. 0 25. 0 17. 3 Eagle Rock, NE Hbr., Sanak Is. 1. 0 2. 0 1. 5 5. 0 20. 0 7. 0 Nagai Is. 1. 5 1. 5 1. 5 7. 0 9. 0 8. 0 Paul Is. 0. 5 1. 0 0. 7 6. 0 6. 0 6. 0 Chirikof Is. 1. 5 2. 0 1. 7 8. 0 9. 0 8. 5 pasagshak Pt., Kodiak Is. 1. 5 3. 5 3. 0 2. 0 7. 0 3. 0 Cape Chiniak, Kodiak Is. 2. 0 5. 0 3. 3 5. 0 9. 0 8. 1 Chiniak Is. 1. 0 3. 0 2. 8 8. 0 21. 0 16. 3 Peril cape, Afognak Is. 1. 5 2. 0 1. 6 2. 0 6. 0 4. 2 English Bay 2. 5 3. 0 2. 8 9. 0 11. 0 10. 0 Wingham Is. 2. 0 2. 5 2. 2 4. 0 6. 0 5. 5 Cape Spencer 1. 0 4. 0 2. 7 6. 0 15. 0 8. 3 Helm Pt., Coronation Is. 0. 5 3. 0 1. 4 4. 0 16. 0 8. 4
TOTALS 0.5 5.0 2.0 1.0 35.0 8.6 150
Table XXIII Continued
SITE: BLADE WIDTH (CM) STIPE LENGTH(CM) Min. Max. Mean Min. Max. Mean
Fully exposed Murder Pt., Attu Is. 1.5 3.5 2.6 5. 0 14.0 9.1 North Is., Adak Is. 2.5 3.5 2.9 6. 0 12.0 7.8 Zeto Pt., Adak Is. 0.5 2.0 1.1 10. 0 15.0 12.7 Cape Aiak-Lance Pt., Unalaska Is. 1.0 2.5 1.6 9. 0 12.0 10.6 Raven P t., Unimak Is. 2.5 3.0 2.8 7. 0 14.0 9.2 Chignik Bay, Nakchamik Is. 0.5 1.0 0.7 4. 0 5.0 4.5 Aghiyuk Is., Semidi islands 0.5 2.0 1.2 3. 0 15.0 9.9 Kayak Is. 2.5 3.5 2.8 7. 0 13.0 9.5 Cape Ommaney, Baranof Is. 1.0 2.0 1.5 5. 0 11.0 9.0
TOTALS 0.5 3.5 1.9 3.0 15.0 9.1
EXTREME MINIMA & MAXIMA & TOTAL MEANS: 0.5 5.0 2.0 1.0 35.0 8.4 151
XII: APPENDIX I
Summary Descriptions of Field Stations
Volga Island, Sitka, Alaska. (57°02.5'N, 135°20.8'W) Rocky reef with many loose rocks lying in tide pools and surge channels. Moderately to fully exposed to surf.
Mean annual seawater temperature = 8.5 C; salinity = 27.7%c Mean annual air temperature = 6.3 C; precipitation = 96.57 inches. Neither L. sinclairii nor L. longipes present. Site used for transplant studies only. Observations made: 1965: June, December 1966: July, December
Aats Bay, Coronation Island, Alaska«, (55°52.7'N, 134°16'W) Rocky reef (argillite) with many deep surge channels, adjacent to beach of gravel and coarse sand. No data on mean annual seawater temperature and salinity. Mean annual air temperature— 6.3°C; Precipitation = 76.12 inches. L. longipes present in abundance„ Site used for in situ studies of L. longipes, transplant studies, and as a source for all L. longipes used in transplants and laboratory experiments. Observations made: 1965: June, December 1966: June, July, December.
River Jordan, Vancouver Island, British Columbia. (48°25.4'N 124 04-W) Loose rocks in sandy mud bottom. Moderately sheltered to moderately exposed to surf. No data on mean annual seawater temperature and salinity. Mean annual air temperature = 9.7°C; Precipitation = 73.42 inches. L. sinclairii present as small plants in very small quantities, hidden under Egregia, Hedophyllum, and Phyllospadix. Site used for transplant studies only. Observations made: 1965: August, November 1966: January, March, May, June, August. 152
Sooke Harbour, W. of Whiffen Spit, Vancouver Island, British Columbia. (48021.2'N, 123°44'W)
Loose rocks and large flat outcrops with much sandy mud. All plants frequently covered with silt layer. Moderately sheltered from surf. No data on mean annual seawater temperature and salinity or air temperature or precipitation. L. sinclairii present as small plants in small quantities, only occasionally found. Site used for transplant studies only. Observations made: 1965: July, August, November. 1966: January, February, March, May, June, August.
Stanley Park, Vancouver, British Columbia. (48°18'N, 123°06'W) Loose rocks, gravel and mud, with scattered large boulders. Fully sheltered. No data on mean annual air temperature; precipitation = 61.75 inches
Mean annual seawater temperature = 9.4°C: salinity = 27.6%0. Neither L. sinclairii nor L_. longipes present. Site used for transplant studies only. Observations made: 1965: August, November, December 1966: January, February, March, May, June. 1967: January, May, July.
Indian Point, Indian Beach, Clatsop County, Oregon. (45°55.9'N, 123°58.8'W) Sandy beach with rocky outcrops (basalt). Sand level fluctuates 1-2 m through year. Fully exposed to heavy surf.
Mean annual seawater temperature = 10.5 C; salinityQ= approx. 30.97%a. Mean annual air temperature = 11.0 C; Precipitation = 79.7 inches. L. sinclairii present in abundance as dominant plant in terms of cover. Site used for iri situ studies of L. sinclairii, transplant studies, and as a source for L. sinclairii used in transplants and laboratory experiments. Observations made: 1965: February, July, August, December. 1966: One to three times every month except January, March, April, July. 1967: One to five times on one low tide series every month, January through September. 1968: March. 153
Arch cape, Clatsop County, Oregon. (45048.2!N, 123°58.2'W) Sandy beach with large rocky outcrops. Sand level fluctuates 1 - 2 m through year. Fully exposed to heavy surf. Mean annual seawater temperature = 11.3 ; salinity = 30.97%.. Mean annual air temperature = 11.0°C; Precipitation = 81.4 inches. L. sinclairii present in abundance, but only accessible at lowest summer tides. Site used for in situ studies of L. sinclairii. Observations made: 1967: May, June, July, August, September.
Short Sand Beach, Tillamook County, Oregon. (45045.5'N, 123°58'W) Sandy beach with rocky outcrops (sandstone). Sand level fluctuates 1 - 2 m through year. Freshwater stream flows across beach at low tide, often covering some of plants studied. Fully exposed to heavy surf. Mean annual seawater temperature = 10.0 C; insufficient data on salinity. Mean annual air temperature = 11.0°C; precipitation = approx. 81.4 inches. L. sinclairii present in abundance. Site used for in situ studies of L. sinclairii, transplant experiments, and as a source for L. sinclairii used in transplants and laboratory experiments. Observations made: 1966: June, August, September, October, November, December. 1967: Once or twice on one low tide series every month, January through September. 1968: March.