Growth and Nutrient Content of Chamaecyparis Lawsoniana (A

AN ABSTRACT OF THE THESIS OF

MILTON DAVID PLOCHER for the degree of. MASTER OF SCIENCE in Botany and Plant Pathology presented on September 28,1976

Title: GROWTH AND NUTRIENT CONTENT OF CHAMAECYPARIS

LAWSONIANA SEEDLINGS FROM CONTRASTING SOILS IN

COOS COUNTY, OREGON Redacted for Privacy Abstract approved: Donald 13/.Zobel

A study of growth and nutrient content of Chamaecyparis lawsoniana (A. Murr.) Part, was undertaken to determine whether differing soil nutrient conditions have resulted in ecotypic differentiation within Coos County, Oregon.Seedlings were collected from near Coquille River Falls, from a productive forest soil; from North Spit on dune sand; from Seven Devils, an indurated pansoil.; from Rock Creek, in a roadcut through ultramafic rock,Soils from the first three sites and from Iron Mountain, another ultramatic site, were brought to Corvallis for a reciprocal transplant study in the, green-

house. Height growth was measured weekly from March li.;ovEmber, 1974, for the fourteen soil X population treatments.(Rock Creek and North Spit seedlings were not grown on the Seven Devilsell. Cliamee in seedling wet weight over the term of theexperiment was used as another growth measurement.Soil and foliage contents of N, P, Ca, Mg, and S were determined at the beginning and end of the experiment. No one population was capable of significantly. more growth on its native soil than all other transplanted populations.However, the fact that the North Spit population grew the tallest on all soils, and that the Coquille Falls population grew well but stopped growth earlier than North Spit on Coquille Falls soil. suggested real possibilities of ecotypic differentiation.Regression analysis showed that at least ninety percent of the variability in height growthwithin each population was associated with the variation in foliar K, Ca, and N

levels.Potassium was the dominant factor in the regression equations, responsible for 64 to 95 percent of the variability.Adaptation to low K and Ca in its native soil may h.ave resultedin greater growth of the North Spit population on the other soilwith higher K but low Ca contents.The difference in height between North Spit and Coquille Falls populations on Coquille Falls soil appeared to use due to.phen- ology rather than to nutrient conditions.North Spit_ may not possess a mechanism to stopgrowth in the fall, which appears to limit the Coquille Falls population.Rock Creek and e,:Te.... De:s seedlings did not exhibit the capacity to take as fun advantageof the fertile Coquille Falls soil as did the native population. Another possible expression of ecotypic differences. wasthe different relative amounts of growth amongpopulations on the different soils.The significant soil X population interaction for height growth also suggested that innate differences exist among populations in the amount of growth possible on the varioussoils. The argument that foliar nutrient content of the populations onthe different soils was innately different was supported by thedifferent rankings of population foliar nutrient content across thesoils as well as some soil X population interactionswhich were determined significant by analysis of variance. Growth and Nutrient Content of Chamaeeypar is lawsoniana Seedlings from Contrasting Soils in Coos County, Oregon

by Milton David Plocher

A THESIS submitted to Oregon State University

in partial fulfillment of the requirements for the degree of Master of Science Completed September 28, l97b

C omme nce merit June 1977 APPROVED:

Redacted for Privacy Associate Professor of Botany in charge of major

Redacted for Privacy Head of Department of Botany and Plant Pathology

Redacted for Privacy

Dean of Graduate School

Date thesis is presentedSeptember 28, 197E Typed by Susie Kozlik for Milton David Plocher ACKNOWLEDGEMENTS

1 would like to thank Dr. Donald Zobel for his patient support, thoughts, and encouragement throughout this study.Thanks also to Dr. David Moore of the Oregon State University Soils Department for answering my often foolish questions. The number of friends who helped when two hands were not enough is tremendous, but special thanks to Carlyn Swan, Carol Jefferson, Ron Southard, Alan Davenport, and Bruce Nicholson for the ir ass istance. This study was supported by a grant from the National Science

Foundation. With continuing thanks for the love and support of my mother and family I would like to dedicate this thesis to my father and my God. TABLE OF CONTENTS

Chapter Page

1 I. INTRODUCTION

Distribution and Environment 1 Ecotypic Differentiation 3

II STUDY AREAS 12

METHODS 21 Collection 21 Greenhouse Environment 22 Growth Measurement and Analysis 24 Soil Analysis 26 Foliage Characteristics and Analysis 26

IV RESULTS 29 Height Growth 29 Change in Wet Weight 33 Soil. Nutrients 35 Foliar Nutrient Content 37 Foliage Color 41 The Relationship of Growth to Foliar Nutrition 43

DISC USSION 47 LITERATURE CITED 59 APPENDICES Appendix I 65 Appendix II 67 LIST OF TABLES

Table Page

13 1 Study Area Locations oC 2 Temperature Data, 15

17 3 Precipitation Data

4 Woody Vegetation of the Study Areas 19

5 Seedling Survival Per Treatment 23

6 Methods Used for Analysis of Soils andPlant Samples 27

7 Coefficients of Equations for Growth Curvesof Experimental Treatments 31

8 Average Height Growth by Treatment 33

9 Significance of Relationships as Determinedby Analysis of Variance, F Test 34

10 Average Change in Wet Weight byTreatment 35

11 Soil Nutrient Content 36

12 Foliar Nutrient Content 38

1.3 Regression of Height Growth on FoliarNutrient Content 44

14 Correlations of Foliar Nutrient Contentwith Height Growth and Change in Wet Weightfor Samples Grouped by Population 45 LIST OF FIGURES

Figure Page

1 Map of Study Areas 14

2 Seedling Height Growth 30

3 Seedling Height Growth Computed from Regression Equations 32 GROWTH AND NUTRIENT CONTENT OF CHAMAECYPARIS LAWSONIANA SEEDLINGS FROM CONTRASTING SOILS IN COOS COUNTY, OREGON

Io INTRODUCTION

Chamaecyparis lawsoniana (A. Murr.) Parl. (Port Orford Cedar) has been studied for its economic usage but no extensive ecological research on it has been undertaken.Whittaker's (1960) community study of the Siskiyou Mountains and Youngberg's (1958) seedling nutrition study are the only works in which Chamaecyparis lawsoniana plays a major role. This study was part of a larger investigation of autecology and synecology of C. lawsoniana in Oregon and California, and of C. taiwanensis and C. formosensis in Taiwan.The purpose of this part of the study was to investigate soil and plant nutrient relationships of C. lawsoniana.The possibility of ecotypic differentiation in C. lawsoniana in response to edaphic conditions was also under investigation.

Distribution and Environment

Chamaecyparis lawsoniana grows abundantly in some areas but has a limited range from coastal Douglas County, Oregon to the Mad River in California, and inland over the crest of the Coast and Siskiyou Mountain Ranges (Fowells, 1965; Glenn M. Hawk, personal communication).Disjunct populations are found further inland in the upper 2 Trinity and Sacramento River drainages in California (Griffin and Critchfield, 1972).It occurs from sea level to above 1700 m. The climate throughout the range is tempered by the maritime influence.The majority of the precipitation falls from October through March with annual averages of 160 to 190 cm along the coast and up to 250 cm on the slopes of the Coast Range.Snowfall is rare on the coast, averaging only 5 cm per year but may exceed250 cm at higher elevations (Fowells, 1965). Thermograph data from 1975 (D. B. Zobel, unpublished) indicated that daytime temperatures within forests over the range average - 1 to 6 C in January and 15 to 17C in July.Average maxima for January are 0 to 7 C and for July 21 to 24 C. Minimum averages are - 3 to 3 C for January and8 to 11 C for July.Temperatures rarely exceed 40 C or drop below -10 C.The growing season within the range varies from 170 to 250 frost free days (Zobel,unpublished). Chamaecypar is lawsoniana grows well on many different soils within its range.Although it is most abundant on the medium textured coastal terrace soils and the clay loam and sandy loam soils of the Coast Range, C. lawsoniana also grows on swampy sites and dry rocky areas (Fowells, 1965).In the Siskiyous on soils of diorite, gabbro, and serpentine C. lawsoniana is restricted to the more mesicravines, sheltered slopes and some open northerly slopes (Whittaker,1960). In Whittaker's (1960) study area, C. lawsoniana is dominant on 3 serpentine sites, shares dominance with Pseudotsuga menziesii and Pinus monticola on diorite, and is a less abundant codominant with Pseudotsuga on gabbro. Chamaecyparis lawsoniana grows in all physiographic locations on the western slopes and sometimes over the ridgesof the Coast Range in Coos and northern Curry County, Oregon.Toward the south and interior of the range it is found increasingly on serpentine and related ultramafic soils (Griffin and Critchfield, 1972), becoming more restricted to sites with year-round seepage (Glenn M. Hawk, personal communication).

Ecotypic Differentiation

One of the objectives of this study was to determine whether C. lawsoniana exhibited ecotypic differentiation due to edaphic conditions. Turesson (1922) defined an ecotype as "an ecological unit to cover the product arising as a result of the genotypical response of the ecospe- cies to particular habitat." Clausen, Keck, and Hiesey(1940), while investigating different species complexes across California, found altitudinal and latitudinal ecotypes exhibiting different plantheights and flowering times within many species complexes, e.g.Potentilla gracilis, P. drummondii, P. breweri, Horkelia fusca,Achillea borealis, A. lanulosa, and A. millefolium. Many furtherexamples 4 are now available, including some with tree species.Elevation and moisture differences produced localized ecotypic variation in western white pine (Squillace and Bingham, 1958).In a seed germination study Squillace and Bingham (1958) found that progenies of trees from moist low elevation sites grew more rapidly on good planting sites than those from dry slope, higher elevation sites.Higher elevation seedlings grew more rapidly than low elevation sources when outplanted at higher elevations.On a larger geographic scale conifers exhibit differentiation in seedling vigor, chemical composition, and dry matter content at autumnal hardening (Langlet, 1971).Langlet reviews ecotypic differentiation from a genecological viewpoint covering geographic and climatic variations in herbaceous and woody species, including forest trees. Edaphic factors that lead to ecotypic differentiation are water relations of the substratum and the physical and chemical character of the soils and soil materials (Mason, 1946b).In some situations plant competition may be a factor in variation (Mason ,1946b). Competition or release from competition was shown to be a factor in serpentine endemics and ecotypes (Kruckeberg, 1954) and also in non-serpentine situations as in. Hall's (1955) comparison of juniper populations on Ozark glades and old fields.Within the Cupressaceae, Musselman, Lester, and Adams (1975) determined that Thuja occidentalis populations from wet lowlands and relatively dry uplands 5 showed differences in root length, photosynthesis per unit leaf area, and respiration rate between populations not overcomeby gene flow. Habeck (1958), working with similar upland andlowland T. occidentalis populations, noted significant differences inpercentage of seed germination and greater plasticity in root systemsof upland seedlings. This study investigated relations of soil and plantnutrients with growth in C. lawsoniana.Soil chemical conditions have been linked with many edaphic ecotypes.Gain (1974), using reciprocal cultivation of Vernonia patula, showed that the erect annualpopulation grew best on base-rich, neutral, loamy soil.The decumbent perennial occurred in open grassland on base-poor, acidic,sandy soil.Lime- stone and other calcareous substrateshave also caused ecotypic variation.Kapoor and Ramakrishnan (1974) and Cooper(1976) found ecotypes of Echinochloa colonum, andFestuca ovina and Briza media respectively on limestone soils.Cooper (1976) suspected differences in populations on pure CaCOlimestone and nearby dolomitic lime- 3 stone and conducted experiments todetermine response to both Ca and Mg.The dolomitic population of Festuca ovinashowed reduced root dry weight with increased Caconcentration.Magnesium did not affect Festuca ovina but shoot and rootyields of the limestone population of Briza media were significantlyreduced as Mg concentration was increased.The dolomitic population of B. media wasaffected 6 only at the highest Mg level (Cooper, 1976).Kapoor and Rarnakrishnan (1974) found that calcareous populations of Echinochloa co o:Turn showed greater selectivity inuptake of nutrients.The non- calcareous population was successful or its soil due to better efficiency at obtaining Ca at low concentrations in the soil.Vino- gradov (1949) showed that Pinus sylvestris in southcentral Russia adapted to chalk and limestone soils.Attempts were made to germin- ate seeds from sandy river terraces on adjacentchalk and limestone soils and to grow seed from the limestone on the sandysoil.No river terrace seeds germinated on the limestone soil andthe limestone seeds that germinated on the sandy soil did notsurvive. No limestone populations of C. lawsoniana areknown but ultramafic populations are common. Ultramafic soils areknown for endemic species (Mason, 1946a, 1946b; Tadros,1957).Ecotypes also abound on serpentine and neighboring soils.Kruckeberg (1967) showed by transplant studies that ultramafic ecotypes wereclearly present in herbaceous perennials: Achilleamillefolium, Fragaria virginiana, Pru.nella vulgaris and Rurne:5.. acetosella.Spisaea douglasii var. menziesii and Gaultheria shallonshowed slight ecotypic response and a delayed response wasnoted in Pinus contorta. Differences in growth for Pinus contorta werenoticeable in the outdoor soil frames on ultramafic soil 38months after germination. 7 Robinson, Edgington and Byers (1935) concluded that physical characteristics of serpentine soils did not render them unfavorable for plant growth. Gordon and Lipman (1926) originally proposed high pH and deficiency of N and P as the cause of serpentine infertility but Walker (1954) showed that, since fertilizer did not alleviate the problem, N and P deficiencies were not the entire answer. He proposed that infertility of serpentine soils was caused by:1. deficiency in N and P, 2. molybdenum deficiency, 3. soil alkalinity, 4. toxicity of nickel, chromium, and other heavy metals, and 5. status of Ca and Mg in the soil.Walker (1948) supports the possibility of Mo deficiency but Millikan (1948) points out that the apparent Mo deficiency may really be a Fe deficiency, with unavailable Fe made available with increased Mo. Millikan (1948) also points out that Mo deficiency is not evident in native plants.Alkalinity cannot cause infertility in ultramafic soils ofsouthwest Oregon, because they are not alkaline. The toxicity of Ni, Cr, and other heavy metals in serpentine soils has been investigated by Spence (1957), Soane andSaunder (1959), Spence and Millar (1963), Proctor (1971a, 1971b), White (1971) and Lee, Brooks, Reeves, and Bosewell (1975).Spence and Millar (1963) found N and P to be deficient but no toxicitydue to Ni. Proctor (1971a) confirmed Spence and Millar's(1963) conclusions on N and P but showed that Niis toxic, but not acutely toxic 8

(Proctor, 1971b).White (1971) showed that Ni and Cr toxicity may be a major factor on serpentine soils of the Siskiyou Mountains, southeast of the present study area. Serpentine soils consistently have low Ca:Mg ratios, often less than 1.0.Walker (1954) concluded that low Ca level was the main reason for intolerance on serpentine soils.Loew and May (1901) considered Mg toxic and concluded that a Ca:Mg ratio of 1.0 was necessary for good plant growth.Vlamis and Jenny (1948) and Vlamis (1949) supported the idea that a Ca deficiency is responsible for infertility.Growth of ecotypic populations of Agropyron spicatum in nutrient solution showed that serpentine populations requiredless Ca than non-serpentine populations (Main, 1974).In contrast Proctor (1970, 1971b), concludes that Mg toxicity is the main reason for poor plant growth on serpentine and that serpentine endemics and ecotypes may be able to exclude Mg and thus appeartolerant.Main's (1974) experiments pointed out that it appears that higher external Mg is needed to maintain sufficient internal Mg in serpentine than non- serpentine plants.Jenkins on (1974) has shown differential growth, nutrient uptake and mycorrhizal formation inreciprocal transplants of Pinus ponderosa from a range of soils: highlyfertile granite test soil, and seven infertile ultramafic soils chosen forthe range of chemical and parent rock differences.The growth differences could be related to differences in Ca uptake, as progenieswith better 9 growth had higher Ca concentrations in roots, tops, or both.Differ- ences in growth between the seedlings on different ultramafic soils could be explained by Ca uptake ability of seedlings and soil Ca availability.Further investigations are underway to determine whether ecotypes from these different soils are present within ponderosa pine. Within the Cupressaceae, Cupressus macnabiana and C. sargentii are endemic to serpentine with disjunct populations from Mendocino south in the California Coast Range and scattered in the northern California Coast Ranges and the foothills of the Sierra

Nevada (McMillan, 1956). The Black lock soil series has also been the site of differentiation in conifers.The pygmy forests of the Black lock soils along the Mendocino coast of California are noted for Pinus bolanderi and Cupressus pygmaea, adapted to the highly acid, pH 3.5 to 4.0, condition (Jenny, Ark ley, and Schultz, 1969).Critchfield (1957), treating P. bolanderi as a subspecies of P. contorta, noted that it could be distinguished from other coastal P. contorta by its serotin- o s cones and narrower leaves. McMillan (1956), working with both P. bolanderi and Cupressus pygmaea showed that the coastal plain(from more highly acid soils) and the Anchor Bay (from less acid soils) populations of C. pygmaea exhibited differentiation in growth and seed color. When grown on 10 acid soil the coastal plain population grew more than the Anchor Bay

population.On serpentine soil the Anchor Bay population grew taller than the coastal plain seedlings.McMillan (1956) also tested response to altered acid soils.When N was added in increasing amounts P. bolanderi increased in growth and C. pygmaea was inhibited.Both grew well when N and P were added but notwith just P. Jenny et al. (1969) noted that the indurated pan of the Blacklock soil is an additional limit to nutrient availability.When roots pene- trate the pan, where cracked, the plants are no longer dwarfed.The water availability also affects growth.Standing water is common in the winter with the other extreme, little available water in the surface horizons, prevalent in the summer months.Water, nutrients, and pH may combine to limit enzymatic reactions or metabolicpath- ways in the plants, causing the dwarfing(Jenny et al. ,1969). Ecotypic differentiation on coastal sand has not been extensively

studied.Kumler (1969) determined that Senecio s "lvaticus had both a Coast Range mountain race and acoastal sand race.Senecio from Douglas-fir clear cuts required additional N and P to survive in the rosette stage beyond 64 days on infertilecoastal sand while the coastal sand race bolted, flowered, and set seedwithout supplement. When grown in potting mixture both racesbolted and flowered in five months. 11

During this study, growth and foliar N, P, K, Ca, Mg, and S were measured in C. lawsoniana seedlings grown on fourcontrasting soils, including an ultramafic soil, a Black lock soil, and coastal dune sand.Heavy metals in the soils and foliage were not examined. 12

II.STUDY AREAS

Five areas within or just outside Coos County, Oregon were used during this study (Table 1, Figure 1).The sample sites are distributed from near sea level at North Spit to 850 m near Iron

Mountain. The sites were chosen for their contrasting soils.North Spit is stabilized coastal dune sand covered with 20 cm of litter (Beaulieu and Hughes, 1975).Because of the litter layer and the depression in the landscape, the sample site probably retains moisture longer than the surrounding dunes.Chamaecyparis lawsoniana at Seven Devils is growing above an indurated pan of the Black lock soil series.The soil at the Coquille Falls site is a sandy loam derived from the Tyee Formation sandstone and siltstone (Baldwin, 1974).Rock Creek study area is a roadcut with both gabbro and serpentinite (A. R. Niem, Department of Geology, Oregon State University,personal communication).The Iron Mountain soil is derived from serpentinite (Dott, 1971). All sites have a climate that is influenced by their proximity to the Pacific Ocean. Most precipitation isreceived from October through May.Thermograph data were available for 1975 on the Iron Mountain site and the Coquille Falls site (Table2).Temperatures from the Coos County Forest site were takenapproximately 4 km 13

Table 1.Study Area Locations.

Study ElevationTownship/Range Latitude (Abbreviation) (meters) Longitude

Coquille Falls 515 SE1/4, NE1/4, Section 20Lat. 42°43'N. (CF) T 33 S, R11 W Long. 124°1'W.

Rock Creek 660 SE1/4, NE1/4 Section 13 Lat. 42°44'N. (RC) T 33 S, R 12 W Long. 124°4'W.

Iron Mountain 850 SE1/4, SE1/4, Section 34Lat. 42°40'N. (IM) T 33 S, R 12 W Long. 124°7'W.

North Spit 10 NE1/4, NW1/4, Section4 Lat. 43°26'N. (NS) T 25 S, R 13 W Long. 1240 15'W.

Seven Devils 80 SE 1/4, SE1/4, Section 27Lat. 43°17'N. (SD) T 26 S, R 14 W Long. 124° 20'W. 14

124°1N

1 To mi.

io Km.

NORTH SPIT

COOS B Y

SEVEN DEVILS

COOS COUNTY

44° N

A POWERS

POPULATION SOIL

ra"COQUILLE FALLS ROCK CREEK

RON MOUNTAIN Figure 1.Map of Study Areas. Table 2.Temperature Data, °C.(Johnsgard, 1963, and D. B. Zobel, personal communication). NB = North Bend and North Bend CAA AP, MF = Marshfield, PW = Powers U. S. Weather Bureau Station, IL = Illahe (preceding from Johnsgard), CR = Coquille RiverFalls, AG = A guess Pass, CC = Coos County Forest (from Zobel). CR, AG and CC data were taken at 1 m under naturalChamaecyparis stands.

Average Maximum Average Minimum

Calendar Period NB MF PW IL CR AG CC NB MF PW IL CR AG CC

January 11.2 10.9 11.6 7.7 4.4 7.8 3.3 2.6 1.1 1.4 -0.4 3.2 February 12.1 13.9 13.2 11.2 6.0 1.2 8.4 4.2 3.1 2.3 2.8 2.2 -2.0 3.4 March 13.0 13.5 14.9 14.7 6. 9 9.0 4.8 3.5 2. 8 2. 9 1. 9 3, 5 April 14. 3 14.7 17. 4 19.4 7. 7 6. 7 9. 3 6. 0 4. 4 4. 4 4. 7 1. 3 -O. 4 3. 4 May 16.2 16.6 19.9 23. 1 16.2 17.4 13.5 7.9 6.4 6.6 7.0 5. 2 3.6 5.9 June 18.0 18.6 22.7 21.5 19.7 20.6 15.5 9.8 8.6 8.4 9.1 6.8 5.9 8.6 July 19.2 20.1 25.5 31.4 22.0 24.1 17.2 11.0 10.2 10.0 11.3 8.9 7.8 9.6 August 19.7 20.6 26.2 31.4 17.7 19.3 15.6 11.0 10.3 9.5 10.7 6.5 5.6 8.9 September 19.2 20.2 25.2 28.4 22.5 25.3 16.3 9.9 8.6 7.6 9.1 9.0 9.5 8.0 October 17.3 18.3 20.5 19.2 9.7 9.5 12.5 8.6 5.2 6.0 7.3 3.8 2.8 7.1 November 14.3 14.4 15.4 11.8 5.8 3.2 9.2 5.9 4.7 3.4 4.4 1.0 -1.7 3.9 December 11.8 11.4 12.4 9. 1 9. 1 4.4 9.8 4.8 2.7 2.6 3. 4 4. 1 -0.3 4. 7 Annual 15.6 15.9 18.7 19.5 - 7. 3 5. 9 5.4 6.2 (1931-(1902- (1931-(1939- (1975) (1975) (1975) (1931-(1902-(1931-(1939- (1975) (1975) (1975) 1955) 1930) 1955) 1952) 1955) 1930) 1955) 1952) 16 inland from the Seven Devils site and at approximately the same elevation.No current data were available for the North Spit site, although the North Bend weather station (Johnsgard, 1963) is just across the bay, and should represent the site well.Both North Spit and Seven Devils have summer temperatures moderated by coastal fog.The weather station at Powers is the closest to both the Rock Creek and Coquille Falls areas, although it is still 27 km away and at lower elevation.The Iron Mountain study area is on the divide between the Rogue and Coquille River drainages and is not tempered as much by the ocean's influence as the other areas.The Illahe weather station is closest to Iron Mountain but is 800 m lower and 7 km away, and thus probably only roughly comparable. Rainfall data are unavailable for all sites, although North Bend and Marshfield probably well approximate precipitation at North Spit and Seven Devils, respectively (Table 3).Precipitation is probably slightly higher at the three inland sites than down in the valleys at Powers and Illahe.Snowfall is a regular winter occurrence at Rock Creek and Coquille Falls but there is no significant accumulation. Snow remained on the ground at the Iron Mountain site from January to April in 1975. The vegetation of the study areas can be divided into three types on the basis of the major conifers.North Spit and Seven Devils are characterized by C. lawsoniana, Picea sitchensis, Pinus Table 3.Precipitation Data (Johnsgard, 1963). Precipitation (cm) Calendar Period North Bend Marshfield Powers Illahe

January 25. 1 27. 9 26. 7 36. 6 February 20. 3 22. 9 19. 6 30. 7 March 19. 3 19. 1 20. 3 24. 4 April 10. 4 12. 5 10. 4 11, 4 May 6. 8 8. 1 7. 4 9.1 June 4. 6 4. 3 4. 3 4. 3 July 1. 0 1. 0 0.8 1. 3 August 1. 5 1.0 1.0 1. 3 September 4. 3 6. 1 3. 0 4. 3 October 13. 7 10. 7 12. 7 18. 8 November 22. 9 25. 4 21. 6 33. 0 December 28.2 24. 9 28. 7 42. 7 Annual 158.2 163. 8 156.5 217. 9 (1931-1955) (1902-1930) (1931-1955) (1939-1952) 18 contorta, and Tsuga heterophylla, placing these sites in the Picea sitchensis zone of Franklin and Dyrness (1973).Chamaecyparis lawsoniana is codominant with Pseudotsuga menziesii on the Coquille Falls site.Tsuga heterophylla is the climax species in this section of the Coast Range. At Rock Creek and Iron Mountain Pinus monticola, P. lambertiana, Pseudotsuga menziesii, Arbutus menziesii and Calocedrus decurrens dominate the overstory along with C. lawsoniana.Pinus jeffreyi, a serpentine indicator, occurs near the Iron Mountain area.Chamaecyparis lawsoniana grows larger than either Pseudotsuga or Pinus monticola on these two latter sites. Rock Creek and Iron Mountain are characteristic of the mixed ever- green or Pseudotsuga-sclerophyll zone as it occurs on serpentine soils (Franklin and Dyrness, 1973). All sample sites have understory vegetation dominated by erica- ceous shrubs (Table 4).The greater diversity in the understory and the abundance of sclerophyllous species tend to emphasize the fact that Rock Creek and Iron Mountain represent a different vegetation zone than North Spit and Seven Devils, orCoquille Falls. Coquille Falls and Iron Mountain study areas are included in a community study of C. lawsoniana (Glenn M. Hawk, unpublished). Coquille Falls is in Hawk's C. lawsoniana-Tsugaheterophylla/ Polystichum munitum-Oxalis oregana community (near plot 33). Iron Mountain is characteristic of the C. lawsoniana-Lithocarpus 19

Table 4.Woody Vegetation of the Study Areas (Glenn M, Hawk, unpublished).

Study Area Tree Species Shrub Species

Coquille Falls Chamaecyparis lawsoniana Berberis nervosa Pseudotsuga menziesii Gaultheria shallon Tsuga heterophylla Lithocarpus densiflora Vaccinium ovatum

Iron Mountain Chamaecyparis lawsoniana Amelanchier pallida Arbutus menziesii Arctostaphylos columbiana Calocedrus decurrens Berberis nervosa Pinus monticola Castanopsis chrysolepis Pseudotsuga menziesii Ceanothus pumilus Gaultheria shallon Lithocarpus densiflora Quercus chrysolepis Q. sadleriana Q. vaccinifolia Rhamnus californica Rhododendron macrophyllum R. occidentale Rosa gymnocarpa

Rock Creek no data available but floristically similar to Iron Mountain

North Spit Chamaecyparis lawsoniana Gaultheria shallon Picea sitchensis Mvrica californica Pinus contorta Rhamnus purshiana Pseudotsuga menziesii Rhododendron rnacrophyllum Tsuga heterophylla Vaccinium ovatum

Seven Devils Chamaecyparis lawsoniana Gaultheria shallon Picea sitchensis Ledum glandulosum var. columbianum Tsuga heterophylla Menziesia ferruginea Myrica californica Rhamnus purshiana Vaccinium ovatum V. parvifolium 20 densiflora community (plot 38).North Spit and Seven Devils, while not included in a major community, are represented as Hawk's plots 68 and 70, respectively. 21

III. METHODS

Collection

Soils were collected in August 1973 from the first 15 cm below the litter at Coquille Falls and Iron Mountain.North Spit soil was taken from the second layer of sand below the litter, the first 10 cm being a leached horizon.The soil collected at Seven Devils was a combination of the Al and A2 horizons above the indurated pan.All soils were sifted through a 1.25 cm screen before use in the seedling study. Cuttings were taken from small C. lawsoniana trees in five areas in June 1973.Only five percent of the cuttings rooted after four months. A second attempt to root cuttings was made in October

1973.After six months the cuttings, though treated with fungicide and placed in a mist bench, failed to root.Since similar rooting methods had worked previously the problem could have been 1.too short a time for rooting to occur or, 2.drying of foliage in transport to Corvallis resulting in dead foliage unable to root.Hartmann and Kester (1975) state that more easily rooted softwoods, including C. lawsoniana, may take six months to a year to root.The low percentage of surviving cuttings probably resulted from acombination of these factors. 22 Sample sites were then examined for seedlings to be used in place of rooted cuttings.Seedlings were abundant at North Spit, Seven Devils, and Coquille Falls but few were found on the ultramafic soil of Iron Mountain. Rock Creek was selected as an alternate site with an ample number of seedlings growing in an ultramafic soil similar to that of the Iron Mountain site.Seedlings were collected in the open on disturbed areas (roadsides, skid roads, androadcuts). In March 1974 seedlings from the four study areas were taken to Corvallis and transplanted into soils from North Spit,Seven Devils, Coquille Falls and Iron Mountain. Seedlings were weighed, measured for total length and stem length and paired randomly for transplanting.The seedlings were planted in 2.2 liter fluted metal pots with approximately 2 liters of soil.Thirty-four seedlings in seventeen pots were planted for each of fourteen treatments; survival was good in most cases (Table5).

Greenhouse Environment

The environment of the greenhouse was set to duplicate as closely as possible that of Coquille Falls study area in July.The desired temperatures were 14 C night and 22 C day;however, the average recorded by thehygrothermograph was 15.5 C minimum at night and 25 C maximum during the day throughoutthe experiment. 23

Table 5.Seedling Survival Per Treatment.

Soil Seedling Populations Sites Coquille Falls Rock Creek North Spit Seven Devils

Coquille Falls 2 in 15 pots 2 in 11 pots 2 in 15 pots 2 in 16 pots 1 in 2 pots 1 in 3 pots 1 in 2 pots 1 in 2 pots 32 seedlings 25 seedlings 32 seedlings 34 seedlings

Iron Mountain 2 in 16 pots 2 in 10 pots 2 in 8 pots 2 in 16 pots 1 in 1 pot 1 in 6 pots 1 in 9 pots 1 in 1 pot 33 seedlings 26 seedlings 25 seedlings 33 seedlings

North Spit 2 in 14 pots 2 in 14 pots 2 in 17 pots 2 in 15 pots 1 in 3 pots 1 in 3 pots 1 in 0 pots 1 in 2 pots 31 seedlings 31 seedlings 34 seedlings 32 seedlings

Seven Devils 2 in 9 pots none none 2 in 7 pots 1 in 6 pots planted planted 1 in 6 pots 24 seedlings 20 seedlings 24

A hygrothermograph run concurrently during July, 1974, on the Coquille Falls site had 15 C average minima and 25 C average maxima.Humidity varied from 40 to 80 percent at Coquille Falls and 50 to 90 percent in the greenhouse. Plants were watered with distilled water to limit the nutrient variability to that within the soils and the seedlings.Water stress was eliminated by frequent waterings, two to three times weekly at the start, and once a week after the middle of July.All pots were surface flooded except for the Seven Devils soil.Compaction of the Seven Devils soil caused runoff of surface water which was alleviated by sub-irrigation.Natural light was supplemented with Gro-Lux flourescent lighting (11 40-Watt bulbs/m of bench), set for 15 hour days.Pots were randomly assigned to an original position and then rotated randomly each week to limit the effects of position in the room.

Growth Measurement and Analysis

Height growth was chosen as the most readily measurable check for differences in seedling growth (Minckler, 1950). A spot of acrylic paint was placed above the soil level on the stem of each seedling, to serve as a permanent reference point.Measurements of total height from the spot to the tip of the terminal or longest branch (if the terminal had died) were taken at weekly intervals. 25 At the end of the experiment (November, 1974), after 34 weeks of growth, all plants were removed from the pots, weighed, and measured for total length and root length.Seedlings were dried and foliage dry weight was determined. Height growth was analyzed comparing the twelve treatments on Coquille Falls, Iron Mountain, andNorth Spit soils.The Seven Devils soil was not included because all seedling populations were not replicated on it.The seedlings were randomly grouped into four groups of four seedlings withineach treatment to average the initial size differences.Sixteen seedlings were used because that was the largest easily grouped number available on each treatment,excluding transplant deaths and single seedling pots.The average cumulative growth of the four seedlings in each group wasdetermined for each sample date. A regression equation was computedfor height growth as a function of time.The model regression equation was y = a + bx + cx2,where x = number of two week periods in the greenhouse.The coefficients from the forty-eight groups(four groups for each of twelve treatments) were compared usinganalysis of variance to determine whether there was anysignificant variability of the coefficients of the he ight growth equationsassociated with treatment combinations. The seedlings from both Coquille Fails andSeven Devils sites which were grown on the Seven Devils soil werecombined into three 26 groups of four.The regression equation for each group was then computed.The resulting coefficients were compared with the coefficients of the twelve other treatments using theleast significant difference, using t for a probability level of .05, todetermine if their growth was similar.

Soil Analysis

Soils collected for use in the greenhouse wereanalyzed before and after seedling growth. Analysis included pH,total nitrogen (N), phosphorous (P), potassium (K), calcium (Ca), magnesium(Mg), and organic matter.This analysis was performed by the Soil Testing Laboratory, Oregon State University(for methods see Table

6).Particle size determinations were carried outfollowing Schroeder (1953).

Fo lia e Characteristics and Anal sis

Color measurements, using the Munsell ColorCharts for Plant Tissues, were taken at time of transplant(March), in late April to early May, in August and at the end ofthe experiment in

November. Seedling foliage nutrient contents weredetermined for 1) 15 to 20 additional seedlings collected at the studysites in March 1974, and 2) for forty-one composite samples atthe end of the experiment, 27

Table 6.Methods Used for Analysis of Soils and PlantSamples. Soils Testing Laboratory, Oregon StateUniversity, Corvallis, Oregon.

Soil Anal ysis Methods For Total Nitrogen - RegularMicro-Kjeldahl Method (Bremmer, 1965) For Extractable Phosphorous - DiluteAcid-Flouride Method (Bray and Kurtz, 1945) For Extractable Calcium, Magnesium,and Potassium - Ammonium Acetate Method (Peech, et al. ,1947) For Organic Matter - Walk ley-BlackTitration Method (Walk ley and Black, 1934) For pH - use 1:2 soil to solution ratioand Glass Electrode pH Meter (Jackson; 1958)

Plant Analysis Methods For Total Nitrogen - RegularMicro-Kjeldahl Method (ARS Labora- tory, USDA, Corvallis) For Total Phosphorous - Reagentsused: Ammonium vandate, Ammonium Molybdate.Procedure available Soils Testing Laboratory O. S. U.Corvallis, Oregon For Calcium and Magnesium - Reagent:Strontium solution 1500 ppm. Procedure availableSoil Testing Laboratory, O. S. U. ,Corvallis, Oreg on For Potassium- Reagent:.1 N NaC1 solution.Procedure available Soils Testing Laboratory, 0. S. U. , Corvallis , Oregon Calcium, Magnesium and Potassium are determined usingatomic absorption spectrophotometry For Sulfur- Reagent:bariumchloride.Procedure available Soils Testing Laboratory, 0. S. U. , Corvallis, Oregon 28 each including one third of the seedlings from each treatment (except that each sample for Seven Devils on Seven Devils soil included one half the seedlings).Foliage included those leaves that could be stripped from the seedlings and the woody tissues that were removed with the leaves.Samples were analyzed for total nitrogen (N), phosphorous (P), potassium (K), sulfur (5), calcium (Ca), and magnesium (Mg).All analysis was done by the Soils Testing Labora- tory, Oregon State University (for methods see Table6). Nutrient contents were statistically analyzed using multiple regression analysis with average height growth and averagechange in total seedling wet weight within each composite seedlingsample as dependent variables and nutrient contents of the compositesamples as independent variables (Mader andHowarth, 1968).Regressions were calculated 1) for each population onall soils, and 2) for all populations on each soil type.Analysis of variance of growth and nutrient contents of the composite samples were donefor all treatments on the Coquille Falls, Iron Mountain, andNorth Spit soils. 29

IV. RESULTS

Height Growth

There was significantly more height growth of populations on the Coquille Falls soil than on the other three, and more on the Iron Mountain soil than on the other two soils (Figure 2).The differences among populations on North Spit and SevenDevils soils were not significant (Table 7). The coefficients for slope from the growth curve equations (b, Table 7) were compared using the least significant differencewith t = .05.On Coquille Falls soil, slopes for the Coquille Falls and North Spit populations were significantly greaterthan those of Rock Creek and Seven Devils seedlings.Curvature of the growth curves of the North Spit and Coquille Falls populations aresignificantly different (Table 7, as seen in Figures 2 and 3) with growthof the Coquille Falls population leveling off sooner. There is significant growth of all populations on the IronMoun- tain soil; here the North Spit population grewsignificantly more rapidly than the others.The difference of North Spit from the others is evident in both slope and curvature(Figure 3, Table 7). No significant difference in growth occurredbetween populations on the North Spit soil or onthe Seven Devils soil. 30

North Spit ,Coninile Falls Coquille Falls Coquille Falls Falls 250 e 0 Rock Creek Coquille ft-Et Seven Devils/CoquilleFalls North Spit, Iron Mountain Coquille Falls Iron Mountain Rock Creek /Iron Mountain Seven Devils iron Mountain North Spit 'North Spit 200 Coquille Falls/ North Spit Rock Creek/North Spit Seven Devil s 'North Spit Coquille Falls Seven Devils Seven Devils Seven Devils ,-0

150

3 a 100

I I ri 1 r---Tr-TTrr-r-T-- 8 10 12 14 1618 20 22 24 2628 30 32 34 Timeweeks Figure 2.Seedling Height. Growth. 31

Table 7.Coefficients of Equations for Growth Curves of Experi- mental Treatments, y = a + b + cx2 where y = increase in plant height in cm and x = time in number of two week periods.In the text coefficient b is referred to as the "slope" and c as the "curvature." Treatment Coefficients (populations oil) a c

Coquille Falls/Coquille Falls -40.16 31.66 -1.05 Rock Creek/Coquille Falls -35.27 23.80 -0.69 North Spit/Coquille Falls -57.79 30.92 -0.78 Seven Devils/Coquille Falls -47.39 23.07 -0.56 Coquille Falls/Iron Mountain -15.06 10.23 -0.16 Rock Creek/Iron Mountain -16.63 9.13 -0.16 North Spit/Iron. Mountain -29.30 17.52 -0.32 Seven Devils/Iron Mountain -17.45 8.03 -0.01 Coquille Falls/North Spit 5.86 0.69 0.08 Rock Creek/North Spit 1.11 0.46 0.03 North Spit/North Spit 5.99 0.73 0.10 Seven Devils/North Spit 1.62 -0.46 0.13 Coquille Falls/Seven Devils 2.44 0.84 0.02 Seven Devils/Seven Devils -0.36 0.80 0.06 6.915 4.159 0.183 L. S. D.(.05) 32. North Spit/Coquille Falls 250 .._Coquille Falls/Coquille Falls Rock Creek/Coquille Falls _Seven Devils/Coquille Falls North Spit /Iron Mountain

to- Coquille Falls/ Iron Mountain Rock Creek/Iron Mountain 200 Seven Devils/Iron Mountain A A North Spit/North Spit Coquille Falls/North Spit Rock Creek/ North Spit Seven Devils/North Spit _Coquille Falls/Seven Devils 'Seven Devils/Seven Devils

3 100

......

I I III T--rrMmr- 10 12 14 16 18 20 22 24 26 28 30 3234 Time - weeks Figure 3.Seedling Height. Growth Computedfrom Regression Equations. 33

Maximum height growth was ranked consistently foreach population across soils: Coquille Falls > Iron Mountain >North

Spit > Seven Devils (Table 8).North Spit seedlings grew the most, and Rock Creek the least, on all soils (Table8),

Table 8.Average Height Growth by Treatment (in mm). Populations Soils Coquille Rock North Seven Falls Creek Spit Devils

Coquille Falls 199 168.5 233.5 174 Iron Mountain 105 87 153 107 North Spit 41 16 42.5 31

Seven Devils 21 - 25

Analysis of variance of height growth for all treatmentsshowed that it varied significantly with soils and withpopulations (Table 9). Height growth was also significantly correlated atthe five percent level with soil X population interaction.

Change in Wet Weight

Change in wet weight was highly correlatedwith height growth.

The r2between weight change and height growthfor various populations ranged from .93 to .96.Change in total plant wet weight was highly significantly correlated with soilsand significantly related to 34

Table 9.Significance of Relationships as Determined by Analysis of Variance, F Test.NS = not significant,* = significant at 5 percent level, ** = significant at the 1 percent level. Data from Seven Devils soils are not included. Source of Variation Variable Soil Population Soil X Population

Height

A Wet Weight NS (just)

Nitrogen NS

Phosphorous NS Potassium Calcium NS Magnesium

Sulfur NS NS NS 35 soil X population interaction but was not correlated significantly with populations (Table 9).The change in wet weight was not ranked consistently for each population across the different soils (Table 10). The relationship between height and wet weight was not the same across the populations.

Table 10.Average Change in Wet Weight by Treatment. Populations Soils Coquille Rock North Seven Falls Creek Spit Devils Coquille Falls 31.60 g 32.65 g 24.53 g 28.89 g Iron Mountain 6.00 g 7.74 g 13.25 g 3.72 g North Spit 1.65 g -0.24 g 1.15 g 0. 66 g Seven Devils 0.55 g - 0.55 g

Soil Nutrients

The soils were chemically quite different (Table11).The Coquille Falls soil was chosen initially to represent aproductive forest soil.Soil analysis indicated that Coquille Fallssoil was highest in total N, P, K, and Ca, with Ca:Mgratio averaging 3.7:1. The Ca:Mg ratio is presented as calculatedfrom data in both percent and milliequivalents/100 grams ofsoil, but the latter will be used in discussion to be consistent with the restof the literature. Ultramafic soil from Iron Mountain hadthe highest Mg content and had an average Ca:Mg ratio of 0.16:1.Iron Mountain soil also Table 11.Soil Nutrient Content; samplo-th:s are composites of smaller samples taken from each pot of each treatment atthe end of the e:x:periment. Initial samples are from initial field collections. Done by Soil Testing Laboratory, Oregon StateUniversity, Corvallis.* taken from Bullard, et ad. (1957).

Percent Sample 9/0 Organic (Meg/ 100g) Soil (texture) Time Population pH %N yP %K. %Ca. %Mg Matter Ca:Mg Ca Mg Ca:Mg

Coquille Falls initial 5.2 0.24 0.0040 0.0400 0.216 0.035 3.50 6.17 10.8 2.9 3.72 (sandy loam) final Coq. Falls 5.6 2.69 0.0040 0.0230 0.244 0.036 9.91 6.78 12.2 3.0 4.07 Rock Creek 5.5 2.95 0.0040 0.0270 0.232 0.035 7.99 6.63 11.6 2.9 4.00 North Spit 5.7 2.00 0.0040 0.0320 0.228 0.038 4.48 6.00 11.4 3,2 3.56 Seven Devils 5.7 1.78 0.0040 0.0290 0.224 0.036 7.52 6.22 11.2 3.0 3.73 Iron Mountain initial 6.3 0.12 0.0004 0.0072 0.032 0.119 7.30 0.27 1.6 9.9 0.16 (loam) final Coq. Falls 6.5 1.12 0.0004 0.0054 0.064 0.107 5.22 0.60 3.2 8.9 0.36 Rock Creek 6.7 1.35 0.0004 0.0044 0.064 0.168 6.08 0.38 3.2 14.0 0.23 North Spit 6.8 1.47 0.0004 0.0044 0.054 0.180 6.08 0.30 2.7 15.0 O. 18 Seven Devils 6.6 1.48 0.0004 0.0064 0.080 0.156 6.66 0.51 4.0 13.0 0.31 North Spit initial 5.1 0.03 0.0030 0.0036 0.006 0.002 1.10 3.00 0.3 0.16 1.88 0.16 4.38 (sand) final Coq. Falls 5.3 0.22 0.0027 0.0016 0.014 0.002 0.08 7.00 0.7 Rock Creek 5.3 0.22 0.0025 0.0008 0.014 0.002 0.80 7.00 0.7 0.16 4.38 North Spit 5. 4 0.16 0.0027 0.0008 0.014 0.002 0.80 7.00 0.7 0.16 4.38 Seven Devils 5.2 0.25 0.0032 0.0016 0.014 0.002 1.01 7.00 0.7 0.16 4.38 0.36 1.14 Seven Devils initial 4.2 0.09 0.0026 0.0036 0.008 0.004 4.50 2.00 0.4 0.82 1.22 (clay loam) final Coq. Falls 4.5 0.14 0.0011 0.0036 0.020 0.010 5.76 2.00 1.0 Sever Devils 4.4 0.17 0.0015 0,0044 0.014 0.011 7.25 1.27 0.7 0.89 0.79 Coastal Oregon * 0.19 0.0001 0.0105 to tc to 0.64 0.0010 0.0170 Southwestern Oregon* 0.05 0,000450.0150 3 to 1 1 2 to 4 to to to 0.09 0.0052 0.0250 rn 37 had the highest pH and lowest P content.The total N was about half that of the Coquille Falls soil. North Spit sand was the lowest in K, Ca, Mg, and organic matter contents.Phosphorous content was greater than Seven Devils and Iron Mountain but less than Coquille Falls.Total N was only one tenth the value for Coquille Falls. Soil from the Seven Devils site was the most acidic, pH 4.2 to 4.5, and was low in total N.Contents of both Ca and Mg were low and the Ca:Mg ratio was about 1:1. Organic matter was the only component analyzed that changed its relative value over the course of the experiment.Initially Iron Mountain had the most organic matter.At the end of the experiment the maximum organic matter content was found in the Coquille Falls soil. The soil from Rock Creek site was similar to Iron Mountain but slightly higher in pH, K, Ca, and Mg and lower in total N and organic matter.The Ca:Mg ratio was 0.45:1, indicating the high Mg of its ultramafic parent rock.

Foliar Nutrient Content

Results of foliar analysis on field collected seedlings differed between populations for each of the nutrients (Table 12).The Coquille Falls population was high in N and P and low in Ca. 38

Table 12.Foliar Nutrient Content.

Population/Soil %N %P %K %S %Ca %Mg Ca:Mg CF initial 1.07 0.20 0.69 0.11 0.60 0.20 3.00 CF/CF mean 1,04 0.27 1.05 0.20 0.65 0.17 3.74 RC/CF mean 1.02 0.28 1.11 0.04 0.65 0.21 3.17 NS/CF mean 1.140.32 1.28 0.09 0.58 0.19 2.98 SD/CF mean 0.92 0.24 1.15 0.1.7 0.18 0.62 3.50 RC initial 0.57 0.09 0.58 0.11 1.11 0.30 3.70 CF/IM mean 0.93 0.24 0.88 0.14 0.39 0.50 0.78 RC/IM mean 0.88 0.25 0.89 0.12 0.38 0.48 0.80 NS /IM me an 0.98 0.31 0.99 0.22 0.32 0.48 0.67 SD/IM me an 1.07 0.29 0.91 0.12 0.38 0.49 0.77 NS initial 1.22 0.21 0.84 0.19 0.54 0.25 2.16 CF/NS mean 1.02 0.33 0.85 0.19 0.59 0.25 2.40 R C/NS mean 0.67 0.19 0.63 0.08 0.68 0.29 2.37 NS/NS me an 0.97 0.35 0.91 0.26 0.52 0.32 1.63 SD/NS mean 1.23 0.33 0.68 0.24 0.25 0.68 2.74 SD initial 0.68 0.09 0.53 0.09 0.11 1.2411.27 CF/SD mean 0.93 0.15 0.83 0.24 0.70 0.17 4.11 SD/SD mean 1.13 0.14 0.84 0.16 0.22 0.71 3.23 McNabb 0.98 0.09 0.66 0.09 0.80 0.18 4.44 (on serpentine) Zobel CRF 0.89 0.13 0.59 1.20 0.21 5.71 AG 0.67 0.07 0.41 0.88 0.24 3.67 CCF 1.14 0.12 0.68 0.50 0.11 4.55 Youngberg (1958) (whole seedlings) Greeley 1.39 0.24 0.90 0,92 Wind River 0.57 0.12 0.74 0.51 Corvallis 0.55 0.12 0.72 0.53 39 Seedlings from North Spit were high in N, P, K, and S and low in Ca. The Rock Creek collection was high in Mg and Ca and low in P. Seven Devils foliage samples were high in Ca but low in Mg, P, and

K. During the experiment total N increased in the Rock Creek and Seven Devils populations on all soils (Table 12).Foliar N content decreased in the North Spit population and decreased slightly in the Coquille Falls population on each of the soils.The Coquille Falls soil produced the largest increase in total N in the Rock Creek population and the smallest decrease in the North Spit and Coquille Falls populations.The Seven Devils seedlings responded with the greatest increase on the North Spit sand. Potassium in the foliage of all populations increased on all soils.The increase in percent K for each population followed a trend across the soils similar to soil K content: CoquilleFalls > Iron Mountain > North Spit > Seven Devils. The foliar Ca content of all populations grown on the Iron Mountain soil definitely decreased (Table 12).Calcium content in the Coquille Falls foliage remained the same on North Spitsoil but increased slightly on the Coquille Falls and SevenDevils soils. North Spit foliage showed little or no change in Ca onNorth Spit or Coquille Falls soils.All treatments of the Rock Creek and Seven Devils populations resulted in decreased foliar Cawith the greatest 40 decreases occurring on Iron Mountain soil. The effect of the ultramafic soil of Iron Mountain was apparent in the Mg analysis of each population (Table 12).Foliar Mg in Coquille Falls seedlings registered only small changes on Coquille Falls, North Spit and Seven Devils soils but increased by 150 percent on the Iron Mountain soil.Magnesium in the Rock Creek population was unchanged on North Spit soil, decreased on Coquille Falls and increased by 60 percent on the Iron Mountain soil.Seedlings from North Spit decreased in Mg after growing on Coquille Falls soil and increased on North Spit sand.On Iron Mountain soil the North Spit foliar Mg content increased by 94 percent.The Seven Devils population increased in percent Mg on each soil but increases on Coquille Falls, North Spit, and Seven Devils were overshadowed by a 400 percent jump on the Iron Mountain soil. The Ca:Mg ratio for seedlings grown on Coquille Falls soil ranged from 2.98 to 3.74.Foliage grown on North Spit sand had Ca:Mg ratios of 1.63 to 2.74.The ultramafic nature of the Iron Mountain soil was reflected in the Ca:Mg ratios of 0.67 to 0.80 for populations grown on it.The Seven Devils soil produced the highest Ca:Mg, 4.11, in the Coquille Falls population and a ratio of 3.23 in Seven Devils seedlings.The North Spit population had the lowest Ca:Mg ratio on all soils due to its lower Ca content than the other populations on the same soil. 41 No consistent trend was evident in P or S content for the different populations on the different soils.North Spit and Coquille Falls were initially the highest in foliar P content.North Spit remained highest on all soils. Analysis of variance (Table 9) showed that sulfur was not significantly linked to soil, population, or soil X population inter-

action.All other nutrients varied significantly with the seedling populations, with N, P, K, and Ca being highly significant.Foliar N and P contents were not significantly related to soils, but Ca, Mg, and K varied with soil at the .0.1 level of significance.The soil X population interaction was not significant for Ca, but it was for N, P, K, and Mg.

Foliage Color

Color of the foliage over the term of the experiment did not effectively separate the different treatments.The majority of the new growth on all treatments in March wasMunsell colors (Hue, with the fraction Value/Chroma) 7.5 GY6/8, 7.5 GY 5/6, 7.5 GY 4/6, 5 GY 5/8, and 5 GY 5/6 (Appendix 2).These colors are difficult to separate and do not indicate a clear differencein new foliage color at the beginning of the experiment.By November, new foliage was all consistently greener,mostly (60 percent or greater) 7.5 GY 6/10.New foliage colors in May and August, for 42 seedlings grown on Coquille Falls and Iron Mountain, exhibited an increasing trend in green color toward 7.5 GY 6/10.Foliage color on North Spit and Seven Devils soils, with the exception of the Seven Devils population, changed toward yellow with colors from the 2.5 GY hue being common. On the North Spit sand the Rock Creek population showed the greatest change from March to May, with 68 percent of the seedlings exhibiting 2.5 GY colors, 35 percent being 2.5 GY 5/6.Forty-seven percent of the seedlings from Coquille Falls and 31 percent from North Spit had foliage in the 2.5 GY range. Twenty-eight percent of the Coquille Falls population on the Seven Devils soil had foliage yellower in May than in March. However, recovery to a greener color had occurred by the end ofthe experiment. Old foliage color in March was composed of the same colors as new growth, with the percentages varying slightly.In November, older foliage colors for most treatments were 7.5 GY5/6, 7.5 GY 4/6, 5 GY 5/8, and 5 GY 5/6. An exception was the Seven Devilspopula- tion on the Seven Devils soil which was 30 percent 2.5 G5/8, greener than most of the others. Color measurements were made difficult by charges in greenhouse shadows during the observations.Throughout the experiment C. lawsoniana foliage appeared slightly more greenthan Munse 11's 7.5 GY hue but not as green as the 2.5 G colors. 43 The Relationship of Growth to Foliar Nutrition

Multiple regression analysis of height growth on nutrient variables, by population, showed that K accounted for 64 to 95 percent of the variability within all populations (Table 13). All nutrients made significant contributions to height growth on the Coquille Falls population.Potassium accounted for 75 percent of the variability and the addition of Ca and total N increased theR2to 93 percent. Variability in height growth of the Rock Creek population could be explained by K alone,R2= .95. Sixty-four percent of the North Spit population's variation in growth could be linked to K.The addition of Ca and N to the equation brought theR2to .90.All nutrients except P contributed significantly. All nutrients contributed significantly to the height growthof the

Seven Devils population.Ninety percent of the variability was due to K, Ca, and N, with K alone accounting for 76 percent. Regression analysis of height growth on foliar nutrientcontent, by soil, indicated no single nutrient dominant (Table13).Phosphorous, N, and Mg contributed the most to the regression,in contrast to that within populations and across soils, as presented above. Correlations of foliar nutrient concentrations withchange in wet weight were similar to those with height growth(Table 14).Wet 44

Table 13Regression of Height Growth on Foliar Nutrient Content. Samples are grouped by population and by soil.**indicates a contribution significant at the 1 percent level, * indicates significance at the 5 percent level. Order of entry into the regression Population first second third fourth fifth sixth

Coquille Falls K** Ca** N** S* mg** P* Regression R2 .75 .81 .93 .94 .96 .97

Rock Creek S=* P* N** Ca** Mg Regression R2 .95 .96 .97 .99 .999

North Spit K* Ca** N* S* Mg * Regression R .64 .88 .90 .96 .99

Seven Devils K** Ca** N** Mg S RegressionR2 .76 .84 .90 .98 .98 Order of entry into the regression Soil1 first second third fourth fifth

Coquille Falls N.** K** Mg* Ca Regression R2 .40 .77 .77

Iron Mountain P* Mg* Ca* N K* Regression R2 .38 .59 .67 .71 .84

North Spit ID* Mg* K** Ca* Regression. R .59 .75 .91 .92 Seven Devils RegressionR2 .80 .89 .99

1Sulfur was not not included in the Regression 45

Table 14.Correlation Coefficients (r) of Foliar Nutrient Content With Height Growth and Change in Wet Weight for Samples Grouped by Population.

Population N P K Ca Mg S Coquille Falls Height .61 .69 .86 -.19 .04 .00

A Wet Weight .70 .58 .90 .15 -.31 .05 Rock Creek Height .92 .82 .97 -.14 -.20 -.50

A Wet Weight .79 .66 .90 .11 -.49 -.56

North Spit Height .72 -.42 .80 .11 -.31 -.54

A Wet Weight .76 -. 49 .85 .29 -.46 -.64 Seven Devils Height -.86 .03 .87 -.51 .19 -.26

A Wet Weight -.89 -.07 .89 -. 30 -. 06 -. 17 46 weight exhibited a larger negative correlation to change in Mg and Ca showed a slightly more positive correlation than that with height growth (Table 14).Correlations for all populations were similar for K, Mg, and S.Differences evident were the negative correlations of growth with N and Ca by the Seven Devils population and with P inthe North Spit population. 47

V. DISCUSSION

Height and rate of growth have been used as indicators of difference within species in many studies.Minckler (1950) showed ecotypic differentiation in shortleaf and loblolly pine from different geographical regions based on height growth.Localized altitudinal ecotypes of western white pine were separated on rate of growth and amount of foliage dry matter (Squillace and Bingham, 1958). Kruckeberg (1967) separated ultramafic and non-ultramafic ecotypes of Pinus contorta on height growth of seedlings in outdoor soil frames after 38 months. Chamaecyparis lawsoniana exhibited differences in total height growth and rate of growth.The Rock Creek population was consistently shortest and slowest in its growth.This difference may at least partially be accounted for by the age and condition of the seedlings when transplanted.The Rock Creek seedlings initially were larger and older than the other populations.Removal from the site and transport resulted in loss of more fine roots than other seedlings. Morrison (1969) showed that there was a high correlation between total growth and root area, and that complete nutrient uptake was accounted for by elemental reserves near root surfaces for jack pine and white spruce.The lack of finer roots could explain the slower growth of the Rock Creek seedlings.Seedlings from North Spit were 48 the tallest on each soil.No initial differences were noted to account for this growth. The controlled climate of the greenhouse narrowed the explanation of differences in growth to edaphic factors and inherent genetic characteristics of the population.This investigation is limited to the more observable differences of the soils and plant responses to the soils.Seven Devils soil and North Spit sand both exhibited irrigation problems that may help to explain the slower growth of all populations on these soils.The silty clay loam from Seven Devils compacted so that surface irrigation became ineffective and sub-irrigation was substituted.Pots containing North Spit sand did not drain as expected, the seedlings were waterlogged untilthe twelfth week of the experiment, when the watering schedule was changed.This change in watering is reflected in the late start of growth on these soils as seen in Figure 2. Slow growth on thesesoils was also affected by the low nutrient levels ofthe soils themselves, low in everything but P. Soil nutrients vary for each soil sample one takes but a range of nutrient contents for an area can be compiled.Bullard et al, (1957) reported data for soil nutrient contents within the Douglas-fir region of coastal and southwestern Oregon (Table10).Soil from Coquille Falls was the only one I studied to have comparablelevels of all nutrients sampled.Nitrogen and P contents on all soils fell 49 within the range given by Bullard et al.(1957), but all sites except Coquille Falls had lower levels of K.Potassium content of North Spit sand was only ten percent of the lower value listed by Bullard et al. (1957).Iron Mountain soil is at the lower end of the Ca range and North Spit and Seven Devils are below all reported data

(Bullard et al.,1957).Magnesium content is low on North Spit and Seven Devils but high on Iron Mountain, a result of the ultramafic origin.Soil nutrient contents suggest a greater nutrient availability on Coquille Falls and Iron Mountain soils that is reflected in the growth curves of C. lawsoniana on these soils. Organic matter content, compared to surface soil values of two plantations (Bullard et al. ,1957), was initially low on all soils except Iron Mountain.North Spit sand was the only soil remaining low in organic matter at the close of the experiment.The increases in organic matter on other soils may be an artifact of plant roots remaining in the soil sample. Initial foliar nutrient content compares favorably with C. lawsoniana sapling foliar nutrient contents of McNabb (unpublished) and Zobel (unpublished, Table 12).The Rock Creek seedling samples, when compared with Zober s data from my Iron Mountain site, support the similarity of the two sites.No data were available for direct comparison with the North Spit site,Zobel's (unpublished) Coos County Forest data, though geographically close 50 to Seven Devils and North Spit, were from saplings grown on a different soil. The pygmy forest soils of the Mendocino coast studied by Westman (1975) are similar to the Black lock at Seven Devils.Initial Seven Devils soil nutrient contents are higher than all of Westman's A horizon data except N, which is slightly lower on Seven Devils. Westman's (1975) Al horizon data are close to or slightly greater than Seven Devils Ca, K, and N contents.Magnesium content in the Westman Al is five times the initial Seven Devils content and twice that of the two final Seven Devils percentages.Final Seven Devils contents for Ca and N are greater than Westman's pygmy forest data. All Seven Devils soil P values are ten times greater than those of the pygmy forest. Trees at North Spit and Seven Devils seem to have many of their roots in the organic surface soil.This organic layer could serve as a nutrient source for the trees which was notavailable to my seedlings.Most of the nutrients in the pygmy forest soils (Westman, 1975) are in the Al horizon.If North Spit and Seven Devils are similar, then the most nutrient-rich horizon was not ,3.sed forthis study and North Spit and Seven Devils in the natural state maybe better substrates for growth than this study indicates. All populations grown on their native soils increased in nutrient content during the experiment with three exceptions.North Spit 51 decreased in N, which may be a result of growing without the organic layer to draw on.Seven Devils decreased in Mg by more than 50 percent but still had the highest foliar Mg content. Rock Creek, serving in place of Iron Mountain, decreased in Ca on the Iron Mountain soil. Youngberg's (1958) data represent nursery grown seedlings but his samples offer the only nutrient comparison available.The Greeley site is well irrigated and seedlings exhibited good growth. Seedlings at Corvallis and Wind River show higher uptake of K and N. These results (Table 12) fit within the range of this study's results and point out again the possible importance of K in growth of C. lawsoniana. Nutrient deficiency levels have not been established for C. lawsoniana. However, at the end of the experiment it showed none of the deficiency symptoms of C. obtusa indicated by Tsutsumi

(1962).Foliar color changes toward yellow on Seven Devils soil and North Spit sand had reversed by August.This recovery suggests that the yellowing was a result of the irrigation problems onthose soils discussed earlier.The severity of the color change in the Rock Creek population on North Spit sand may have been aggravatedby the lack of fine roots on the Rock Creek seedlings.The closest possibly applicable deficiency levels are those for Thuja plicata,another member of the Cupressaceae, determined by Walker,Gessel, and 52

Haddock (1965).If those levels were correct for C. lawsoniana, deficiencies should have been noticed for N and P.The lack of deficiency symptoms indicate that Walker, Gessel, and Haddock's (1965) deficiency levels for N and P cannot be applied directly to C. lawsoniana. Foliar nutrient data (Table 12) show, and analysis reaffirms, that when nutrients are observed across soils for all populations, K is ranked Coquille Falls (CF) > Iron Mountain (IM) > North Spit

(NS) = Seven Devils(SD).Calcium is ranked IM < CF = NS = SD, and Mg is ordered IM > NS > CF = SD.Nitrogen and P overlapped on all populations on Coquille Falls, Iron Mountain and North Spit soils.Foliar P on Seven Devils was lower than on all other soils but Seven Devils was not included in the analysis of variance and P was not significantly linked to soil. When foliar nutrient contents were grouped by population regardless of soil (Table 12), foliar N contents were ranked North Spit > Coquille Falls > Rock Creek.Seven Devils, though not included in the analysis of variance, had the highest foliar N on all soils except Coquille Falls.Potassium and P were both ranked NS > CF > RC = SD and Ca was ranked RC SD > CF > NS.Mag- nesium was equal in all populations on each soil. The above rankings are somewhat arbitrary since many of the treatment mean values are identical or within .01 or .02 percent of 53 each other.Even those treatments that can be separated by means may have ranges that overlap.The importance of these rankings lies in the fact that they do show differences on both soils and populations.These differences as well as the soil X population inter- actions determined significant by analysis of variance support the argument that nutrient content of the populations on the different soils was innately different.This same point can be supported for growth by noting that the rankings of average total height growth and change in wet weight show different population responses to the same soils (Tables 8 and 10). Regression analysis showed that ninety percent of the variability in height growth within all populations could be explained by variation in foliar K, Ca, and N.Potassium was the dominant nutrient responsible for 64 to 95 percent of the variability.Leyton (1958) proposed that when multiple regression analysis of nutrient contents showed a positive correlation with heightgrowth those nutrients would be the ones restricting growth. Mader andHowarth (1968) and Kawada, Nishida, and Yoshioka (1973) supportLeyton's hypothesis, working with red pine and hinoki (C. obtusa), respec tively.Chamaecyparis lawsoniana showed positive correlationof growth with K for all populations and with N for three.This adds another conifer in support of Leyton's (1955,1956) suggestion, prompted by work with Japanese larch and Scots pine,that height 54 growth is limited by only N and K.White and Leaf (1965) found K to be positively correlated with height growth in red pine.The low level of extractable K in the soil at three of my sites supports the possibility that K may be restricting growth. Nitrogen concentration is largest in the Seven Devils population on Seven Devils soil but is negatively correlated with height growth, r = -.86, and change in wet weight, r = -. 89, for the Seven

Devils population.The Seven Devils population had the highest N on the three soils where seedlings grewthe least.Nitrogen may be a non-limiting factor.If that is the case, then the negative correlation with both growth measurements may be due to a dilutioneffect. The North Spit population is the only one that has asubstantial negative correlation for P with height growth, r = -.42,and change in wet weight, r = -.49. Mean foliar P content for NorthSpit is highest of all populations on all soils.Phosphorous is probably a non-limiting nutrient for the North Spit population in this study. No one population is capable of significantly moregrowth on its native soil than other transplanted populations. The North Spit population grows taller than allother populations on each soil.This growth can now be examined in the light of the nutrient data.Seedling height growth differences between soil types can be linked to the K content of the soilsthemselves. 55

Coquille Falls produces the most growth and has the highest K content.Iron Mountain soil, though low in K by comparison (Bullard et al. ,1957), has more extractable K than either North Spit or Seven Devils and supports considerable growth.North Spit soil has low original K content, but North Spit seedlings have the highest K content of all populations on each soil (overlap with Coquille Falls onNorth

Spit).Since regression results suggest that K is limiting growth, North Spit's higher K contents indicate that it may be limited least, and consequently grow taller and faster than the otherpopulations. Low K in native North Spit sand may lead to development of adaptive mechanisms for K uptake which would be advantageous when transplanted to soils with higher K content. Though not as obvious as K in the regression, soil N contentof the various soil types varies in parallel fashion to heightgrowth and soil K content: CF > IM > NS = SD. Calcium content is lowest on all soils in the North Spit popula-

tion.Ability to grow well with low Ca content may beanother ad- vantage of the North Spit population.All soils except Coquille Falls were adequate to low in Cabut there was sufficient Ca for North Spit to grow taller. One further possible explanation of NorthSpit's better growth is that the greenhouse environment of highhumidity and low light intensity, with no local drought, and a climatewhere late growth is 56 possible, favored North Spit by closely resembling its native site. Coquille Falls on Coquille Falls is the only other treatment to have significantly different growth from the otherpopulations on the Coquille Falls soil.However, it is on its native soil.Nutrient differences do not appear to explain the difference in this one treatment.The difference in curvature appears to be a difference between the North Spit and inland populations.Curves for Coquille Falls and Rock Creek both level off while the North Spit populationhas not yet peaked.The difference may be a phenological one.North Spit may lack a mechanism stopping growth in the fall that is presentin the other three populations. Growth of the North Spit population on Iron Mountainsoil, greater than the other three populations, maybe a result of the Ca and Mg relations of the ultramafic soil.Iron Mountain has low Ca content and high Mg, but North Spit seems to havelow Ca require- ment, since it has the lowest Ca content ofall populations on each soil.Magnesium does not appear to be at a toxic leveland though significant in the regression it is the fifth nutrient toenter. Data for saplings on ultramafic soilsthroughout the range of C. lawsoniana (Zobel, unpublished) show thatthe foliar Ca:Mg ratio on the Iron Mountainsoil of this study is low compared tothe other field samples, none less than 2.2:1.None of the other causes for serpentine infertility listed by Walker(1954) were evident in this 57 study.Heavy metals were not investigated but the failure of the population from an ultramafic area to grow best on ultramafic soil shows that Ni and Cr are probably not strong selective forces on Iron Mountain.Proctor (1971a) showed that plants from different serpentine sources do not necessarily react the same on the same soil.Though the soil was similar, the Rock Creek population may have been sufficiently different from the native Iron Mountain population that no ecotypic adaptation would be noticed. The late start and slow growth on the Seven Devils and North Spit soils limited the detection of possible ecotypes but the different nutrient requirements, non-limiting N for Seven Devils seedlings and P for North Spit, showed that ecotypic differences may exist.The ability of the North Spit population to utilize K and outgrow other populations on their native soils while remaining low in foliar Ca content indicates definite adaptation and indicates the likelihood of ecotypes within C. lawsoniana. Greater growth of the Coquille Falls population on Coquille Falls than the Rock Creek and Seven Devils populations may be exhibiting an adaptation to its native soil that the other two populations cannot equal. In addition to the above specific possibilities for ecotypes,the different population responses to nutrients and the significantsoil X population interaction for height growth indicated by analysis of 58 variance suggest that innate differences inthe amount of growth are possible for the individual treatments. To further clarify the possibilities forecotypes within. C. lawsoniana more study is needed.I suggest the following as continuing investigations with C.lawsoniana:

1. Control of genetic variability.

a.longer term investigations using rootedcuttings.

b.seed collection from individual parenttrees followed by germination and growth studies.

2. Longer term transplant studies includingfield outplantings.

3. Nutrient culture experiments using:N, P, K, Ca, Mg, Ni, and Cr.

4. Sand culture orpot/fertilizer experiments using these populations and varying K levels. This study has barely scratchedthe surface of the ziutrient relations of C. lawsoniana.Much remains to be done. 59

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Append Lc I.Foliar Nutrient Content (as determined by the Soils Testing Lal-,oratory, 02egon State University). CF = Coquille Falls, IM = Iron Mountain, NS = North Spit, SD = Seven Devils, and RC = Rock Creek.* mistake in sample (mistakes listed at end of table). Sample X (mm. ) (grams) Pop. /soil Total Ht. LI Wet 9!)N P .X,K %Ca %Mg 0/IS Growth Weight

CF/CF #1 0.94 0.24 1.08 0.65 0.18 0.34 206 31.19 #2 1.05 0.28 1.08 0.66 0.18 0.20 192 29.51 #3 1.12 0.28 1.00 0.63 0.16 0.07 206 34.74 RC/CF #1 1.00 0.28 1.24 0.71 0.19 0.05 195 35.47 #2 0.94 0.26 1.04 0.63 0.20 0.01 165 38.10

#3* 1.13 0.30 1.04 0.60 0.23 0.07 117 22.00 NS/CF #1 1.16 0.32 1.28 0.55 0.19 0.10 227 22.45 #2 1.16 0.32 1.28 0.61 0.19 0.11 251 26.84 #3 1.10 0.32 1.28 0.57 0.20 0.07 234 24.23 SD/CF #1* 0.86 0.24 1.12 0.61 0.17 0.19 201 36.14

#2 0.96 0.22 1.16 0.61 0.19 0.20 180 2(5.10 #3 0.94 0.26 1.16 0.63 0.17 0.13 165 2.5.15 CF/IM #1 0.97 0.22 0.80 0.39 0.48 0.20 121 6.21 #2 0.94 0.28 0.92 0.42 0.51 0.18 113 5.61

#3 0.88 0.22 0.92 0.36 0.51 0.03 104 6.18 RC/IM #1* 0.85 0.26 0.88 0.39 0.45 0.:.1 102 ,,-"J. 98

#2 0.90 0.26 0.96 0.36 0.48 0.08 18 8.93 #3 0.90 0.24 0.84 0.40 0.51 0.06 106 8.89 NS/IM #1 0.89 0.28 1.00 0.32 0.51 0.20 143 11.33 #2 1.05 0.36 1.04 0.33 0.45 0.214 203 11.91 #3 1.00 0.30 0.92 0.31 0.48 0.23 177 17.07 SD/IM #1 1.18 0.34 O. 96 0.34 0.51 0.13 108 0.52 #2 1.00 0.26 0.92 0.39 0.45 0.17 131 2. _'8

#3 1.04 0.28 0.84 0.40 0.51 0.07 118 8.4.,:: CF/NS #1* 1.09 0.34 0.80 0.59 0.26 0.10 48 1.13 1.44 #2* 1.06 0.36 0.92 0.54 0.25 0.41 40 2.44 #3 0.92 0.28 0.84 0.64 0.23 0.05 49 0.21 RC/NS #1 0.57 0.16 0.60 0.73 0.28 0.14 13 -0. 13 #2 0.74 0.22 0.64 0.73 0. 29 0.03 19 #3 0.70 0.20 0.64 0.59 0.30 0.07 18 -0.73 66

Appendix I.Continued. Sample X (mm. ) X (grams) Pop. /soil Total Ht. A Wet %N %P %K %Ca %Mg V3S Growth Weight

NS/NS #1* 0.97 0.34 0.88 0.53 0.32 0.34 49 1.09

#2 1.01 0.38 0.96 0.51 0.32 0.31 35 1.15

#3 0.92 0.34 0.88 0.52 0.32 0.13 41 1.20

SD/NS #1 1.24 0.34 0.72 0.65 0.26 0.15 39 0.90

#2 1.20 0.30 0.64 0.75 0.23 0.13 37 0.71

#3 1.24 0.34 0.68 0.64 0.26 0.44 33 0.37

CF/SD #1 0.88 0.14 0.84 0.64 0.16 0.18 22 0.72

#2 0.94 0.18 0.84 0.73 0.17 0.15 19 0.69

#3 0.98 0.14 0.80 0.73 0.18 0.39 26 0.19

SD/SD #1 1.10 0.14 0.76 0.67 0.22 0.11 28 0.38

#2 1.16 0.14 0.92 0.75 0.22 0.21 37 0.92

*SampleRC/CF #3contained7seedlings, two of which wereRC/NS

SampleSD/CF #1contained9seedlings, two of which wereCF/CF SampleRC/IM #1contained 10 seedlings, two of which were SD/IM SampleCF/NS #1contained 10 seedlings, two of which wereSD/NS

SampleCF/NS #2contained9seedlings, two of which were NS/NS Sample NS/NS #1 contained 10 seedlings, two of which were CF /NS

Samples are mean of7to 10 seedlings except SD/SD #1 which has 15 seedlings included. Appendix II.Color of new growth foliage; numbers are percentage of seedlings per treatment observed having that color. Seven Devils/ Rock Creek/ Coquille Falls/ North Spit North Spit North Spit Color MarMay Aug NovMar May Aug Nov Mar May Aug Nov 2.5 G 4/6 2.5 G 6/8 2.5 G 5/8 3 7.5 GY 7/10 3 3 3 6 7.5 GY 6/10 6 70 84 10 65 39 77 7.5 GY 7/8 9 7 7.5 GY 6/8 6 16 9 6 3 10 3 41 7 3 6 7.5 GY 5/8 3 7.5 GY 7/6 3 7.5 GY 6/6 3 7.5 GY 5/6 26 19 3 6 9 3 16 59 3 7.5 GY 4/6 6 7.5 GY 6/4 3 7.5 GY 5/4 7.5 GY 4/ 4 5 GY 6/10 5 GY 7/8 3 3 3 5 GY 6/8 6 3 13 6 3 39 6 14 32 6 5 GY 5/8 29 6 53 16 23 6 3 6 5 GY 4/8 5 GY 7/6 5 GY 6/6 3 9 3 6 10 3 5 GY 5/6 6 3 19 6 6 3 5 GY 4/6 3 5 GY 5/4 2.5 CY 8/8 2.5 CY 7/8 3 10 2.5 CY 6/8 3 6 6 13 3 3 3 10 2.5 GY 5/8 Appendix II.Continued.

Seven Devils/ Rock Creek/ Coquille Falls/ North Spit North Spit North Spit Color Mar May Aug Nov Mar May Aug Nov Mar May Aug Nov

2.5 GY 7/6 3 7 2.5 CY 6/6 6 6 24 2.5 GY 5/6 6 3 9 35 3 2.5 GY 5/4 7.5 YR all 5 YR all 3 2.5 YR all 2.5 Yall 7 Appendix H.Continued.

North Spit/ Seven Devils/ Coquille Falls/ Seven Devils Rock Creek/ North Spit Seven Devils Seven Devils Iron Mountain Iron Mountain Nov Mar May Aug Nov Mar May Aug Nov Color Max May Aug Nov Mar May Aug Nov Mar May Aug 2.5 G 4/6 2. 5 G 6/8 3 2.5 C 5/8 9 7.5 GY 7/10 3 19 3 5 3 38 85 7.5 GY 6/10 28 79 15 37 65 4 24 63 35 59 88 61 7.5 GY 7/8 3 13 3 4 6 3 4 4 4 7.5 GY 6/8 22 3 25 6 5 21 5 13 4 16 4 3 3 7.5 GY 5/8 3 7.5 GY 7/6 9 4 7.5 GY 6/6 9 3 15 5 4 4 3 7.5 GY 5/6 42 6 33 11 63 12 13 18 16 10 7.5 GY 4/6 14 15 5 5 13 8 9 3 7.5 GY 6/4 7.5 GY 5/4 7.5 GY 4/4 3 3 3 5 GY 6/10 5 GY 7/8 3 3 3 5 8 4 10 24 6 14 58 4 5 GY 6/8 6 13 3 5 4 12 12 4 32 10 3 32 11 8 5 CY 5/8 8 17 30 11 5 3 3 5 CY 4/8 5 GY 7/6 9 15 3 10 5 GY 6/6 13 3 5 11 16 18 3 26 4 5 GY 5/6 3 37 5 5 CY 4/6 4 3 5 CY 5/4 3 4 2.5 GY 8/8 2-.5 CY 7/8 6 2.5 CY 6/8 3 5 12 8 2.5 CY 5/8 Appendix II.Continued.

North Spit/ Seven Devils/ Coquille Falls/ Seven Devils/ Rock Creek/ North Spit Seven Devils Seven Devils Iron Mountain Iron Mountain Color Mar May Aug Nov Mar May Aug Nov Mar May Aug Nov Mar May Aug Nov Mar May Aug Nov

2.5 GY 7/6 19 2.5 GY 6/6 9 3 5 12 4 3 3 2.5 GY 5/6 3 3 5 4 4 8 9 6 10 4 2.5 GY 5/4 3 7.5 YR all 3 3 3 5 YR all 3 2.5 YR all 2.5 Yall Appendix H.Continued.

Coquille Falls/ North Spit/ Seven Devils/ Rock Creek/ Coquille Falls/ North Spit/ Iron. Mountain Iron Mountain Coquille Falls Coquille Falls Coquille Falls Coquille Falls Color MarMay Aug NovMar May Aug NovMarMay Aug Nov MarMay Aug Nov Mar May Aug NovMar May Aug Nov 2.5 G 4/6 2.5 G 6/8 3 2.5 G 5/8 7.5 GY 7/10 12 4 12 6 3 7.5 GY 6/10 24 24 82 72 40 88 91 34 68 88 28 76 100 23 72 87 38 78 7.5 GY 7/8 6 3 7.5 GY 6/8 16 3 3 12 6 12 12 3 3 31 13 6 41 42 34 6 7.5 GY5/8 3 7.5 GY 7/6 7.5 GY 6/6 3 3 3 7.5 GY5/6 49 3 5912 21 3 3 3 23 14 4 68 10 44 3 13 7.5 GY 4/6 22 4 7 6 15 7.5 GY 6/4 7.5 GY5/4 3 7.5 GY 4/4 5 GY 6/10 5 GY 7/8 6 3 5 GY 6/8 52 55 4 36 6 25 29 3 6 10 16 19 25 3 3 3 16 5 GY 5/8 16 3 3 6 4 21 10 3 12 6 3 3 5 GY 4/8 11 6 6 24 6 33 4 9 6 6 5 GY 7/6 5 GY 6/6 5 GY 5/6 5 GY 4/6 3 5 GY5/4 2.5 GY8/8 2.5 GY 7/8 3 2.5 GY 6/8 3 10 3 -4 2.5 GY5/8 3 1 Appendix H.Continued.

Coquille Falls! North Spit/ Seven Devils/ Rock Creek/ Coquille Falls/ North Spit/ Iron Mountain_ Iron Mountain Coquille Falls Coquille Falls Coquille Falls Coquille Falls Colo: Mar May Aug Nov Mf.-U.' May Aug Nov Mar May Aug Nov Mar May AugNov Mar May Aug Nov Mar May Aug Nov

2.5 GY 7/6 2.5 GY 6/6 6 3 2.5 GY 5/6 6 3 2.5 GY 5/4 7.5 YR all 9 5 YR all 2.5 YR all 2.5 Yall