VEGETATIONAL ANALYSIS OF THE

COASTAL FOREST ZONE

IN OLYMPIC NATIONAL PARK,

A Thesis

Presented to

the Department of Biology

Western Washington State College

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

Andrew Michael Kratz

June 1975 WASHINGTON NATURAL HERITAGE PROGRAM BLDG 17 AIRDUSTRIAL CENTER • OLYMPIA WA 98504 • 206.753.2449

September 20, 1978

Dr. J. F. Franklin USDA Forest Service 320 Jefferson Way State University

Dear Jerry,

Hows it feel to be clean again? A little lime seems to have taken care of the moss on the north side of my nose.

I am enclosing a copy of my thesis, with apologies again for the three year delay. I will confess to a good deal of dissatisfaction with my thesis, which is one reason I have never entertained the idea of publishing the information. Soils data were poorly collected out of ignorance. Samples should have been air dried soon after collection, not with as much as a two week delay. Texture analysis was incomplete for "lack of time" and not knowing how to deal with heavy soils which once dried formed equivalents to adobe bricks - Calgon would not cause dispersion of the clays and grinding either reduced the minibricks to sand size or reduced the sand component to clay or silt size. In other words, there isnt much information in my soils data.

Tree growth rate data was hampered by a dull increment borer and my inexperience telling me that either I was doing something wrong or there was something wrong with the trees (not all the cores came out corkscrewed). I finally did get a sharp one.

I have distinct reservations on the use of prominence values beyond all other doubts of the implications of the other number games I used. Multiplying cover and frequency (acutually square root of frequency) renders indistinguish- able the very different situations of high cover with low frequency and the reverse.

For example:

SPECIES A - All 20 Daubenmire microplots with 5-25% coverclass (mean cover 15% and frequency = 100%) has a PV=150.

SPECIES B - 6 microplots in the 95-100% coverclass, remainder with none (mean cover = 29.25% and frequency = 30%) has a PV=160. September 20, 1978 Page 2

SPECIES C - 5 microplots in the 95-100% coverclass, others with none (mean cover = 24.38% and frequency = 25%) has a PV=122.

SPECIES D - 15 microplots in the 5-25% coverclass, 5 microplots in the 0-5% coverclass. This species has actual coverage varying from 4 to 10%. (Mean cover = 11.88% and frequency = 100%). PV=119.

Using PVs tends to obscure the similarity in distribution of species A and D and masks the differences between C and D. Species A and B would be weighted more similarly than would species B and C.

These are of course "theoretical" considerations and perhaps in the overall analysis of communities it doesnt make too great a difference. Nevertheless, I have little faith in number-manipulation that has such great potential for obscuring ecological "reality".

I want to thank you for having me along in the Hoh; it was good to be a part of the team. Though my field experience has been limited, that venture was the most outstanding - I wish it could have been longer. The group spirit was tremendous and the interdisciplinary approach made the campfire talks all the more interesting. Im looking forward to seeing the results.

Take care, and let me know if theres something I can do.

Yours truly,

Andrew Kratz

AK:pl

Enclosure iii

ACKNOWLEDGEMENT

I wish to thank my advisor, Dr. Richard W. Fonda, for his guiding influence and assistance during the full course of this study. I must extend a very special thanks to my youngest brother, Richard E. Kratz, for the very considerable help, companionship, and encouragement he gave me over many long miles during my summer in the field. Grateful acknowledgement is made to

Mi. Michael Bortel for his valuable help with computer analysis of data; to the administration and staff of Olympic National Park, and in particular

Mr. William Lester and Mr. Michael Kalahar, for their cooperation and provision of shelter when most needed; and to Drs. Ronald J. Taylor and -

Franklin C. Raney for their constructive critique of the manuscript. Finally,

I wish to thank the numerous unnamed people at home and on the Olympic

Peninsula who helped in so many ways to make this study possible. iv TABLE OF CONTENTS

ACKNOWLEDGEMENT iii

LIST OF FIGURES vi

LIST OF TABLES vii

INTRODUCTION 1

STUDY AREA 3

Location 3

History 3

Geology 3

Climate 7

METHODS 12

Vegetation 12

Field work 12

Data analysis 12

Tree growth rates 15

Soils 17

RESULTS 18

Vegetation 18

Picea sitchensis-Alnus rubra/Rubus spectabilis community 18

Picea sitchensis/ shallon community 18

Picea sitchensis/ community 18

Picea sitchensis/Carex obnupta community 23

Picea sitchensis/Maianthemum dilatatum community 23

Picea sitchensis/bryophytes community 24

Picea sitchensis-Tsuga heterophylla/Blechnum spicant community 24

Tsuga heterophylla-Picea sitchensis/Polystichum munitum community 25

Tree growth rates 26

Soils 26 V

DISCUSSION 33

Pattern 33

Disturbance 36

Phytogeography 37

SUMMARY 38

LITERATURE CITED 39

VITA 41 vi

LIST OF FIGURES

Figure

Study area and numbered sample plot locations 5

Estimated evapotranspiration for 15 cm water storage capacity soil on Tatoosh Island, calculated by monthly means for a 30 year period from 1931 to 1960 10

Hytherographs from the tropical rain forest in Guyana, the boreal forest in Manitoba, Canada, and the coastal Picea forest in Washington 11

Sampling plot 14

Phenogram showing the grouping of stands into communities 16

Transect taken near Norwegian Memorial extending 200 m into the forest from the edge of the beach 35 vii

LIST OF TABLES

Table

Percentage frequency of wind direction by qua4rant at Tatoosh Island and Moclips 9

Mean density, mean basal area, and standard errors of the mean of different tree species in the eight communities 19

Size class distribution for mean number of trees/ha in the eight communities 20

Understory species composition of the eight communities 21

Average growth rates and ages with standard errors of the mean for Picea sitchensis 37-57 cm dbh in seven of the eight communities 27

Chemical properties of soils under seven of the eight communities 28

7. Selected descriptions of soils under seven of the communities 30 1

INTRODUCTION

Picea sitchensis occurs naturally in a narrow band along the Pacific

Coast from Kodiak Island, , to Mendocino County, . Along the west coast of Washington P. sitchensis is an element in a coastal forest band that is several kilometers wide. This forest band on the coastal plain of the western Olympic Peninsula is dominated by Tsuga heterophylla and Thuja plicata, but it is replaced by a narrow belt of Picea sitchensis-dominated forest adjacent to the beach.

There is little published information on these forests of Picea sitchensis. Few papers deal with the synecology of Picea communities, and even the autecology of the species is poorly understood. Fowells (1965) provides a good silvicultural summary. Picea sitchensis is generally considered to be a prolific species, but it produces less than its most common associate, Tsuga heterophylla. Flowering occurs between late March and April; seed fall lasts from about October through February. are scattered adequately for 0.5 km from their source, and they will germinate on almost any kind of seedbed if moisture is abundant. Silvicultural establishment is best on mineral soils with side shade and overhead light, but these conditions seldom occur in nature. Picea seedlings can also become established on rotten wood or moss, but on thick layers of moss they are subject to drought during dry spells. Along the Olympic Coast Rubus spectabilis, Sambucus racemosa, and Gaultheria shallon compete with, and easily outgrow, Picea seedlings.

Frequently only the seedlings on rotten wood survive, because of low cover on these logs. Picea sitchensis is a vigorous, fast-growing, shallow- rooted tree that readily overtops Tsuga heterophylla and Thuja plicata. It is classed as shade tolerant, but less so than these two competitors

(Franklin and Dyrness, 1973: 48). 2

Jones (1936) designated a climax forest of Tsuga heterophylla and Picea sitchensis along the Olympic coast, but he did not describe communities of this forest. Daubenmire (1969) mentioned that Picea sitchensis plays a climax role in a narrow strip along the ocean, but that it is seral to Tsuga heterophylla away from the beach. Franklin and Dyrness (1973) described the general composition of forests in the Picea sitchensis Zone along the coast of Oregon, but did not present compositional data. Cordes (1972), working on , appears to have made the only synecological study of forests dominated by Picea sitchensis. He used the Braun-Blanquet approach.

To date there has been no study of Picea sitchensis-dominated forest communities on the Pacific Coast using the approaches of American ecologists.

This study was undertaken to answer the following questions: 1) what is the extent of the band of forest dominated by Picea sitchensis on the Olympic coast? 2) what kinds of communities make up this zone? 3) what environmental factors control the pattern of plant communities? I will show that the Picea sitchensis zone is restricted to within 200 m or less of the beach along the Olympic coast, and that it comprises at least eight plant community types. Community pattern, however; could not be related to any single limiting factor.

Botanical nomenclature follows Hitchcock and Cronquist (1973). 3

STUDY AREA

Location.- Olympic National Park is located on the Olympic Peninsula in the northwest corner of the State of Washington. The Park includes most of the Olympic Mountains and a separate 8,100 ha Pacific Ocean coastal strip stretching 80 km from just above the Ozette Indian Reservation on the north to the border of the Quinalt Indian Reservation on the south. The average width of this strip is about 2 km. Park property is interrupted along the

Pacific beach only by the Ozette, Quillayute, and Hoh Indian Reservations.

This coastal strip constitutes the study area (Fig. 1).

History.- Indians have lived along the coast for at least 2,000 years

(Kirk, 1974) and perhaps for 5-6,000 years (Kirk, pers. comm.) with little modification of the natural vegetation before the twentieth century

(Fagerlund, 1954). Europeans first entered the Olympic Peninsula in the late

18th century, and by 1840 fur trade and white settlement had greatly increased.

There has been no logging in the coastal strip, however, largely because there was no practical way to transport timber from the area. Only local land clearing and lumber needs have been met.

There was a brief riverine gold rush in 1894 along the coast; the most productive area was between the Quillayute River and Cape Flattery. The vegetation was hardly disturbed, however, because the gold was removed from riverbeds rather than from the land.

The coastal strip was added to the Olympic National Park in 1953, although it had been under National Park Service administration for some years prior to 1953.

Geology.- Little study has been made of the geology of this coast. The most recent and detailed work covers the area from Point Grenville north to the Hoh River (Rau, 1973). The following discussion is based primarily on the work of Rau (1973) and Danner (1955). 4

Fig. 1. Study area and numbered sample plot locations (•). Totoog.h Island INSET A Cape Flattery

Cape Alava °C46 tt Indian Reservation 35 Portage iHeicid 3738 -39 Point jof Arches

32 -31 34 :33 30 River Olympic ional Park River 0sr.• 0 Mount s Olympus

0

-n Whale Cre Ouinault Indian Reservation Point Grenville N • Moclips

20 19 18 Aberdeen Rialto Beach Grays Harbor

10 20 30 km

INSET B

N ° Norwegian Mlmorial • 14. 15 67 (-7

10 .11 .13 transect

28 41

r(:) 27 .3 Destruction Island 2

Ca mpground 23o Kalalach 24. 26

25 5 km

1 km The geology of the coast is complicated because of eustatic, isostatic, and other tectonic changes in land-sea level relations. In the late Pleisocene eustatic changes were small; a rise in sea level of about 2 m may have occurred during the Hypsithermal. Isostatic and other tectonic changes are usually large, measured in hundreds of meters, but are hard to distinguish between in areas that have been glaciated (Heusser, 1960: 189).

Rock outcrops along the coast are largely sandstones, shales, and conglomerates of marine origin, but some are volcanics composed of several kinds of dark, basic lava. The oldest rocks along the coast are possibly of lower Cretaceous age (Soleduck Formation). Sedimentary rocks of Oligocene,

Miocene, and Pliocene age are also found along the beach. Several offshore islands and reefs are formed of volcanic rock occurring with Miocene sediments.

During the Eocene the present coast was under the sea. Lava flows of the Metchosin volcanics spread out on the sea floor, but since uplift of the

Olympic Mountains only remnants exist along the coast, mostly at Point of

Arches, Portage Head, and probably Point Grenville (Fig. 1). After lava deposition the area underwent several erosion cycles.

Most of the coastal plain is mantled with unconsolidated marine or freshwater sediments including sand, gravel, clay, and peat which were deposited at several different times during the Pleistocene. Some of the deposits in the area between Point Grenville and the Hoh River may have come from alpine glaciers that reached the present coastline. After the deposition of the younger sand and gravel deposits, and probably before much vegetation had developed, windblown sand and silt covered the coastal area to a depth of

1 m or more.

In the area between Whale Creek and Kalaloch (Fig. 1) late Pleistocene sand and gravel deposits are revealed in wave-cut cliffs. These materials once formed a broad surface that extended from the foothills of the Olympics to sea level at some distance from the present coast. Destruction Island,

6 km offshore, was once part of this surface, perhaps only 6,000 years ago.

Eastward erosion has produced cliffs ca. 25 m high along the present coastline, but the cliffs at Kalaloch are lower because of downwarping.

Crustal movements during the Pleistocene in the Soleduck Valley area tilted and folded some of the older Pleistocene sediments. Southward, the crust was downwarped throughout the Pleistocene epoch, with the central axis in the Kalaloch area.

Thousands of years of erosion formed the present coastline, and these forces are likely to continue. Land surveys in this century show that there is an average erosion rate of 1.1 m per year in some places south of the

Hoh River. From 1902 to 1962 two adjacent government lots lost 51 and 60 percent of their total surfaces. This high rate of erosion is partly because of the susceptability to erosion of sand, gravel, and clay deposits. Part of the Hoh rock assemblage, another deposit which is common along the beach, is highly erodable. Technically this portion is a tectonic melange, consisting of chaotically mixed resistant rocks set in a fine-grained matrix of softer materials. These deposits are structurally weak because of the clay minerals which expand when wetted by waves and precipitation. Consequently, the Hoh melange slumps periodically where it is exposed along the coast.

Climate.- The climate of the Picea sitchensis Zone is the wettest and mildest of any northwestern vegetation zones (Franklin and Dyrness, 1973).

The Cascade Mountains to the east shield the Olympic Peninsula from arctic air flows in the winter and continental air flows in the summer. The west coast of the peninsula is, however, exposed to gale force winds and heavy rains during winter. On the coast, winds blow most frequently from the east during fall and winter, but from the south or west during spring and summer (Table 1). The strongest winds are associated with winter storms moving eastward across the Pacific Ocean (Phillips and Donaldson, 1972).

Precipitation averages 1800-2300 mm per year. It is distributed unevenly through the year with a maximum in winter; December is the wettest month, July is the driest. Assuming the soil has a 15 cm waterholding capacity in the root zone, the will experience a water deficit of less than 1 cm during the summer months as indicated by the difference between potential and actual evapotranspiration (Fig. 2). Actual moisture stress experienced by the plants in any given habitat is likely to differ from this estimation because of soil type, water table depth, drainage patterns, rooting characteristics of the individual plants, etc. Figure 2 represents an average for a 30 year period and does not reflect the extremes which can have important effects on the vegetation. Rainfall intensity is light to moderate.

Fog is most frequent during the summer, with fog drip resulting from condensation in the canopy.

Mean monthly temperatures range from 6 C in winter to 14 C in summer.

During the warmest summer months the afternoon temperatures are about 15-20 C and the lows are around 7-12 C. In winter the average maximums are around

10 C and the average minimums are near 2 C. Maximums reach 0 C or lower on an average of about 7-12 days during the year (Phillips and Donaldson, 1972).

Relative humidity ranges between 80-95% throughout the day year round.

Annual sunshine along the north Pacific coast is less than 40% (Heusser, 1960:

30).

A comparison of hytherographs from Tatoosh Island, Washington, Fort Good

Hope, Manitoba, and Georgetown, Guyana reveals that the spruce zone in the

Olympics has the general range of precipitation and the overall uniformity of temperature of the tropical rainforest, but that mean monthly temperatures are more similar to those of the warmest months in the boreal forest (Fig. 3). TABLE 1. Percentage frequency of wind direction by quadrant at Tatoosh Island and Moclips. Calm periods are not included. Data based on an 11 year period from 1948 to 1958 (Phillips and Donaldson, 1972).

Tatoosh Island Moclips N E S w N E S W J 4.8 54.2 28.7 11.9 5.1 66.8 11.9 6.9 F 6.0 42.2 32.2 18.8 5.1 58.0 12.9 13.9 M 7.8 38.9 29.6 22.4 10.8 37.9 16.0 24.2

A 9.1 26.6 34.2 28.4 8.7 25.6 19.8 33.7 M 8.0 15.6 40.1 34.2 9.1 16.9 17.2 45.2 J 6.2 14.2 47.9 30.0 8.3 9.5 10.9 56.3

J 4.9 8.7 62.8 22.2 11.0 7.9 14.4 45.4 A 6.2 14.1 63.9 13.9 8.0 11.4 13.0 43.4 S 12.9 27.7 42.6 14.3 10.2 19.3 13.1 31.1

0 9.7 /1/1.8 30.5 13.5 9.9 40.1 22.0 13.4 N 5.7 53.5 27.4 12.6 11.0 55.2 14.5 11.4 D 3.0 /1/1.0 33.3 18.8 9.8 50.6 19.4 10.2 10

30 -

25 - AVERAGE MONTHLY PRECIPITATION

AVERAGE POTENTIAL MONTHLY EVAPOTRANSPIRATION

20 - AVERAGE ACTUAL MONTHLY EVAPOTRANSPIRATION

10 - WATER DEFICIT

5 -

1 I 1 i I I I I I JAN FEB MAR1 APR MAY JUN JUL AUG SEP OCT NOV DEC

Fig. 2. Estimated evapotranspiration for 15 cm water storage capacity soil on Tatoosh Island, calculated by monthly means for a 30 year period from 1931 to 1960. Average actual evapotranspiration and potential evapotranspiration are different only from June through August. Data from Phillips and Donaldson (1972). 11

30 —

10 7 c9 2 1 12 6 GEORGETOWN, GUYANA

20--

8

10 10 — 11

12 3 2 TATOOSH ISLAND, WASHINGTON

10

-10 -. FORT GOOD HOPE, MANITOBA

-20 — 11

1 -30 — 1 1 I I 5 10 15 20 25 30 35 MEAN MONTHLY PRECIPITATION (cm)

Fig. 3. Hytherographs from the tropical rain forest in Guyana, the boreal forest in Manitoba, Canada, and the coastal Picea forest in Washington. Months are numbered consecutively (1 = January). Data from Phillips and Donaldson (1972) for Washington, Fonda (pers. comm.) for Guyana and Manitoba. Data for Washington based on the 30 year period from 1931 to 1960 (Phillips and Donaldson, 1972). Data for Guyana and Manitoba based on unknown period (Fonda, pers. comm.). 12

METHODS

Vegetation

Field work.- A preliminary survey of the Picea forests in the study area showed that this coastal zone included several different community types which

could be largely differentiated on the basis of understory vegetation. A general classification of the vegetation by the major understory components was then devised and a total of 38 sample plots were studied during summer 1973, with between three and fifteen plots per tentative community type.

All stands but one were sampled using a single 15 x 25 m macroplot

randomly located in a homogeneous portion of the stand (Fig. 4). A 25 m line was run down the center of the macroplot to establish its length, then a 7.5 m

line was repeatedly run perpendicularly to the centerline to establish the width and locate the understory plots within the macroplot. The herbaceous understory was sampled by systematically placing a 20 x 50 cm frame at 20 points

inside the macroplot, and were sampled in eight 1 m radius circular plots regularly spaced within the macroplot. Shrub and herb coverage was estimated by the cover classed of Daubenmire (1959), and his mid-points were used in analyzing the data. All trees larger than 2.5 cm dbh were measured on the

15 x 25 m macroplot. Tree reproduction was counted in the 1 m radius circular plots. Bryophytes and lichens were treated together as a single component of

the understory.

Data analysis.- After all data had been collected and summarized by stand, coefficients of similarity for stands were calculated based on prominence values (PV) of all species. The values for understory species were calculated as PV = mean % cover x frequency l (Beals, 1960). An alternate formula was used for tree species (PV = basal area x density 2 ), because there were no frequency values for trees. Both methods of calculating prominence values 13

Fig. 4. Sampling plot. Trees were tallied on 15 x 25 m plot, shrubs on 1 m radius plots, and herbs on 20 x 50 cm frames. • ..!.1,-4,•=s-A.

Im RADIUS f-20x50cm FRAME

I5m 10 15 20 25

25m 15 yielded numbers between 0.28 and 945, and growth forms (tree, shrub, herb) were equally weighted; the highest prominence value in a given stand could be held by a tree, shrub, or herb.

Similarity coefficients were calculated by comparing all stands two at 00)b2w a time using the formula C = (1 where C is the coefficient of similarity, a + a and b are the sums of the prominence values calculated for stands A and B, and w is the sum of the lowest common value of the two stands being compared.

A phenogram was then constructed from the similarity matrix following the methods of Mountford as described in Southwood (1966: 342-344). The phenogram (Fig. 5) shows a grouping of the stands based on their similarities.

Division of the stands into communities was made by the selection of a minimum percentage similarity for members of the communities. In Fig. 5 the 55% similarity level shows the grouping of stands into community types described under Results. The phenogram required modification of the original community classification, after which mean values of cover and frequency for understory species, and basal area and density for tree species of each community were calculated. These values appear in Tables 2, 3, and 4. The number of plots in each finalized community type is shown in Table 2.

Tree growth rates

To assess differences in growth conditions increment borings from Picea in seven of eight communities were examined for growth rates. Only trees of

37-57 cm dbh were drilled since annular ring width decreases as a function of tree diameter even under otherwise constant conditions. Because annular rings are usually larger on the downhill side of a tree, all trees were drilled from the south side, except in the Picea-Alnus/Rubus stand in the

Rialto Beach area where they were drilled from the east side, since the coast runs east-west there. 16 0 9 PICEA/CAREX

22-

16 34 PICEA/GAULTHERIA 27 11 7 17 13 39 TSUGA-PICEA/POLYSTICHUM 15 12 37- PICEA-TSUGA/BLECHNUM 36 30- 8 25 PICEA/BRYOPHYTES 32 4 26- 28 PICEA/MAIANTHEMUM 18 2_ 40 41 PICEA-ALNUS/RUBUS 38 21 19 5 20 35 PICEA/POLYSTICHUM 31 14 29 33 1_

I I I I I r I 10 20 30 40 50 60 70 80 90 100 PERCENT SIMILARITY

Fig. 5. Phenogram showing the grouping of stands into communities. 17

Data on Picea growth rates were subjected to analysis of variance using a completely randomized design and the level of significance chosen at 5% before testing.

Soils

I sampled soils of each community shown in Table 2, except for the

Picea-Tsuga/Blechnum community which was revealed after the phenogram was constructed. Soil pits were dug in or adjacent to 17 of the macroplots, and the profiles were described. A composite 1 kg sample of soil was collected from each horizon, and air dried within 14 days. Samples were passed through a 2 mm sieve and subsequent analyses were carried out on the 2 mm fraction.

Twenty-six of the samples from 5 and 30 cm depth in 14 soil pits were analysed for pH by the glass electrode method, NO 3-N by calcium oxide extraction with color developed by phenoldisulfonic acid and read on a colorimeter, Mg and Na by ammonium acetate extraction for atomic absorption and flame spectrophotometer, respecitively. These were standard analyses performed by technicians at Washington State University, Pullman. Two of the pits were less than 30 cm deep, however, so that only 5 cm deep samples were analysed for them.

Data on chemical properties of the soil were subjected to analysis of variance using a completely randomized design and the level of significance chosen at 5% before testing. 18 RESULTS

Vegetation

Picea sitchensis-Alnus rubra/Rubus spectabilis community.- This was one

of the most common communities facing the coast. Picea sitchensis and Alnus

rubra dominated the community (Tables 2 and 3). The two species had similar

density, but Picea had a larger total basal area. Tsuga heterophylla was minor in this community. One stand, south of La Push, had Picea seedlings,

but the other two stands had no tree reproduction.

The shrub layer was about 2 m tall and had a mean cover of 83%; herbaceous

cover was 57%. The dominant understory species were Rubus spectabilis,

Polystichum munitum, Sambucus racemosa, Gaultheria shallon, and Athyrium

filix-femina (Table 4).

Picea sitchensis/Gaultheria shallon community.- This community was commonly encountered on level to steeply sloping surfaces where it sometimes covered areas of a few hectares. It was generally exposed to storm winds off the ocean. Picea sitchensis dominated the canopy (Tables 2 and 3). Tree reproduction was sparse (Table 3).

The shrub layer ranged from 1.5-3 m in height. Shrub cover averaged 85%, herbaceous cover 15%. The dominant shrub species, Gaultheria shallon, was sometimes accompanied by Rubus spectabilis (Table 4). Polystichum munitum was the most important herb species and was present in all stands of this community.

Picea sitchensis/Polystichum munitum community.- This common community occurred on level to steeply sloping surfaces. It occasionally was found next to the beach but was usually protected from ocean winds by a line of young trees or Gaultheria shallon. Picea sitchensis clearly dominated this community.

Tsuga heterophylla and Alnus rubra were also present but were always minor 1 q

TABLE 2. Mean density, mean basal area, and standard errors of the mean of the different tree species in the eight communities.

Mean Mean Density Bas1 Area Community Tree (1i/ha) (m /ha) Plots

Picea-Alnus/Rubus Picea 151 + 49 63 + 20 Tsuga 27 + 15 <1 + <1 3 Alnus 133 + 80 20 + 11 Pyrus 9 + 9 _<1 T <1 Total 320 33 Picea/Gaultheria Picea 373 + 44 74 + 9 Tsuga 9+ 9 <1 + <1 6 Alnus 13 + 9 2 + 1 Total 396 76 - Picea/Polystichum Picea 368 + 59 79 + 11 Tsuga 53 + 22 7 + 4 10 Alnus 72 + 29 8 + 3 Total 493 94 Picea/Carex Picea 222 + 105 27 + 2 Tsuga 18 + 18 4 + 4 3 Alnus 53 + 27 6 + 2 Populus 18 + 18 5 + 5 Total 311 42 Picea/Maianthemum Picea 860 + 171 67 + 7 Tsuga 33 T 33 1 + 1 4 Thuja 40 + 40 2 + 2 Total 933 70 Picea/bryophytes Picea 869 + 139 69 + 5 Tsuga 229 + 101 6 + 3 Thuja 139 + 97 3 + 2 5 Alnus 5 + 5 ,1 + <1 Pyrus 16 + 16 (1 + <1 Total 1259 79 Picea-Tsuga/Blechnum Picea 213 + 80 146 + 21 Tsuga 360 + 173 36 + 5 2 Total 573 182 Tsuga-Picea/Polystichum Picea 147 + 52 39 + 10 Tsuga 693 + 154 67 + 8 4 Total 840 105 20 TABLE 3. Size class distribution for mean number of trees/ha in the eight communities.

1 cm tall- 3-15 16-30 31-61 62-92 93-122 123+ Community 2 cm cm cm cm cm cm cm Tree dbh dbh dbh dbh dbh dbh dbh

Picea-Alnus/Rubus Picea 519 44 - 53 27 9 18 Tsuga - 18 9 - - - - Alnus - - 9 124 - - - Pyrus - 9 - - - - -

Picea/Gaultheria Picea 103 4 62 253 49 - 4 Tsuga 51 4 4 - - - - Alnus - - 4 9 - - -

Picea/Polystichum Picea 343 51 51 176 67 21 3 Tsuga 432 13 13 19 8 - - Thuja - - 21 51 - - -

Picea/Carex Picea 1,029 18 98 80 27 Tsuga 309 - - 18 - Alnus 206 - 18 27 - Populus - - - 9 -

Picea/Maianthemum Picea 154 113 413 313 20 Tsuga - - 33 - - Thuja - - 27 13 -

Picea/bryophytes Picea 124 91 469 293 16 Tsuga 1,050 101 112 16 Thuja - 59 75 5 - Alnus - 16 - - - Pyrus - 16 - - - Rhamnus 247 - - - -

Picea-Tsuga/Blechnum Picea 1,699 - 13 67 53 40 40 Tsuga 4,324 147 93 80 40

Tsuga-Picea/Polystichum Picea 3,088 - 7 93 33 13 - Tsuga 65,164 47 327 307 7 - 7

nearly all individuals in this size class are 1-30 cm tall. TABLE 4. Understory species composition of the eight communities. Only species with at least 1% mean cover in at least one community are listed. C = mean cover, F = frequency; "+" means a value of less than one, "-" means absent.

Picea-Alnus Picea/ Picea/ Picea/ Picea/ Rubus Gaultheria Polystichum Carex Maiamthemum Species C F C F C F C F C F bryophytes + lichens 18.1 58.3 15.8 53.3 10.4 32.0 7.3 23.3 19.9 61.7 Polystichum munitum 41.7 70.0 6.9 21.7 73.4 88.5 21.2 33.3 4.1 16.7 Blechnum spicant + 3.3 + 2.5 + 4.5 + 1.7 1.1 8.3 Athyrium filix-femina 7.8 23.3 1.5 2.5 1.6 6.0 2.0 10.0 - - Dryopteris austriaca 1.6 10.0 1.0 5.0 + 4.0 - - - - Pteridium aquilinum + 1.7 4.0 11.7 - - 2.0 3.3 - - Maianthemum dilatatum 1.0 6.7 + 5.8 1.1 13.0 + 10.0 71.5 98.8 Lysichitum americanum + + + 1.7 + + + 5.0 Tiarella trifoliata + 1.7 - - + 2.0 - - - - Carex obnupta + + + 2.5 + + 55.2 73.3 + + Galium triflorum + + + + 2.4 19.5 + 6.7 Montia siberica 1.6 21.7 + + + 12.0 - - - - Gaultheria shallon 11.7 33.3 79.9 100.0 + 15.0 5.4 20.8 + 18.8 Rubus spectabilis 58.7 95.8 4.3 22.9 + 3.8 + 8.3 + + Menziesia ferruginea - - + 8.3 + 3.8 - - - - Vaccinium parvifolium + 4.2 + 4.2 + 1.3 + + Sambucus racemosa 12.9 41.7 - - + 3.8 - - - - TABLE 4 continued

Picea/ Picea-Tsuga/ Tsuga-Picea/ bryophytes Blechnum Polystichum Species C F C F C F bryophytes + lichens 43.3 97.0 13.6 62.5 37.5 86.3 Polystichum munitum + 1.0 14.7 30.0 12.8 26.3 Blechnum spicant + + 34.8 60.0 3.8 10.0 Athyrium filix-femina - - 2.4 12.5 1.8 6.3 Dryopteris austriaca - - 1.1 7.5 1.2 10.0 Pteridium aquilinum + + - - - - Maianthemum dilatatum + 7.0 4.5 45.0 + 5.0 Lysichitum americanum + 1.0 - + 1.3 Tiarella trifoliata - - 3.8 40.0 + + Carex obnupta - - + 2.5 - - Galium triflorum + 4.0 - - + 12.5 Montia siberica - - - 1.9 27.5 Gaultheria shallon + 7.5 + 1.0 25.0 Rubus spectabilis + 2.5 + + + 6,3 Menziesia ferruginea + + - - + 15.6 Vaccinium parvifolium + 10.0 + 12.5 2.3 43.8 Sambucus racemosa - - - - 23 components (Table 2). There was some tree reproduction in four of the stands of this community, and it was about equally divided between Picea and Tsuga

(Table 3).

All shrub species had a mean cover of less than 1% and total cover for the shrub layer was less than 2%. The herb layer was strongly dominated by

Polystichum munitum which had a much higher cover than in the Picea/Gaultheria community (Table 4). Galium triflorum was the next most important species, followed by Athyrium filix-femina and Maianthemum dilatatum.

Picea sitchensis/Carex obnupta community.- This community comprised a very small percentage of the Picea-dominated forests on the coast. It occurred in depressions where the water table could be high during the rainy season.

Many of these depressions apparently are old stream channels. Picea sitchensis dominated the community (Tables 2 and 3). Picea had the greatest mean density and mean basal area of all tree species in the community. Tsuga heterophylla and Alnus rubra were minor components, and Populus trichocarpa was found in only one of the three stands sampled. Trees grew primarily near the edges of the community. Tree reproduction occurred almost exclusively on fallen logs and was found in all stands sampled.

The shrub layer was depauperate, and Gaultheria shallon accounted for nearly all shrub coverage (Table 4). The herb layer had a total coverage of

84% and was dominated by Carex obnupta. Polystichum munitum consistently occurred in or next to the community; Athyrium filix-femina and Pteridium aquilinum were less important herbs.

Picea sitchensis/Maianthemum dilatatum community.- This uncommon community usually covered only small area. Often it formed a narrow band parallel to the edge of the coastal cliffs, but it also occurred on well 24 drained sand and gravel areas in the lee of thick growths of young trees :Ind driftwood piles at the beach edge. Picea sitchensis dominated the tree layer

(Tables 2 and 3). Tsuga heterophylla and Thuja plicata were found occasionally in the community near Kalaloch.

Total shrub cover was less than 1% (Table 4). The herb layer was lush,

Maianthemum dilatatum accounting for most of the cover.

Picea sitchensis/bryophytes community.- This community was often adjacent to the Picea/Maianthemum community. It occupied the tops and edges of cliffs facing the ocean, and gravels behind the driftwood pile at the winter high tide line. This community was common and extensive near Kalaloch.

The tree layer was composed of thin, often twisted trees, and it was dominated by Picea sitchensis. Tsuga heterophylla also occurred in the canopy but had less than 33% of the density and basal area of Picea (Tables 2 and 3). Thuja plicata was a minor component when present. The number of tree seedlings in this community was intermediate compared to the other communities studied (Table 3). Tsuga was clearly represented by more individuals in the two smallest size classes than was Picea (Table 3).

The shrub layer had a total cover of less than 1% (table 4). The herb layer was of equally minor importance with a total cover of just over 1%.

Bryophytes and lichens were the most important understory component with a total cover of 43%. This coverage is equal to that of bryophytes and lichens in the Tsuga heterophylla-Picea sitchensis/Polystichum munitum community, so that the absence of shrubs and herbs is more important than the high bryophyte and lichen coverage.

Picea sitchensis-Tsuga heterophylla/Blechnum spicant community.- This community was only encountered at Cape Alava, where it occurred on old marine 25 terraces or other surfaces transitional between Picea-dominated and

Tsuga-dominated forests. Only Picea and Tsuga were sampled in the tree layer of this community, although Thuja plicata occurred nearby. Size class distribution indicated that Tsuga will eventually dominate this community

(Table 3). Tsuga accounted for 67% of the seedlings and had a large number of individuals between 3-61 cm dbh. Picea was largely represented by individuals with diameters greater than 31 cm , and there is a break in its size class distribution in the 3-15 cm range.

Vaccinium parvifolium was the most important of the shrub species, all of which totaled less than 1% cover. The herb layer had 31% and 92% total cover in the two stands sampled, yielding a mean of 61%. Blechnum spicant accounted for a majority of this layer while Polystichum munitum accounted for over half of the balance (Table 4). Maianthemum dilatatum, Tiarella trifoliata, and Athyrium filix-femina were also common.

Tsuga heterophylla-Picea sitchensis/Polystichum munitum community.- This community marks the shift from Picea forests to Tsuga forests. Stands of this community occupied aggraded surfaces or older surfaces behind the sea cliffs.

The tree layer was dominated by Tsuga heterophylla (Table 2). Picea sitchensis was present in low density, although the trees were large (Table 3). Tree reproduction, particularly for Tsuga, was higher than in any other community studied.

The shrub layer had a total cover of 4%. Vaccinium parvifolium was the dominant shrub species and was present in all stands sampled. The herb layer was open, total cover being 23%. The most important herb species were

Polystichum munitum and Blechnum spicant (Table 4). 26

The averages for this community exclude stand 12 which contained a higher cover of Lysichitum americanum than in any other stand sampled in this

community. Stand 12 also lacked a fern component.

Tree growth rates

With one exception, no significant differences were found among the growth

rates of Picea trees in the different community types or between locations along the coast. Only the trees in Picea/bryophytes communities at Norwegian

Memorial grew more rapidly than trees in the same community at Kalaloch (Table 5).

Soils

Soils in the Picea forests varied widely in chemistry, texture, and structure. For the soil chemistry data which has been analyzed statistically there was a high mean square value within groups. Although among groups mean square values were often high, the variability from site to site was too great to produce statistically significant differences. There were no significant differences for pH, Mg, Na, and NO 3-N among: different community types; vegetation with and without Tsuga; areas drained vs areas receiving drainage; well drained vs less well drained; and exposed to the ocean vs less exposed.

Communities exhibited wide and overlapping ranges for these factors. The

Tsuga-Picea communities showed a trend toward higher pH than the Picea-dominated communities. The concentrations of NO 3-N, Mg, and Na ions tended to be lower in the Tsuga-Picea communities than in Picea communities (Table 6).

Soil texture is variable along the coast (Table 7). All examples of the Picea/Carex community were in depressions on sand, but other communities showed more variation in soil type. Picea/Maianthemum occurred over sand and cobbles as well as on clay. Picea/bryophytes occupied old sand and gravel bars

and sandy clays. Picea/Gaultheria and Picea/Polystichum were found on sands 27

TABLE 5. Average growth rates and ages with standard errors of the mean for Picea sitchensis 37-57 cm dbh in seven of the eight communities. Locations: K = Kalaloch, R = Rialto Beach, N = Norwegian Memorial.

Mean Growth Mean Tree No. of Community Location Rate (cm/year) Age (years) Trees

Picea-Alnus/ Rubus R .978 + .307 53 + 18 2 K .445 + .036 90 + 0 2 Picea/ Gaultheria N .787 + .030 63 + 3 2 K .678 + .044 63 + 8 2 Picea/ Polystichum N .879 + .081 54 + 4 4

Picea/ Carex N .828 + .091 50 + 0 2

Picea/ Maianthemum N .653 + .104 68 + 3 2 R .495 + .023 93 + 8 2 K .391 + .069 113 + 16 4 Picea/ bryophytes N .772 + .048 55 + 0 2 K .330 + .079 160 + 40 2 Tsuga-Picea/ Polystichum N .805 + .084 63 + 3 4

significant difference (5% level) 28

TABLE 6. Chemical properties of soils under seven of the eight communities. Data are from two pits in each community except Tsuga-Picea/Polystichum.

Mg Na No -N Community (meq/100g) (meq/100g) (PPm)

Depth = 5 cm

Picea-Alnus/Rubus 4.0 1.91 1.14 6.10 4.6 0.72 0.39 1.40

Picea/Gaultheria 4.6 9.55 4.15 7.72 4.2 1.60 0.34 1.06

Picea/Polystichum 4.6 2.44 1.03 3.46 4.4 1.60 0.44 2.06

Picea/Carex 4.5 2.65 0.52 6.10 4.9 2.22 0.37 0.16

Picea/Maianthemum 3.8 4.16 1.30 6.20 5.8 1.60 1.34 0.70

Picea/bryophytes 3.8 5.02 2.50, 5.80 4.2 3.60 1.21 5.00

Tsuga-Picea/Polystichum 5.1 1.07 0.29 2.20

no corresponding 30 cm value. 29

TABLE 6 continued

Depth = 30 cm

Picea-Alnus/Rubus 4.7 2.22 0.85 4.00 4.8 0.43 0.30 3.10

Picea/Gaultheria 4.7 4.38 3.25 1.70 5.2 0.48 0.15 0.08

Picea/Polystichum 5.2 1.18 0.55 0.85 4.9 0.77 0.17 0.30

Picea/Carex 5.3 0.77 0.19 0.15 5.7 0.98 0.19 0.30

Picea/Maianthemum 4.1 2.65 0.78 2.06

Picea/bryophytes 4.3 2.22 0.64 2.90 4.1 1.91 0.85 2.20

Tsuga-Picea/Polystichum 5.2 0.38 0.17 0.16 10

TABLE 7. Selected descriptions of soils under seven of the communities. Locations: K = Kalaloch, R = Rialto Beach, N = Norwegian Memorial.

Community Location Horizon Depth(cm) Structure Texture

Picea-Alnus/ Rubus 0 4-0 litter 1 A 0-5(10) medium crumb 5-40 medium subangular blocky C 40+ unconsolidated - sandy loam Picea/ Gaultheria 0 3-0 litter 1 A 0-6 single grain 6-22 single grain - sand C 22+ single grain - sand Picea/ Gaultheria 0 18-0 litter 1 A 0-10 medium crumb B 10-19 medium subangular blocky 1 B 19-43 medium subangular blocky - 2 loam C 43+ massive - clay loam Picea/ Polystichum 0 11(9)-0 litter 1 Al0-8 strong crumb to weak subangular blocky A 8-18 strong crumb to weak 3 subangular blocky - clay B 18-32 medium subangular blocky - 2 clay loam B 32-50 medium subangular blocky - 3 clay loam C 50+ massive - clay loam Picea/ Polystichum 0 4-0 litter 1 A 0-3(8) single grain 3-26(36) single grain - sand C 26+ single grain - sand Picea/ Carex 0 3-0 litter 1 A 0-12 single grain 12-20 single grain - sandy clay loam C 20+ single grain - sand Picea/ Maianthemum" 0 5-0 litter 1 R A 0-40(45) single grain - sand 40-65(85) single grain C 65+ single grain - sand 31

TABLE 7 „continued

Picea/ Maianthemum 0 5-0 litter 1 K Al 0-5(8) crumb to subangular blocky B 5-12 massive 1 C 12+ massive - clay Picea/ bryophytes 0 2-0 litter 1 N Al 0-9 crumb B 9-20 subangular blocky 1 B 20-39 subangular blocky 2 B 39-75 subangular blocky - loam 3 C 75+ massive - sandy clay Tsuga-Picea/ Polystichum 0 6-0 litter 1 N A 0-10 strong crumb to weak subangular blocky B 10-30 medium subangular blocky - clay C 30+ massive - clay and on clay loams. Picea-Alnus/Rubus stands occurred on disturbed areas independent of soil type. Tsuga-Picea/Polystichum stands were on heavier soils associated with old marine terraces. Profile development ranged from poor- to well-developed, and the soils exhibited a similar wide range in structure (Table 7). The solum for all soils was less than 1 m thick. 33

DISCUSSION

Pattern.- The coastal forests dominated by Picea sitchensis in Olympic

National Park comprise at least eight communities. The distribution of these communities cannot be accounted for by a single limiting factor. Cordes (1972) reported that the communities he described under the Braun-Blanquet approach were distributed along the coast in response to two major factors: parent material and intensity of salt spray reaching the forest floor. Along the

Olympic coast I did not find significant differences in salt, as measured by

Na ion concentration in the soil, among the communities I described. Nor did I find significant differences among community types for other chemical

factors or soil texture. General growth conditions, as measured by the growth rate of Picea in the different communities, also showed no significant differences among community types. The distribution of these communitites along the coast seems to be controlled by a complex interaction of several physical factors, and perhaps by competition among understory species.

Daubenmire (1942) demonstrated that species under the canopy of the

Pinus ponderosa forests of the Northern Rockies had narrower ranges of tolerance

to environmental gradients than did the overstory trees. The parallel between

the Pinus ponderosa forests and the coastal Picea sitchensis forests is clear.

Picea sitchensis ranges from Alaska to California, and on the Olympic Coast

it is distributed through nt least eight communities and dominates the canopy

in five of them (Table 2). Picea must have a broader tolerance range than

many of the understory species associated with it along the Olympic Coast.

Competition among these understory species of more limited tolerance ranges

is presumably greater than for Picea, and the pattern of community distribution

may reflect this competition.

Although the actual factors governing the distribution of communities

are not known, the general pattern of distribution can he described. Typically the Olympic coast is eroding eastward, so that the distance between the beach and Tsuga-dominated forests is short. Near Norwegian Memorial, however, is one of the few aggraded beach surfaces in the study area, and it supports a well developed forest. In this area a transect was run perpendicularly from the edge of the beach for 200 m into the forest (Fig. 6).

Herbs were tallied by recording the species that occurred at each meter mark.

Shrubs were tallied by counting stems along a 7.5 m perpendicular line on each side of the transect center line every 5 m along the transect length. All trees within 7.5 m of either side of the transect line were tallied by species.

The transect began in a Picea/Gaultheria community at the edge of the beach, traversed a Picea/Carex community (50-75 m) in an old drainage channel at the base of an old coastal bluff, passed through a Picea/Polystichum community (75-125 m) on the foreslope of the bluff, over into a Tsuga-Picea/

Polystichum community (125-175 m) on the rear slope, and ended in the transition zone to the Tsuga-dominated forest. This transect shows some of general relations between these four communities. Picea/Gaultheria usually occupies exposed sites close to the beach along this coast. It frequently occurs just beyond the winter waves or on steep slopes or cliff tops that are exposed to high wind speed and presumably salt spray. Picea/Carex communities, on the other hand, are always protected because they occupy sheltered topographic depressions of drainage channels. Picea/Polys t.ichum communities are partially protected, and frequently are adjacent to Picea/Carex communities.

Picea/Polystichum never occur in the depressions, but are often found on protected slopes that are closer to the beach than is the Picea/Polystichum community in the transect. The position of the Tsuga-Picea/Polystichum community on the top and back side of old coastal bluffs or raised marine terraces is typical of this part of the Pacific Coast. The large number 35 375 SHRUB LAYER 15 15 m contacts/25 m TREES/ha 250 10 HERB LAYER 125 contacts 1 25 rn

PICEA SITCHENSIS LGOEMIELER

TSUGA HETEROPHYLLA ALNUS RUBRA

GAULTHERIA SHALLON

RUBUS SPECTABILIS

VACCINIUM PARVIFOLIUM

FERNS

CAREX OBNUPTA

FORBS

BRYOPHYTES LICHENS

0 25 50 75 100 125 150 175 200 Distance along tronsect (m)

Fig. 6. Transect taken near Norwegian Memorial extending 200 m into the forest from the edge of the beach. 36 of Tsuga trees, concurrent with Vaccinium parivolium as an important shrub species, marks the shift away from Picea-dominated forests.

The pattern of other communities may be related to this transect. Sites similar to the Picea/Polystichum communities often support Picea-Alnus/Rubus on unstable slopes. These sites are less exposed to storms than those of the Picea/Gaultheria communities. Picea-Tsuga/Blechnum communities were only encountered in the Cape Alava area, where they occupied the top of an old marine terrace in a position analagous to the 125-150 m area of this transect.

Picea/Maianthemum communities are variable in their topographic location, but they appear to occupy sites that are more open to light than the adjacent

Picea/bryophytes communities.

Along much of this coast, any one of the Picea communities, except

Picea/Carex and Picea/Maianthemum, may grade directly into Tsuga-dominated forests.

Disturbance.- Disturbance is an important factor in the Picea forest zone on the Olympic coast. The high rate of erosion coupled with the susceptability of geologic deposits to slumping creates a state of dynamic tension between development of Picea forests and their destruction. Most ecologists in the Pacific Northwest consider Picea sitchensis a seral species that is replaced by Tsuga heterophylla, but that may be maintained in areas of repeated disturbance. In the Hoh Valley seral Picea forests develop on river terraces that are continuously being created and destroyed (Fonda, 1974).

The persistence of Picea in old growth forests along the Oregon coast has been attributed to windthrow and overstory mortality (Hines, 1971). In

British Columbia Picea has been maintained by windthrow, fire, and erosion

(Day, 1957). Succession in most mature coniferous forests near this coastal area is toward replacement of Picea sitchensis by Tsuga heterophylla

(Franklin and Dyrness, 1973; Krajina, 1969). Along the coast Picea-dominated 37 forests must be largely a product of disturbance. The Olympic coast is frequently disturbed, and Picea forests are enabled to develop on these surfaces. On the other hand, Cordes (1972) has shown that beaches on Vancouver

Island are aggrading, so that the persistence of the Picea forest belt is partly due to the continual build up of new surfaces for colonization by

Picea. The older parts of the seral Picea forest on Vancouver Island are being replaced by Tsuga-dominated forests. The persistence of Picea along the coast appears also to be due to its ability to tolerate more salt than can Tsuga (Cordes, 1972).

Phytogeography.- The Picea sitchensis Zone is variously reported in the literature as being one to several kilometers wide. Franklin and Dyrness (1973) reported that it averages a few kilometers wide in Oregon and expands to several kilometers on the Olympic Peninsula. This may describe the distribution of Picea sitchensis as a species, but it does not correspond to the distribution of communities dominated by Picea sitchensis. I have found that the Picea sitchensis-dominated forest, other than the extensions up river valleys, is restricted to a narrow band along the open Olympic coast.

This band varies in width from less than 50 m to about 200 m. It is generally a continuous strip broken irregularly at slumped areas by shrub thickets or

Alnus communities. Farther inland many of the Picea trees are large and old and are greatly outnumbered by Tsuga heterophylla and Thuja plicata. There is a similar narrow band of Picea-dominated forest types on the west coast of Vancouver Island (Cordes, 1972). 5.„ I suggest that the term "Picea sitchensis Zone" be re-defined to apply 40-00c only to this very narrow band of Picea sitchensis-dominated forests along rAC along the coast. At the same time the Tsuga heterophylla Zone should be enlarged to include the Tsuga-Picea-dominated communities along the edge of this Picea sitchensis Zone. 38 SUMMARY

Eight plant communities were recognized and described for the

Picea sitchensis forest:

Picea sitchensis-Alnus rubra/Rubus spectabilis

Picea sitchensis/Gaultheria shallon

Picea sitchensis/Polystichum munitum

Picea sitchensis/Carex obnupta

Picea sitchensis/Maianthemum dilatatum

Picea sitchensis/bryophytes

Picea sitchensis-Tsuga heterophylla/Blechnum spicant

Tsuga heterophylla-Picea sitchensis/Polystichum munitum

Texture, structure, and chemistry of the soils in these communities varied widely within and among the communities.

No single limiting factor was found to govern the distribution of the plant communities, but their general distribution along the coast was described. The role of competition in determining the community composition and distribution needs investigation.

The Picea sitchensis-dominated communities were restricted to within 200 m of the beach.

5. I have proposed that the term "Picea sitchensis Zone" be re-defined to include only the Picea-dominated forests along the coast. This proposal also called for the expansion of the "Tsuga heterophylla Zone" to include the transition zone to the Picea-dominated forests. 39 LITERATURE CITED I 4,01(1 Beals, E. 1960. Forest bird communities in the Apostle Islands of Wisconsin. ,(% \$ Wilson Bull. 72: 156-181. AS h/""" el Cordes, L. D. 1972. An ecological study of the Sitka spruce forest on the evt west coast of Vancouver Island. Ph.D. thesis, Univ. Brit. Columbia, m PY 1101“4 If Vancouver. 454 pp. Woe-- KRA7: " ; ue/ AA( • -6 41 .4 Danner, W. R. 1955. Geology of Olympic National Park. Univ. Wash. Press, Seattle. 68 pp.

Daubenmire, R. 1952. Forest vegetation of northern Idaho and adjacent Washington, and its bearing on concepts of vegetation classification. Ecol. Monog. 22: 301-330.

Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis. Northw. Sci. 33: 43-64.

Daubenmire, R. 1969. Ecological plant geography of the Pacific Northwest. Madroho 20: 111-128.

Day, W. R. 1957. Sitka Spruce in : a Study in Forest Relationships. Forestry Comm. Bull. No. 28. 110 pp.

Fagerlund, G. O. 1954. Olympic National Park. Natural History Handbook No. 1. U.S. Dept. Int., Wash., D.C. 67 pp.

Franklin, J. F. and C. T. Dyrness. 1973. Natural Vegetation of Oregon and Washington. U.S.D.A. Forest Ser. Gen. Tech. Rep. PNW-8. 417 pp.

Fonda, R. W. 1974. Forest succession in relation to river terrace development in Olympic National Park, Washington. Ecology 55(5): 927-942.

Fowells, H. A. (comp.) 1965. Silvics of Forest Trees of the United States. U.S. Dep. Agric. Hanb. 271. 762 pp.

Heusser, C. J. 1960. Late-Pleistocene Environments of North Pacific . Amer. Geogr. Soc. Spec. Publ. No. 35. 308 pp.

Hines, W. W. 1971. Plant communities in the old-growth forests of north coastal Oregon. M.S. thesis, Oreg. State Univ., Corvallis. 146 pp.

Hitchcock, C. L. and A. Cronquist. 1973. Flora of the Pacific Northwest. Univ. Wash. Press, Seattle. 730 pp.

Jones, G. N. 1936. A Botanical Survey of the Olympic Peninsula, Washington. Univ. Wash. Press, Seattle. 286 pp.

Kirk, R. 1974. Hunters of the Whale. William Morrow and Co., New York. 160 pp.

Krajina, V. J. 1969. Ecology of forest trees in British Columbia. Ecol. West. N. Amer. 2: 1-146.

40

\J-1 Phillips, E. L. and W. R. Donaldson. 1972. Washington Climate for these fov <4‘ Counties: Clallam, Grays Harbor, Jefferson, Pacific, and Wahkiakum. Coop. Exten. Serv., Wash. State Univ., Pullman. 88 pp.

Rau, W. W. 1973. Geology of the Washington Coast between Point Grenville and the Hoh River. Wash. Dept. Nat. Res., Geol., and Earth Res. Div. Bull. No. 66. 58 pp.

Southwood, T. R. E. 1966. Ecological Methods with Particular Reference to the Study of Insect Populations. Methuen and Co. Ltd., London. 391 pp.

7 74-44c( Al 20 ai.-Y /y4 .,, iv _,,,,-?,4.157 f`-ftV0, , i,ji c , Pr 14 .4 7 ..?..? air 14. PI-4-,v4„54, 74:, v - r r r iii) " OP 51 k.....1r.r 4 4944/004i - ....). 4184 Th, - 6 44) /iv( ...... Dot

7W.0 NV kil :A If 41

VITA

Andrew Michael Kratz was born in Statesville, North Carolina on

February 1, 1950. From 1952 until 1972 he lived in Los Angeles, California.

He graduated with high honors from Granada Hills High School in January 1968 and entered Occidental College in March of that year. Andy graduated with a

B.A. in Biology from Occidental College in June 1972. He then entered the graduate program at Western Washington State College in the fall of 1972.

In May and June 1973 he worked on an environmental impact statement for the

Campus Planning Office at WWSC. From September 1973 until June 1974 he held a teaching assistantship. Andy was granted the M.S. degree in Biology in

June 1975.