THE ECOLOGY OF THE MACROFUNGI
AT THE
LANPHERE-CHRISTENSEN DUNES PRESERVE,
ARCATA, CALIFORNIA
by
Sue Sweet Van Hook
A Thesis
Presented to
The Faculty of Humboldt State University
In Partial Fulfillment
of the Requirements for the Degree
Master of Arts
December, 1985 THE ECOLOGY OF THE MACROFUNGI
AT THE
LANPHERE-CHRISTENSEN DUNES PRESERVE
ARCATA, CALIFORNIA
by
Sue Sweet Van Hook
We certify that we have read this study and that it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a thesis for the degree of Master of Arts.
Major Professor
V 1 Y Approved by the Graduate Dean Acknowledgements
I wish to thank Dr. David L. Largent, my major professor and mentor, for years of excellent guidance and encouragement in the field of mycology; the members of my graduate committee, Drs. Dennis Anderson, Francis Meredith, and Richard Hurley, for their critical review of the thesis; mycologists, Dr. Orson K. Miller, Dr. Joseph F. Ammirati, and Dr. Harry D. Theirs, for their verification of species identifications; Hortense M. Lanphere for the background material on the fungi at the preserve and for her supportive correspondence; Marjorie Moore for data entry and graphics; The Nature Conservancy, for permitting the study at the Lanphere-Cristensen Dunes Preserve; and my family for their labor, love and patience. Table of Contents
Page
I. Introduction
A. Literature review 1 B. Objectives 5
II. The Study Area
A. Location 6 B. Geology 6 C. Climate 8 D. Plant Communities 9
III. Methods and Materials
A. Areas studied 10 B. Plots within areas 16 C. Measurements of precipitation and air temperatures 17 IV. Results A. Precipitation 25 B. Air temperatures near the ground 27 C. Climatalogical data from Eureka, California 32 D. Fungal Synecology 34 1. Characterization of species 34 2. Seasonal succession 35 3. Seasonal periodicity 41 4. Mid-dates of fruiting 42 5. Average Cluster Size and Cluster Range 61 6. Abundance 61 7. Phanerogam Associates 64 8. Distribution 65 V. Discussion
A. Variations in precipitation and temperature 68 B. Clusters, a method for recording sociability 70 C. Synecology of fungal flora 71 1. Characterization of the species 71 2. Seasonal succession 74 3. Seasonal periodicity 83 4. Mid-dates of fruiting 86 5. Abundance 90 6. Distribution 91 Table of Contents (continued)
Page
VI. Conclusion 96
VII. Appendix A 99
VIII. Appendix B 100
IX. Appendix C 103
X. Appendix D 104
XI. Literature Cited 149 I. Introduction
Α. Literature review
The literature on fungal synecology has been reviewed and critiqued by Hueck (1952), and by Cooke (1953).
Both investigators emphasized the difficulties arising from the great variability in earlier synecological methods and the absence of a unified approach to the study of fungal communities.
Hueck (1952) attributed the variability in methods to a lack of appropriate techniques for best characterizing a mycoflora and consequently he proposed that a plot should
be chosen so that there appears to be both an abundant and
an homogeneous flora of fungi as well as a homogeneity of
vascular plant vegetation. The plot, whether a series of quadrats or an area of irregular shape, should be
delimited in such a way that repeated checks are
possible. The plots should be sampled 6-8 times per year for several years until no new species are found. Scales,
modified from Braun-Blanquet (1932), Lange (1948), and
Haas (1932), should be chosen for recording abundance,
sociability, and substrate.
The early 1950's was a period of transition in the
study of fungal synecology. New methods of sampling were
combined with earlier techniques. Questions that arose
from sampling fungus populations using this new approach
involved: the use of stationary plots versus routes
(Parker-Rhoades, 1951; Kalamees, 1968); the sizes and 2 numbers of plots (Parker-Rhoades, 1951; Cooke, 1955;
Kalamees, 1968; Richardson, 1970; and Fogel, 1976); the periodicity of sampling (Richardson, 1970); and the description of fungi being sampled (Parker-Rhoades, 1951;
Kalamees, 1968; Hering, 1966; Richardson, 1970).
The objective of these various mycocoenological methods is to be able to characterize major and minor components of a fungal flora. Hering (1966) labeled those species representing 5 percent of the total biomass or total number as dominants. Richardson (1970) also used the 5 percent of total biomass criterion to determine the major species of a pine plantation in Scotland.
Parker-Rhoades (1951) designated species having 4 or more records as "prevalent" species and found that constancy of fruiting was an important criterion in placing a species in the "prevalent" category.
It is important to note that comparisons of fungal productivity or occurrence between years within a site, or between similar site types in the same year, are reliable only when both the abundance and sociability of the fungi have been recorded. Kalamees (1968) emphasized the importance of sociability over abundance saying, "It is the only indicator demonstrating, but partly (with the
present methods), the connection of fruiting bodies with the respective mycelia, thus giving a certain idea on the distribution of mycelia in the soil." He recognized that
in order to determine dominant species, it is necessary to 3 count clusters of fruiting bodies that represent a single mycelium.
Since the 1950's, only two studies in fungal synecology have been done in North America. Cooke (1955) studied the relationship of cryptogam to phanerogam communities in eastern Washington and adjacent Idaho and
Fogel (1976) studied sporocarp phenology of hypogeous fungi in a Douglas fir stand in western Oregon.
Cooke (1955), in describing the characteristics of the fungal flora, was able to make only general statements about constancy, fidelity, vigor, dominance, sociability and phenology. Constancy and species richness increased in the more moist associations. Fidelity to a phanerogam association was greatest among mycorrhizal species. Vigor was difficult to measure, but a loss of vigor was detected in poorer years by the smaller size of sporocarps.
Dominance, defined as the volume of space occupied by all sporocarps of the species observed, was not useful for describing competitive significance. Fall and spring fungal floras differed, with only 10 percent of the species occurring in both seasons.
Fogel (1976) determined that the minimum area that could be used for sampling was 100 square meters.
However, because of the time required to unearth this much area to find the fungi, sporocarp production was estimated from 50 randomly located one-square meter quadrats; sporocarp numbers and biomass were underestimated as a 4
result. Major species were defined as those comprising 5
percent or more of the total dry-weight production. Major species differed in each of the 3 years. The mid-dates of fruiting for each species were determined using
Richardson's (1970) formula, "m= Σ dn/Ν"; where d equals the number of days from the starting date times n, the number of sporocarps, all summed, then divided by N, the total number of sporocarps for a species. Sporocarp
production increased logarithmically in spring when the mean minimum temperature was between 0° and 6° C and the precipitation was less than 5 cm. In fall, production
began when precipitation totaled at least 0.5 cm per month
and increased until the mean minimum temperature dropped to 4° C.
Mycologists concur that the paucity of synecological studies is due to the inherent difficulties associated
with field studies of fungi (Hueck, 1952; Fogel, 1981).
Identification of fungal species often involves taxonomic
problems which must be overcome before a study of fungal ecology can begin. Also, in theory, sociological studies
should be based on the vegetative portion or dominant
stage in the fungal life cycle, as they are for vascular
plants. However, the inability to quantify microscopic
subterranean mycelia has led to a reliance on the
reproductive fruiting bodies, whose ephemeral nature and
irregular occurrence pose additional difficulties (Hueck,
1952). It is often impractical to carry out long-term 5 studies with weekly sampling, to obtain accurate records of fungal populations (Fogel, 1981). Lastly, fungal synecology requires the development of efficient methods of study different from those established for phanerogams
(Hueck, 1952).
Β. Objectives
The primary objective of this study was to develop a new method of recording sociability of macrofungi.
Α second objective was to describe the synecology of the fungal species of a beach pine/dunes system through frequent sampling based on the identification of sporocarps and clusters.
Α third objective was to produce a species list of macrofungi for this ecosystem that includes information on growth habit, habitat, ecological function, and distribution for each species. 6
II. The Study Area
A. Location
The 86 ha Lanphere-Christensen Dunes Preserve of The
Nature Conservancy, managed by Humboldt State University,
is a sand dune/beach pine system located 6.4 km west of Arcata, California on the Samoa Peninsula (Fig. 1). The
preserve is one of four minimally altered habitats
supporting native dune plant populations along the
California coast (Barbour, 1977). The system is limited to the west by the Pacific Ocean, to the east by
pastureland and a tidal slough. The well-defined limits of this ecosystem, in addition to its protected status,
make it attractive for studies.
The Dunes Preserve is habitat for many interesting and
unusual fungi (Smith, 1957; Stuntz, 1962; Largent, Miller, and Ammirati, personal communications).
B. Geology
The sand dune system extends for 35 km along Humboldt Bay from Table Bluff north to Little River (Fig. 1). The
sand dunes are best developed and reach their greatest
heights (27 m) and widths (1.2 km) along the Samoa Peninsula where sand accumulates against a forest south of
the preserve.
The sand consists of fine, light-colored grains of
quartz and feldspar averaging .10-.25 mm, and coarse, dark
particles of basalt .25-.50 mm in size (Johnson, 1963). In areas stablized by plants, the organic content of the 7
Figure 1. Location of the Lanphere-Christensen Dunes Preserve north of Humboldt Bay. 8
soil increases, but no pedogenic horizons exist (Gordon,
1978). The soil pH, measured at 30 cm depth, is highest
just behind the beach and becomes increasingly acidic as the organic content increases (Johnson, 1963).
C. Climate
The climate of the Humboldt Bay region, recorded from
Eureka, California, remains mild the year round. Annual temperatures normally range from 4-16° C. There is an
average of only 5 days per year with minimum temperatures
recorded at 0° C or below. Skies are normally overcast
in winter and foggy during summer months. Typically, there are only 60 clear days in a year, occurring mostly
in late spring and early fall. Successive storms deposit
an average of 99 cm of rain during the wet season from
September to May (National Oceanic and Atmospheric
Administration, 1981).
Prevailing winds blow northnorthwest during spring,
summer and fall with 10.8 kph mean velocity. The wind
blows most strongly from this direction during April-May
and September-October, reaching 64-88 kph maximum
velocities. As winter storms approach the coast, the low
pressure currents circulate counterclockwise, bringing
warm air from the south. Over the past 71 year period,
the maximum wind velocity recorded in Eureka, California,
has been 90 kph (National Oceanic and Atmospheric
Administration, 1981). 9
D. Plant Communities
Five plant communities, described by Barker (1976), are encountered successively from the mean high tide line inland to the forest.
The upper beach where drift-logs catch the sand is called the strand. The foredune is considered to be the first rise above the strand. The grasses and herbs of the strand and foredune constitute the littoral strip community.
Inland, a low-growing herbaceous mat of perennial herbs, grasses, mosses, and lichens comprises the established dunes community.
Low areas near the water table are called vegetated hollows. These deflation plains, often partially submerged in winter and spring, are colonized by herbs, sedges, rushes, dwarfed shrubs and trees constituting a young forest.
The beach pine forest is composed of conifers, sclerophyllous evergreen shrubs, low-growing perennial herbs, grasses, mosses, and lichens.
Bordering the pine forest are willow swamps with water-tolerant broad-leaved deciduous shrubs and trees and an understory of ferns, sedges, trailing herbs and vines
(Appendix B). 10
III. Methods and Materials
A. Areas studied
The reviews of the ecological literature by Hueck (1952) and Cooke (1953) demonstrated that studies should be conducted for 3-4 seasons (Kalamees, 1968; Richardson, 1970; Fogel, 1976). This is the length of time necessary to obtain an accurate picture of the fungal flora in an area (Hueck, 1952). However, this study was carried out for two years, from September through May 1979-1981, and the sampling interval was shortened from once per week to twice per week in an attempt to compensate for the difference in the number of seasons in which the fungal flora was analyzed.
Approximately 36 ha of the 86 ha preserve are covered with bare moving sand and are unproductive for fungi. Of the remaining 50 ha, 36 ha were divided into three areas of approximately equal habitat size and fungal abundance
(Fig. 2). Three areas were chosen so that each area could be sampled twice in a 6 day workweek. Areas I and II represented the south and middle portions of the beach pine forest respectively. Area III comprised the vegetated hollows, established dunes and littoral strip communities. Mrs. Hortense Lanphere, biologist and former owner of the study area, had named _locations where she collected fungi. Her location names, in addition to new names, were used to refer to 65 specific site locations 11
Figure 2, Study Areas I, II, and III. 12
within each area. The sites were coded by letter and number for entry into a computer (Fig. 3; Appendix C). Parker-Rhoades (1951) found that data collected by traversing a habitat, termed "perambulation", were not statistically different from those collected from permanent transects. Therefore, given the heterogeneous habitat within the beach pine forest of Areas I and II and the vegetated dune hollows of Area III, perambulation was chosen as the best method for surveying this ecosystem.
The open established dunes and littoral strip communities of Area III were surveyed by criss-crossing the terrain in a regular pattern. Sporocarps were easily spotted among the low-growing vegetation from the routes traversed.
The primary purpose of this survey was to record the abundance and sociability of the macrofungi. A new method was tried which involved recording absolute counts of sporocarps as they appeared in clusters that presumably represented the underground mycelia. In this way both the number of individuals and the number of clusters were used to characterize the dominant, subdominant, and common species of fungi in the flora. Dominant species were defined as those comprising 5 percent or more of the total number of sporocarps and/or 5 percent or more of the total number of clusters (Hering, 1966; Richardson, 1970; Fogel, 1976). Subdominant species were defined as those comprising 2-4.9 percent of the total number of sporocarps 13
Figure 3. Study Sites and preserve trail system. 14 and/or the total number of clusters that also produced more than 40 clusters in any one year. Common species were described as those fungi that produced 15 or more clusters, but which comprised less than 2 percent of the total number of sporocarps or total number of clusters in any one year.
Mid-dates of fruiting (Richardson, 1970) were calculated separately for each dominant, subdominant ani common species using the number of sporocarps and then the number of clusters. The function of the cluster data was to allow comparisons between species that typically produce a single large fruiting body and those that produce many small sporocarps. It also permitted comparisons of the sizes of clusters within a species in the two years. Average cluster size was determined by dividing the total number of sporocarps by the total number of clusters for each species.
In any ecological study, the identification of the organisms usually precedes sampling. However, in this study, ecological data were collected concurrently with taxonomic data until the identity of a species was learned.
Each of the areas was sampled for 3 hours biweekly, from September through May during the first year. All species of fungi encountered for the first time were collected as voucher specimens, and assigned a collection number. The following information was also recorded: date, area, site, number of sporocarps occurring in each 15
Figure 4. Voucher data collection sheet. 16 cluster, substrate, and all vascular plant species occurring within a 1, 2.5, and 5 m radius of a fungus might be mycorrhizal hosts for the fungus. Vascular plants that were associated 100 percent with individual fungal species are probably their mycorrhizal hosts (Trappe, 1962). The measurements of 1, 2.5, and 5 m were chosen so as to best characterize the vascular plants surrounding the fungus without making a detailed listing of every plant. Once a species had been identified, only date, area, site, and number of sporocarps in each cluster were recorded. Clusters of more than 50 fruiting bodies were recorded to the nearest increment of 10. Beyond 100, they were denoted as "abundant," but were later assigned a value of 100 to carry out all calculations. Consequently, the total number of sporocarps in the flora is underestimated. Perambulations during the second year took 1-2 hours each day, with each of the 3 areas sampled biweekly, except during the latter 3 weeks of December. The perambulation period was shorter the second year for two reasons: eighty percent of the species had been collected and identified the first season, so less time was spent the second year; there were fewer sporocarps per species to be recorded the second year.
B. Plots within areas Permanent plots were established in the second year to allow further study of constancy of fruiting, seasonal 17 succession, and the growth of perennial mycelia from year to year. Ninety-seven permanent plots were placed among 20 community associations within Areas I, II, and III, with 3-6 plots representing each association (Table I). The plant associations, based on the existing combinations of trees, shrubs and ground cover plants, were further divided according to open or closed canopies and various topographical settings, so as to represent the entire range of microhabitats for fungi. Circular plots 6 square meters in size were located in these association types according to where fungi fruited the first year of the study. A 25 cm dowel rod, with north indicated as a line on the top, marked the center of each plot. During weekly visits a 1.38 m stick with a hole drilled in one end was placed over the rod and swung in a circle to define each plot. The date and number of fruiting bodies for every species present, were recorded. The fruiting bodies were denoted as species A, B, C..., number 1, 2, 3... as they were mapped, so that precise records of sporocarp duration could be kept, and growth of different mycelia noted from year to year (Fig. 5). The numbers of fruiting bodies occurring within plots were added to perambulation data for describing the development of the flora.
C. Measurements of precipitation and air temperatures
Fungi fruit when temperatures are moderate and moisture is available (Watling, 1973). Therefore, part of this study included measurement of precipitation and 18
Figure 5. Plot data sheet 19
TABLE I
Plant associations within Areas I, II, III and associated fungal plots.
Plant Topo- No. οf Association Canopy graphy Plots Plots
BEACH PINE FOREST: AREAS I & II Pinus/grasses open level 5 1,2,63,64,65 Pinus/Arctostaph. open level 5 3,49,59,79,87 Pinus/Arctostaph. moderate level 5 60,70,80,88,92 Plnus/Vaccinium moderate ridge 5 5,7,77,89,93 Pinus/Vaccinium closed ridge 5 6,8,17,57,94 Pinus/Vaccinium moderate hollow 5 11,19,61,73,81 Pinus/Vaccinium closed hollow 5 12,18,71,86,95 Pinus/Cladina open level 6 9,58,82,83,90,96 Pinus/Cladina moderate level 5 10,84,85,91,97 Picea closed ridge 4 14,16,22,69 Picea closed hollow 4 20,68,72,78 Abies closed level 4 13,15,55,56 Myrica closed level 5 4,21,62,66,76
DUNES: AREA III Pinus/Arctostaph. open level 5 26,30,42,43,53 Pinus closed level 5 25,28,44,50,54 Pinus/Carex open hollow 3 67,74,75 Salix/Carex open level 5 27,29,41,48,52 Juncus/Lathyrus open level 5 24,39,46,47,51 Eriogonum/Solidago open level 6 23,31,32,33,34,40 Eriogonum/Solidago open ridge 5 35,36,37,38,45
Total: 97 20 temperature variation within the ecosystem and the correlation of these physical differences with changes in the fungal flora. In the first year, 8 sites were established to measure variations in precipitation between open meadows and dunes, and at different elevations beneath closed canopies within Areas I, II, and III (Table II, Fig. 6).
TABLE II
Weather sites. Distribution of sites among open and closed canopies and hollow and ridge topographies.
Open Closed
Hollow Site 3 Site 2 (3-6 m Site 5 Site 6 elevation) Ridge Site 4 Site 1 (10-15 m Site 8 Site 7 elevation)
Variations within each of the 8 sites were measured with 4 no. 10 tin cans placed evenly from the center (can 1) to the edge (can 4) of open sites, and from a tree's trunk (can 1) to its drip line (can 4) in closed sites.
The cans were painted with beige enamel to reflect light, to reduce heating of collected rain water, and to prevent rust. Plastic funnels were taped to the top of the can to minimize evaporation. Measurements were taken from the 32 cans after each storm (Fig. 7). 21
Figure 6. Location of weather sites 1-8. 22
Figure 7. Can with funnel used to measure rain.
Figure 8. Covered and uncovered maximum/minimum thermometers. 23
Four of the weather sites were eliminated the second season because similar results had been obtained from sites 1 and 7, 2 and 6, 3 and 5, and 4 and 8. Weekly maximum/minimum temperatures were recorded during the second year, between 3 August, 1980 and 14 June, 1981, from Taylor outdoor thermometers located at the four remaining sites 5-8. The thermometers were mounted on wooden stakes 10 cm above the ground surface to obtain temperatures near the soil. They were protected from rain and direct sunlight by yellow plastic gallon jars which were aerated by 15 1x2 cm flaps cut and bent outward (Fig. 8).
Temperatures inside the jars were compared to those from identical thermometers allowed to equilibrate outside the jars, to check for excess heating that might occur within the jars. The comparison was made at closed site 7 and open site 8 on several overcast and several sunny days in fall.
Precipitation for each recording was averaged among four cans for each site, then averaged among all 8 sites the first season, and 4 sites the second season. Total precipitation averaged from four sites the second year was compared to total precipitation obtained from a Humboldt
State University weather station located in a meadow 213 m northeast of site 8. Rain was collected from a standard gauge with a 45 mm diameter opening positioned at a standard height of 2 m above the ground. 24
Average maximum/minimum temperatures among the four sites were compared to those recorded by a Belfort hygrothermograph operating in the weather station northeast of site 8. The hygrothermograph was located at 1.25 m above the ground and recorded daily fluctuations in relative humidity and temperature. A daily record of weather conditions for the Dunes Preserve was kept beginning October, 1980 for reference. All macrofungi collected were either identified to species in the fresh condition, or described before drying for future determinations. Specimens are deposited in the
Humboldt State University Fungal Herbarium, Arcata,
California. 25
IV. Results
A. Precipitation The total precipitation averaged from sites 1-8 (year one) was 88.36 cm. Total precipitation averaged from just sites 5-8 (Year One) was 81.90 cm. During the second year rain totaled 63.40 cm from sites 5-8. The total precipitation recorded at the preserve weather station the second year was 81.4 cm (Table III). Annual precipitation was similar for 11 cans placed in the open, for cans adjacent to tree trunks, for cans located beneath a tree's drip-line, and for cans placed beneath a moderate canopy (Table III). The greatest amount of rainfall was measured from can 4, site 5; the least amount from can 4, site 7 (Table III). Mean precipitation for all sites was 18 cm less than the total precipitation at the preserve weather station. However, 5.5 cm more rain was collected from can 1 in site 8, located in the meadow nearest the preserve weather station (Table III). Weekly totals of rainfall, averaged from sites 5-8 for both years appear in Table IV and Fig. 9. During year one, 16.56 cm of rain had fallen by the end of October. By 7 November, 28 percent of the annual precipitation (24.36 cm) had been reached. Thirty-nine percent (34 .56 cm) of the yearly total had accumulated by the end of the first week in December. Seasonal rainfall by the end of
October of the second year was 4 .70 cm. By 7 November, 9 26
Figure 9. Precipitation averaged over sites 5-8 for Years One (Sept.,1979-Μay,1980) and Two (Sept.,1980-June,1981). 27 percent of the annual precipitation (5.60 cm) had been reached. Thirty-one percent (19.60 cm) of the yearly total had accumulated by the end of the first week in
December (Table IV).
B. Air temperatures near the ground
The mean, standard deviation, and range of maximum and minimum air temperatures for sites 5-8 as well as for the preserve weather station are shown in Table V.
The mean maximum air temperature averaged over sites
5-8 was 20.9° C. Maximum temperatures at 10 cm above the ground surface were greatest at sites 5 and 8, and ranged from 20.0 to 32° C in site 5 and from 19.5 to 33.0° C in site 8. Under the forest canopy, maximum temperatures varied only 8 degrees in site 6, and 7 degrees in site 7. The mean maximum air temperature 1.25 m above the ground recorded from the preserve weather station between January and June, 1981 was 17.7° C; the variation in maximum temperature was between 15.7 and 22.4° C. The mean minimum air temperature near the ground, averaged over sites 5-8, was 3.6° C. Minimum temperatures were lowest in sites 5 and 8, and ranged from
-2.0 to 12.0° C in site 5 and from -4.0 to 9.0° C in site 8. Minimum temperatures varied 11 degrees in site 6 and 11 degrees in site 7. The mean minimum air temperature at the preserve weather station between 28
III TABLE
Annual precipitation at weather sites. Precipitation recorded in cm. Total rainfall for each can, mean rainfall for each site, mean annual rainfall averaged among 8 sites Year One (Sept. 1979 - May 1980), and among 4 sites Year Two (Sept., 1980 - June, 1981).
Year One Year Two Total Total Total
Site 1 Site 5 Site 5 can 1 51.6 can 1 88.6 can 1 77.6 can 2 52.7 can 2 116.9 can 2 86.3 can 3 67.7 can 3 115.9 can 3 87.0 can 4 111.5 can 4 331.4 can 4 224.5 mean 70.9 mean 163.2 mean 123.8
Site 2 Site 6 Site 6 can 1 65.2 can 1 58.2 can 1 49.4 can 2 53.6 can 2 53.4 can 2 47.2 can 3 102.6 can 3 64.6 can 3 50.8 can 4 48.2 can 4 102.3 can 4 82.2 mean 67.4 mean 69.6 mean 57.4
Site 3 Site 7 Site 7 can 1 120.8 can 1 65.1 can 1 63.5 can 2 121.9 can 2 99.2 can 2 79.0 can 3 117.7 can 3 55.9 can 3 42.7 can 4 90.1 can 4 36.3 can 4 28.2 mean 112.6 mean 64.1 mean 53.3
Site 4 Site 8 Site 8 can 1 105.9 can 1 119.7 can 1 86.9 can 2 106.8 can 2 123.4 can 2 92.0 can 3 106.9 can 3 51.8 can 3 40.6 can 4 107.1 can 4 52.6 can 4 32.8 mean 106.7 mean 86.7 mean 63.1
Overall mean: 88.4 (sites 1-8) 81.9 (sites 5-8 63.4 (sites 5-8)
Total weather: 81.4 station 29
TABLE IV Weekly precipitation totals in cm averaged from sites 5-8 for Years One (Sept., 1979 - May, 1980), and Two (Sept., 1980 - June, 1981). Precipitation to date in parenthesis.
week: 1 2 3 4 Year One September 0.8 0.4 -- 0.1 (0.8) (1.2) (1.2) (1.3) October 0.3 3.4 4.7 6.8 (1.6) (5.0) (9.7) (16.5) November 7.8 -- 4.6 4.3 (24.3) (24.3) (28.9) (33.2) December 1.3 -- 4.0 3.5 (34.5) (34.5) (38.5) (42.0) January 0.9 6.8 1.8 -- (42.9) (49.7) (51.5) (51.5) February 2.3 0.7 00 9.2 (53.8) (54.5) (54.5) (63.7) March 2.5 6.8 -- 3.8 (66.2) (73.0) (73.0) (76.8) April 1.7 1.6 5.5 0.3 (78.5) (80.1) (85.6) (85.9) May -- -- 1.4 1.0 (85.9) (85.9) (87.3) (88.3)
Year Two September -- -- 0.3 0.1 (0.0) (0.0) (0.3) (0.4) October -- 3.1 -- 1.2 (0.4) (3.5) (3.5) (4.7) November 0.9 2.7 -- 2.6 (5.6) (8.3) (8.3) (10.9) December 8.7 -- -- (19.6) (19.6) (19.6) (19.6) January 3.0 0.2 9.2 4.7 (22.6) (22.8) (32.0) (36.7) February -- 3.2 1.8 2.2 (36.7) (39.9) (41.9) (43.9) March 2.9 1.7 1.4 3.6 (46.8) (48.5) (49.9) (53.5) April 1.7 0.4 0.5 0.1 (55.2) (55.6) (56.1) (56.2) May -- 0.7 4.2 1.0 (56.2) (56.9) (61.1) (62.1) June -- 1.3 -- -- (62.1) (63.4) (63.4) (63.4) 30
January and June, 1981 was recorded as 3.0° C; minimum temperatures ranged from -0.6 to 10.1° C.
The standard deviation from the mean of both maximum and minimum temperature was between 1.8 and 3.1° C.
TABLE V
Maximum/Minimum Temperatures. Air temperature recorded in o C at 10 cm above ground surface from 3 August, 1980 to 14 June, 1981 (January to June, 1981 for weather station).
Weather Mean all Site Site Site Site Station 4 sites 5 6 7 8
Mean max. 17.7 20.9 25.6 14.5 16.2 27.2
Mean min. 3.0 3.6 2.9 4.7 7.1 1.4
Sd.dev.max. 1.8 2.1 2.8 1.9 1.8 3.1
Sd.dev.min. 2.7 3.0 3.0 2.8 2.9 3.1
Range max. 15.7- 16.9- 20.0- 11.0- 13.0- 19.5- 22.4 25.0 32.0 19.0 20.0 33.0
Range min. -0.6- -0.9- -2.0- 0.0- 1.5- -4.0- 10.1 10.5 12.0 11.0 12.5 9.0
Maximum temperatures were higher and minimum temperatures lower in the sites 5 and 8. Extremes of temperature were less in sites 6 and 7. Weekly maximum and minimum temperatures averaged together from sites 5 and 8 are compared to temperatures averaged from sites 6 and 7 in Fig. 10. Weekly maximum and minimum temperatures 31
Figure 10. Maximum and minimum temperatures in °C averaged for sites 5 & 8 (top), and for sites 6 & 7 (bottom) during Year Two (Sept.,1980-June,1981). Maximum (❑),Minimum (■ ). 32 averaged over sites 5-8 illustrate the general temperature regime for year two (Fig. 11).
At site 7, there was no difference in maximum temperatures registered by the covered and uncovered thermometers. The minimum temperature was 1° C lower on the unprotected thermometer. Both maximum and minimum temperatures varied between the covered and uncovered thermometers at site 8. The maximum temperature was
2-3° C higher and the minimum temperature was 1 0 C higher under the protective jar.
C. Climatological data from Eureka, California
Climatological data recorded from a weather station located atop the United States Post Office building at 5th and H streets, Eureka, Ca. at 13 m elevation for the years
1979-1981 are summarized in Table VI.
TABLE VI
Climatological data from Eureka, Ca. Original measurements have been converted to metric units.
Means, 1979 1980 1981 Extremes
Avg. daily max. temp. 15.3 14.9 15.3 14 .2 Avg. daily min. temp. 8.7 8.4 8.8 8.2 Monthly avg. temp. 12.0 11.7 12.1 11.2 Total precipitation 90.65 77.42 109.96 100.99 Fastest k.p.h. wind 70.4 67.2 88.0 89.6 and wind direction S.E. S. S. S.W. Avg. % possible sun 61 45 53 50 Total clear days 64 65 68 78 Total partly cloudy days 117 106 97 99 Total cloudy days 184 195 200 188 33
Figure 11. Maximum and minimum temperatures in °C averaged over sites 5-8 during Year Two (Sept.,1980-June,1981). Maximum (0), Minimum (■ ) . 34
The mean maximum temperature from the Eureka station was lower than temperatures from sites 5, 7, and 8 and the preserve weather station. It was similar to the mean maximum temperature of site 6. The mean minimum temperature from the Eureka station was higher than all mean minimum temperatures recorded from the preserve.
Total precipitation shown in Table VI was calculated on a calendar year basis. Total precipitation in Eureka,
Ca. from September, 1979 to May, 1980 was 94.2 cm. This sum was approximately 10 cm greater than that recorded from the preserve sites. Eureka's total rainfall for the period September, 1980 through June, 1981 equaled 74.7 cm. This amount equaled that recorded from the mean of four sites and was approximately 7 cm less than that measured at the preserve weather station. However, no can from any site produced a total precipitation reading in either year similar to that recorded at the Eureka station.
The highest wind speed was measured as 88 kph from the south on 13 November, 1981.
The average percent of possible sunshine varied from
61% in 1979 to 45% in 1980 to 53% in 1981. The total number of clear days was between 64 and 68, of partly cloudy days between 97 and 117, and of cloudy days between
184 and 200.
D. Fungal Synecology
1. Characterization of Species
Six species were notes as dominants: Amanita 35 muscaria, Laccaria laccata, Suillus brevipes, Suillus
ρseudobrevipes, Suillus tomentosus, and Suillus umbonatus (Table VII). An additional 17 species were considered subdominant (Table VIII). Twenty more species were determined to be common (Table IX). 2. Seasonal succession
Thirty-four of the 43 dominant, subdominant and common species fruited during fall (Fig. 12). The first set to fruit in late September and early October was:
Chroogomphus vinicolor, Collybia dryophila, Hygrophoropsis aurantiacus, Lactarius rufus, Russula brevipes,
R.pectinata, and Suillus tomentosus. Sporocarps of
Cantharellus cibarius that fruited during June and July were still present in early September. Other species belonging to the Boletaceae appeared during the last two
weeks in October, along with Amanita muscaria, Armillaria
ponderosa, A. robusta, Clitopilus prunulus, Cortinarius vibratilis, Hebeloma crustuliniforme, Laccaria laccata,
Lactarius deliciosus, Rhizopogon occidentalis, Russula rosacea, Tricholoma flavobrunneum, T. flavovirens, and T.
vaccinium. Eleven species began fruiting in early to mid-November: Armillaria ponderosa, A. zelleri, C. Cortinarius collinitus, C. glandίcolor, C. pacificus, phoeniceus, Hebeloma sp.1, Mycena Pura, Paxillus
involutus, R. raoultii, Tricholoma albobrunneum, T. cingulatum, and T. saponaceum. The first sporocarps of
Amanita pantherina, Russula 'griseoviolaceus', 36
TABLE VII
Dominant Species. Species comprising 5% or more of the total numbers of sporocarps and/or 5% of the total number of clusters for Years One and Two. Numbers shown indicate by which category (numbers or clusters) the 5% criterion was met.
Year One Year Two No. No. No. No. Species sporocarps clusters sporocarps clusters
Amanita muscaria --- 190 --- 152 Laccaria laccata 1199 --- 1638 --- Suillus brevipes 989 236 1407 498 S. pseudobrevipes ------194 S. tomentosus --- 150 --- 292 S. umbonatus 986 --- 1038 ---
Yearly Totals: 13570 2515 12115 2969 5% of the Total: 678 126 606 148 37
TABU VIII Subdominant Species. Species comprising 2-4.9% of the total number of sporocarps and/or the total number of clusters which also have more than 40 clusters in any one year. Numbers shown indicate by which category (numbers or clusters) the criteria were met.
Year One Year Two No. No. No. No. Species sporocarps clusters sporocarps clusters Amanita pantherina --- 56 ------Armillaria ponderosa 302 107 --- Boletus edulis --- 94 ------Cantharellus cibarius 517 80 ------Chroogomphus vinicolor 284 65 327 146 Cortinarius glandicolor ------270 C. pacificus 310 ------Hebeloma crustuliniforme ------58 Hebeloma sp. 1 ------436 --- Hygrophorus aurantiacus 370 56 ------Lactarius deliciosus --- 94 --- L. rufus ------255 60 Leccinum manzanitae --- 52 --- 67 Russula pectinate ------414 62 R. rosacea 556 90 --- 62 Tricholoma flavobrunneum ------62 Tricholoma vaccinium 347 ------
Yearly Totals: 13570 2515 12115 2969
2-4.9% of Total: 271-665 50-123 242-594 59-145 38
TABLE IX Common Species. Species with 15 or more clusters and less than 2% of the total number of sporocarps or of clusters in any one year. Numbers shown indicate the number of clusters in the year the criterion was met.
Year One Year Two No. No. Species clusters clusters Amanita porphyria 25 15 Armillaria robusta -- 17 A. zelleri 17 -- Clitopilus prunulus 17 -- Collybia dryophila 22 17 Cortinarius collinitus 37 24 C. phoeniceus 31 32 C. vibratilis 16 -- Mycena Pura 17 -- Paxillus involutus -- 33 Rhizopogon occidentalis -- 20 Russula brevipes 46 -- R. 'griseoviolaceus' -- 28 R. raoultii -- 15 Sarcosphaera ammophila -- 34 Tricholoma albobrunneum -- 24 T. atroviolaceus 20 -- T. cingulatum -- 31 T. flavovirens 30 -- T. saponaceum 22 --
Yearly Totals: 2515 2969 2% of Total: 49 58 39
Figure 12. Seasonal succession, seasonal duration, and mid-dates of fruiting for 43 species. Number mid-date (o); Cluster mid-date (x); Year One (|-----|); Year Two ( ι--- ι) . 40
Figure 12. 41
Sarcosphaera ammophila, and Tricholoma atroviolaceus did not appear until December the first year and January the second year.
3. Seasonal Periodicity
Seasonal periodicity is the length of time that sporocarps of a species are produced continuously. Fungi with a short seasonal duration of 4-6 weeks included:
Armillaria robusta, Boletus edulis, Clitopilus prunulus,
Cortinarius collinitus, C. glandicolor, C. vibratilis,
Hebeloma crustuliniforme (yr 1), Leccinum manzanitae, Paxillus involutus, Russula pectinate, Suillus pseudobrevipes (yr 1), S. umbonatus (yr 1), Tricholoma albobrunneum, T. atroviolaceus, T. flavovirens, and T. saponaceum (Fig. 12). Those that spanned 7-10 weeks of fall included: Amanita muscaria, Armillaria ponderosa, A. zelleri, Cantharellus cibarius, Chroogomphus vinicolor,
Collybia dryophila, Cortinarius pacificus (yr 1), Hygrophoropsis aurantiacus, Lactarius deliciosus, Mycena
Pura, Russula brevipies (yr 1), R. rosacea, Tricholoma cingulatum (yr 1) and T. vaccinium. Seasonal durations of between 12 and 16 weeks were characteristic of: Amanita porphyria (yr 2), Cortinarius phoeniceus (yr 1), Russula brevipes (yr 2), Sarcosphaera ammophila, and Tricholoma flavobrunneum. Species that fruited over a period of
18-24 weeks were: Amanita pantherina, A. porphyria (yr
1), Cortinarius pacificus (yr 2), Hebeloma crustuliniforme
(yr 2), Hebeloma sp. 1, Rhizopogon occidentalis, Russula 42
'griseoviolaceus', R. raoultii, Suillus brevipes, S. pseudobrevipes (yr 2), S. umbonatus (yr 2), and Tricholoma cingulatum (yr 2). Sporocarps of the following fungi
could be found during any month of the rainy season between October and May: Cortinarius phoeniceus (yr 2),
Laccaria laccata, Lactarius rufus, and Suillus tomentosus (Figs. 13-27).
4. Mid-dates of fruiting
For thirty-four species, the number and cluster
mid-dates were the same or less than 1 week apart within each year, thus indicating that cluster size remained
relatively constant throughout the period of fruiting. Species such as Amanita porphyria (yr 2), Hebeloma sp.1
(yr 2), Laccaria laccata (yr 1), and Paxillus involutus
(yr 2) whose 'cluster mid-date' preceded its 'number mid-date' by 8 or more days, had a smaller cluster size at the beginning of the period (Table X). The six species,
Amanita porphyria (yr 1), Cortinarius phoeniceus (yr 1), Sarcosphaera ammophila, Suillus tomentosus (yr 2), S.
umbonatus (yr 2), and Tricholoma cingulatum (yr 2, whose 'number mid-dates' preceded their 'cluster mid-dates' by
more than 7 days, had greater average cluster sizes
earlier in the season (Table X). The mean fall mid-dates the first year for 43 species were 30 November and 29 November for number and cluster
mid-dates respectively. These two mid-dates fell on 6 December and 4 December the second year. This difference 4 3
Figures 13-27. Abundance for each of 43 species. Nos. of sporocarps along vertical axis; total no. of sporocarps for each species given in lower left corner for Year One (left side of graph), Year Two (right side). Abundance greater than 180 written at top of bar. Mean weekly maximum/minimum temperatures in o C for sites 5-8 (Year Two), taken from Figure 12, at top right of figure; Maximum (❑ ), Minimum (■ ). mean weekly precipitation in cm for sites 5-8 Years One and Two, taken from Figure 10, is shown below temperatures. 44
Figure 14. 45
Figure 15. 46
Figure 16. 47
Figure 17. 48
Figure 18. 49
Figure 19. 50
Figure 20. 51
Figure 21. 52
Figure 22 53
Figure 23. 54
Figure 24. 55
Figure 25. 56
Figure 26. 57
Figure 27. 58
TABLE X
Mid-dates of Fruiting. Mid-dates calculated using the formula m= Σ dn/Ν ( Richardson, 1970), for the number of sporocarps and the number of clusters of each species.
Year One Year Two
Species Numbers Clusters Numbers Clusters
Amanita muscaria Nov.16 Nov.23 Nov.16 Nov.15 A. pantherina Mar.17 Mar.15 Mar. 3 Mar. 7 A. porphyria Nov.12 Nov.18 Dec. 6 Dec. 6 Mar.17 Apr. 3 May 6 Apr.27 Armillaria ponderosa Dec.12 Dec.13 Dec.11 Dec.11 A. robusta Nov.17 Nov.12 Nov. 8 Nov.13 A. zelleri Dec. 5 Dec. 6 Dec. 2 Dec. 3 Boletus edulis Nov. 8 Nov. 8 Nov.13 Νοv.14 Cantharellus cibarius Nov.25 Nov.21 Oct.18 Oct.20 Feb.13 Feb.12 Jun.15 Jun.15 Jul. 6 Jul. 6 Chroogomphus vinicolor Nov. 7 Nov.12 Nov. 6 Nov. 9 Clitopilus prunulus Oct.31 Nov. 1 Nov.17 Nov.16 Collybia dryophila Nov.12 Nov. 8 Nov.15 Nov.22 Cortinarius collinitus Nov.30 Nov.29 Dec. 1 Nov.28 C. glandicolor Nov.20 Nov.15 Nov.25 Nov.20 C. pacificus Dec. 9 Dec. 9 Dec.27 Jan. 2 C. phoeniceus Dec. 4 Dec.16 Dec.13 Dec.13 Apr. 6 Apr.10 C. vibratilis Nov.13 Nov.11 Nov.29 Nov.29 Hebeloma crustuliniforme Nov.25 Nov.23 Dec. 3 Dec. 7 Hebeloma sp. 1 Dec.19 Dec.23 Jan.27 Jan.10 Hygrophoropsis aurantiacus Oct.30 Oct.31 Nov. 7 Nov. 6 Laccaria laccata Jan.13 Jan. 5 Jan. 2 Dec.27 Lactarius deliciosus Nov.16 Nov.16 Nov.19 Nov.20 Apr.27 Apr.27 L. rufus Jan.17 Jan.10 Jan.15 Jan.16 Leccinum manzanitae Nov.10 Nov.13 Nov.16 Nov.16 Mycena Pura Nov.14 Nov.14 Dec. 5 Dec. 9 Feb.23 Feb.23 Apr. 7 Apr. 7 May 4 May 4 Mar.28 Mar.28 Paxillus involutus Nov. 7 Nov.11 Nov.16 Dec. 2 Rhizopogon occidentalis Dec.19 Dec.19 Jan.27 Jan.25 Russula brevipes Nov.22 Nov.21 Dec. 2 Dec. 2 R. 'griseoviolaceus' Dec.12 Dec. 9 Jan. 7 Jan. 8 May 4 May 4 Apr. 6 Apr. 5 59
Year One Year Two
Species Numbers Clusters Numbers Clusters
R. pectinate Oct.18 Oct.13 Nov.11 Nov.15 R. raoultii Nov.30 Nov.28 Jan. 6 Feb. 7 R. rosacea Nov.21 Nov.15 Nov.22 Nov.25 Sarcosphaera ammophila Jan. 6 Feb. 5 Jan. 28 Feb. 7 Suillus brevipes Nov.12 Nov.12 Nov.21 Nov.23 S. pseudobrevipes Nov. 1 Nov. 1 Nov.14 Nov.16 Mar. 2 Mar. 2 Feb. 2 Feb. 8 S. tomentosus Nov.18 Nov.25 Dec.18 Dec.27 S. umbonatus Nov. 9 Nov. 5 Nov.14 Nov.22 Tricholoma albobrunneum Nov.20 Nov.23 Nov.20 Nov.19 T. atroviolaceus Dec.15 Dec.18 Jan.11 Jan.10 T. cingulatum Nov.28 Nov.28 Dec.27 Jan.22 T. flavobrunneum Nov.25 Nov.29 Nov.22 Nov.23 Apr.10 Apr.10 T. flavovirens Nov.15 Nov.13 Nov.21 Nov.21 Apr. 7 Apr. 7 T. saponaceum Nov.30 Nov.30 Dec. 3 Dec. 4 T. vaccinium Nov.25 Nov.22
Overall mean mid-date Nov.30 Nov.29 Dec. 6 Dec. 4 Apr.24 May 1 Apr.20 Apr.11 60 of one week in the peaks of the flora between the two
years was also apparent for individual species. The number or cluster mid-dates for Boletus edulis,
Cortinarius phoeniceus, Hebeloma crustuliniforme,
Hygrophoropsis aurantiacus, Leccinum manzanitae, Paxillus involutus, Russula raoultii, Suillus brevipes, S. umbonatus, and Tricholoma flavovirens were approximately one week +/- two days later the second year. Twelve species, Amanita muscaria, Armillaria ponderosa, A. zelleri, Chroogomphus vinicolor, Collybia dryophila,
Cortinarius collinitus, Lactarius deliciosus, L. rufus,
Russula rosacea, Tricholoma albobrunneum, T. flavobrunneum and T. saponaceum had a 'number mid-date' 0-3 days between
years. Nine species, Armillaria ponderosa, A. robusta, A. zelleri, Chroogomphus vinicolor, Cortinarius collinitus,
C. phoeniceus, Leccinum manzanitae, Sarcosphaera ammophila, and Suillus pseudobrevipes had a 'cluster mid-date' 0-3 days apart between years. Seventeen species, Amanita pantherina, A. porphyria, Cantharellus cibarius, Cortinarius pacificus, C. vibratilis, Hebeloma sp.1, Laccaria laccata, Mycena Pura, Rhizopogon occidentalis, Russula brevipes, R. 'griseoviolaceus', R. pectinata, Suillus brevipes, S. tomentosus, Tricholoma atroviolaceus and T. cingulatum, had number or cluster mid-dates separated by 8 or more days between years. 61
5. Average Cluster Size and Cluster Range The average cluster size was greater the first year for 34 of the 43 species. It was less the first year for
Amanita pantherina, Cantherellus cibarius, Hebeloma sp.1, Rhizοροgοn occidentalis, Sarcosphaera ammophila, Suillus tomentosus, Tricholoma albobrunneum, and remained the same for Amanita porphyria both years (Table XI).
Range in cluster size was greater the first year. Thirty-two species produced clusters of between 1, and 30 or more sporocarps, whereas only 22 species fit this category the second year. One species fruited with clusters of 1-5 sporocarps the first year, but 10 species did so the second year (Table XI).
6, Abundance
Abundance for each species is shown in relation to maximum/minimum temperatures (yr 2) and precipitation in Figures 13-27. Species that produced 200 or more sporocarps in a 4-6 week period the first year, such as
Armillaria ponderosa, Cortinarius glandicolor, Hygrophoropsis aurantiacus, and Tricholoma vaccinium, reached the peak of fruiting within a week or two of the first appearance of sporocarps. Similarly, sporocarps disappeared rapidly within one or two weeks after the peak of fruiting had been reached.
Many fungi produced the same pattern of fruiting with a rapid increase and sharp decline in the number of sporocarps in a 4-6 week period. However, fewer 62
TABLE XI Cluster Size and Range. Average cluster size and the cluster range for Years One and Two.
Year One Year Two Cluster Cluster Cluster Cluster Species Size Range Size Range Amanita muscaria 2.0 1-12 1.9 1-20 A. pantherina 1.5 1-9 1.6 1-5 Ā. porphyria 2.3 1-9 2.3 1-7 Armillaria ponderosa 2.8 1-16 1.8 1-5 A. robusta 5.8 1-23 4.7 1-18 Ā. zelleri 6.3 1-22 3.1 1-8 Boletus edulis 1.5 1-7 1.3 1-3 Cantharellus cibarius 6.4 1-35 18.0 1-180 Chroogomphus vinicolor 4.4 1-50 2.2 1-45 Clitopilus prunulus 3.2 1-9 1.1 1-2 Collybia dryophila 6.1 1-23 3.9 1-12 Cortinarius collinitus 3.7 1-29 2.5 1-14 C. glandicolor 16.5 1-50 8.7 1-60 C. pacificus 13.5 1-40 3.1 1-12 C. phoeniceus 7.1 1-15 4.3 1-20 C vibratilis 5.0 1-30 2.0 -2- Hebeloma crustiliniforme 2.8 1-10 1.8 1-8 Hebeloma sp. 1 5.6 1-30 8.1 1-30 Hygrophoropsis aurantiacus 6.6 1-80 1.8 1-55 Laccaria laccata 21.4 1-100 15.6 1-100 Lactarius deliciosus 1.7 1-10 1.6 1-10 L. rufus 8.1 1-30 4.2 1-25 Leccinum manzanitae 2.0 1-12 1.3 1-5 Mycena puna 4.2 1-15 1.8 1-5 Paxillus involutus 6.1 1-28 4.9 1-100 Rhizopogon occidentalis 1.0 -1- 2.1 1-6 Russula brevipes 2.3 1-10 1.0 -1- R. 'griseoviolaceus' 20.8 1-50 3.0 1-12 R. pectinata 10.5 1-25 6.6 1-100 R raoultii 13.0 1-40 2.3 1-13 R. rosacea 6.2 1-30 3.0 1-15 Sarcosphaera ammophila 3.5 1-10 5.2 1-50 Suillus brevipes 4.2 1-30 2.8 1-31 S. pseudobrevipes 3.1 1-45 2.4 1-12 S. tomentosus 1.3 1-15 3.2 1-10 S. umbonatus 8.9 1-55 8.0 1-40 Tricholoma albobrunneum 3.6 1-15 4.3 1-35 T. atroviolaceus 4.3 1-15 3.7 1-8 T. cingulatum 12.8 1-80 6.2 1-54 T. flavobrunneum 4.2 1-14 1.7 1-9 T. flavovirens 2.7 1-10 1.0 -1- saponaceum 5.8 1-40 2.1 1-8 T vaccinium 6.9 1-30
Overall mean 5.9 1-28 3.8 1-27 63 sporocarps were produced. Examples of fungi that exhibited this pattern were: Armillaria robusta, A. zelleri, Boletus edulis, Leccinum manzanitae, and
Tricholoma albobrunneum.
Laccaria laccata produced more than 150 sporocarps per month for many months during fall, winter and spring.
Other species produced between 40 and 125 sporocarps over several months time. These fungi were: Amanita pantherina, A. porphyria, Lactarius rufus, and Rhizopogon occidentalis.
Species belonging to the genus Suillus produced a large number of sporocarps quickly in the fall and continued to fruit in low numbers during the winter and spring. A second, lesser peak of fruiting was reached during late January and early February the second year.
Several additional fungi that fruited most abundantly during fall, produced a few sporocarps in the spring of the second year. These species were: Hebeloma crustuliniforme, Lactarius deliciosus, Russula raoultii,
Tricholoma flavobrunneum, and T. flavovirens.
For five species, Cortinarius collinitus, C. phoeniceus, Russula brevipes, R. raoultii, and Tricholoma saponaceum, the peak of fruiting came during the fourth week of November the first year and during the first week of December the second year.
Abundance varied greatly between the two years among certain fungi. The total number of sporocarps per species 64 was greater the second year for Laccaria laccata, Lactarius rufus, Paxillus involutus, Rhizopogon occidentalis, Sarcosphaera ammophila, Suillus brevipes, S. pseudobrevipes, S. tomentosus, Tricholoma albobrunneum, and T. cingulatum. On the other hand, abundance was less the second year for these species: Armillaria ponderosa,
A. zelleri, Boletus edulis, Cantharellus cibarius,
Clitopilus prunulus, Collybia dryophila, Cortinarius collinitus, C. pacificus, C. phoeniceus, Hygrophoropsis aurantiacus, Lactarius deliciosus, Mycena Pura, Russula brevipes, R. 'griseoviolaceus', R. raoultii, Tricholoma atroviolaceus, T. flavovirens, and T. saponaceum.
Many mycorrhizal species of fungi reached peaks in abundance during the same week of both years regardless of peaks in the amount of rainfall. These species included:
Amanita muscaria, A. pantherina, Armillaria ponderosa, A. robusta, A. zelleri, Boletus edulis, Cortinarius glandicolor, Hebeloma crustuliniforme, Laccaria laccata,
Leccinum manzanitae, Suillus brevipes, S. umbonatus,
Tricholoma cingulatum, and T. flavobrunneum.
In contrast, the weeks with greatest rainfall corresponded exactly to the weeks when Amanita porphyria fruited most abundantly during both years.
7. Phanerogam Associates
Forty of these 43 species were mycorrhizal fungi.
Thirty-seven of these species were found growing within 5 m of Pinus contorts var. contorts 100 percent of the 65 time. Two additional species, Amanita muscaria and
Tricholoma vaccinium, fruited in the presence of P. contorts 80 percent of the time or grew near Picea sitchensis when Pinus was absent (Table XII). Vaccinium ovatum grew within 5 m of 12 species 100 percent of the time. Most notable among these were Lactarius deliciosus with 11 recorded occurrences and
Armillaria ponderosa with 5 records. Cortinarius collinitus, Leccinum manzanitae, and
Suillus pseudobrevipes grew in the presence of Arctostaphylos uva-ursi var. coactilis 100 percent of the time. Sarcosphaera ammophila, a saprophytic fungus, always fruited in the presence of Eriogonum latifolium and
Solidago spathulata. 8. Distribution
Ninety-eight percent of the dominant, subdominant and common fungi occurred in the forest. Of these, 57 percent also fruited in the vegetated dune hollows. One species, Sarcosphaera ammophila occurred in the littoral strip and established dune communities and along the periphery of the vegetated hollows (Table XIII). 66
TABLE XIX
Phanerogam Associates. Fungal species that fruited 100% with Pinus contorts var. contorts, Vaccinium ovatum, and Arctostaphylos uva-ursi var. coactilis. The number of recorded occurrences for each fungal species is given.
Pinus Vaccinium Arctostaphylos
Α. pantherina 6 Α. ponderosa 5 Α. zelleri 2 Α. porphyria 3 Α. zelleri 2 C. collinitus 6 A. ponderosa 5 C. collinitus 6 C. glandicolor 1 A. robusta 2 L deliciosus 11 H. crustuliniforme 2 A. zelleri 2 M. pura 1 L. manzanitae 12 Β. edulis 12 R.'griseoviolaceus' 3 M. pura 1 C. cibarius 11 R. raoultii 3 S. pseudobrevipes 7 C. vinicolor 8 T atrovioloceus 2 Τ atroviolaceus 2 C. prunulus 6 Τ. flavobrunneum 4 Τ. flavobrunneum 4 C. collinitus 6 Τ. saponaceum 4 Τ. saponaceum 4 C. glandicolor 1 C. pacificus 9 C. phoeniceus 9 C. vibratilis 3 H. crustiliniforme 2 L. laccaria 11 L. deliciosus 11 L. rufus 5 L. manzanitae 12 M. pura 1 P. involutus 3 R. occidentalis 7 R. brevipes 6 R. 'griseoviolaceus' 3 R. pectinata 5 R. raoultii 3 R. rosacea 3 Suillus brevipes 17 S. pseudobrevipes 7 S. tomentosus 16 S. umbonatus 5 Τ. albobrunneum 4 Τ atroviolaceus 2 Τ. cingulatum 2 Τ. flavobrunneum 4 Τ. flavovirens 5 Τ. saponaceum 4 67
TABLE XIII
Distribution of fungi among phanerogam communities. Littoral strip (L.S.), Established dunes (E.D.), Vegetated hollows (V.H.), Willow swamps (W.S.), Pine Forest (P.F.).
Species L.S. E.D. V.H. W.S. P.F.
Amanita muscaria χ χ Α. pantherina χ χ Α. porphyria χ Armillaria ponderosa χ Α. robusta χ χ Α. zelleri χ Boletus edulis χ χ Cantharellus cibarius χ Chroogomphus vinicolor χ χ Clitopilus prunulus χ χ Collybia dryophila χ χ Cortinarius collinitus χ C. glandicolor χ χ C. pacificus χ χ C. phoeniceus χ C. vibratilis χ Hebeloma crustuliniforme χ χ Hebeloma sp. 1 χ Hygrophoropsis aurantiacus χ Laccaria laccata χ χ Lactarius deliciosus χ χ L. rufus χ Leccinum manzanitae χ Mycena pura χ Paxillus involutus χ χ Rhizopogon occidentalis χ χ Russula brevipes χ R. 'griseoviolaceus' χ R. pectinata χ R. raoultii χ R. rosacea χ Sarcosphaera ammophila χ χ χ Suillus brevipes χ χ S. pseudobrevipes χ χ S. tomentosus χ χ S. umbonatus χ χ Tricholoma albobrunneum χ χ Τ. atroviolaceus χ Τ. cingulatum χ χ Τ. flavobrunneum χ χ Τ. flavovirens χ χ Τ. saponaceum χ Τ vaccinium χ χ 68
V. Discussion
A. Variations in Precipitation and Temperature Precipitation recorded in Eureka, California was significantly lower than that measured at the preserve. Although total precipitation for years one and two at the preserve did not differ greatly, the pattern in which rainfall accumulated during the two fall seasons was quite variable. In the first year, 28 percent of the annual precipitation had fallen by early November, whereas only 9 percent had accumulated by that time the second year. However, by early December, the percentages of annual precipitation were similar, 39 and 31 percent respectively
(Fig. 28).
Temperature variation was greater in exposed sites, and was less beneath the forest canopy. Mean maximum temperatures were higher at sites in the preserve than at the preserve weather station or the Eureka station. Less air circulation 10 cm above the ground than at 1.25 m or 13 m may explain this difference. This difference may also be explained in part by heat radiated from the soil at night. The jar as well as heat radiated from the jar itself may have kept the minimum temperatures higher at 10 cm above the ground. Minimum temperatures were lower at
1.25 m than at 13 m, perhaps due to the sinking of cooler air at night. Consequently, weather data collected on site is more useful in developing correlations with development of 69
Figure 28. Cumulative precipitation; Year One ( ] ), Year Two ( 2 ). 70 fungal sporocarps than data collected from either the preserve weather station or from the Eureka station. B. Clusters, a Method for Recording Sociability
Censusing clusters, which presumably represent individual mycelia, provides an objective way to define the major species in a flora and also allow comparisons between species that form sporocarps gregariously and those that produce a single sporocarp. For example, three of the six dominant species which produced sporocarps in a gregarious manner, met the criterion for dominant species on the basis of the total number of sporocarps only. The three remaining dominant species produced either one sporocarp or a few sporocarps per cluster and qualified as dominant species on the basis of number of clusters. Average cluster size and average cluster range reflect the abundance of sporocarps each year and serve as an indicator of the relative productivity of fungal sporocarps for any one season. For example, when fewer total sporocarps were produced the second year in the preserve, there was a smaller average cluster size and average cluster range for the dominant, subdominant, and common species. Finally, recording the number of clusters enables determination of whether or not individual mycelia fruit every year and how the mycelia respond to favorable or poor environmental conditions. For example, abundance of sporocarps was less the second year than the first year, but more clusters were recorded. It is quite 71 possible that when fruiting was abundant, as was the case in the first year, individual clusters from separate mycelia were recorded as one cluster with many sporocarps. During the second year when abundance of sporocarps was less, clusters from separate mycelia were more distinct because there were fewer sporocarps produced by each mycelium.
C. Synecology of Fungal Flora
1. Characterization of the Species
Six fungi, Amanita muscaria, Laccaria laccata, Suillus brevipes, S. pseudobrevipes, S. tomentosus and S. umbonatus, qualified as dominant in each of the two years. When compared with four previous synecological studies (Table XIV), the number óf dominant species constant to each year was higher for the preserve, than in the other studies. This relatively high degree of constancy for dominant species may be due to the large area sampled and the greater total number of species observed in the preserve. For each of the other four studies, the percentage of the fungal flora represented by dominant species was higher (31-36%) than the 3 percent of the flora represented by six dominant species in the preserve. The difference of approximately 30 percent may have resulted again from the larger area sampled and the greater total number of species observed in the preserve (Table XIV). 72
Using data recorded from plots only in the preserve, the percentage of the flora composed of dominant species approached the higher percentages for the four other studies (15 percent for preserve plots compared to 31-36 percent for the other studies). If subdominant and common species for the preserve are included along with the dominant species, the percentage of the flora they represent increased to 21 percent, but this percentage was still lower than the percentages composed of dominant species for the other four studies (Table XIV).
Comparison of this study with four previous synecological studies indicates that the larger the area sampled, the greater the number of species encountered, and the more constant are the dominant species between years. The total number of species observed in the preserve was greater and the number of dominant species each year was more constant than in other similar studies (Fogel, 1976; Richardson, 1970; Cooke, 1955; and Parker-Rhoades, 1951), for the following reasons: the size of the area sampled was considerably larger; the period of sampling was longer; the frequency of sampling was greater; phanerogam diversity and mycorrhizal host species among the dunes and forest were greater; finally, the fungi sampled were epigeous species which are more numerous and more easily seen than the hypogeous species. 73
TABLE XIV
Comparison of dominant species and total number of species between preserve study and four other ecological studies. Area given in hectares. Criterion of 5% equals 5% of the total number of sporocarps or total number of clusters or total biomass. Value undetermined (undet.), dominant species plus subdominant (sub) and common species (com).
No.of Total No.Spec. Area Criter- domin. no.of % of constant Studies sampled ion species species flora ea. yr.
Preserve dominants 36.00 5% 6 206 3 6
Preserve plots 0.058 5% 8 52 15 undet.
Preserve &sub &com. 36.0 5% 43 206 21 13
Fogel (1976) 0.005 5% 8 26 31 3-8
Richardson (1970) 0.005 5% 10 28 36 4-7
Cooke (1955) 0.18 5% 6 72 35 undet. Parker- 4 or Rhoades more (1951) 100.00 occur. 60 172 35 undet. 74
2. Seasonal Succession
Seasonal succession of fungi occurred at the species
level and could be broken down according to species that
initiated the flora, species that formed the major portion
of the flora, and species that were stimulated to fruit by
cold temperatures (Table XV). After a dry summer, the fungal season began each autumn when 0-5 cm of rainfall
had accumulated, when maximum temperatures were between 22
and 25° C, and minimum temperatures ranged from
4.2-6.5° C. Chroogomphus vinicolor (yr 2), Collybia
dryophila (yr 1), Hygrορhοroρsis aurantiacus, Lactarius
rufus (yr 2), Russula brevipes (yr 2), R. pectinata (yr
1), and Suillus tomentosus (yr 1) initiated the floral
succession during September and the first week of October
(Table XV).
The major portion of the flora, comprised of 32
species, could be further divided into three subgroups:
early-abundant, longer-lasting, and later-abundant.
Twenty-six early-abundant species (Table XV), fruited
between the second week or October and the third week in
December. Sporocarps appeared between the second week of
October and the second week of November, when
precipitation was between 5.0 and 24.3 cm the first year
and between 3.5 and 8.3 cm the second year. Maximum
temperatures during this period were between 17.9 and
21.2° C and minimum temperatures ranged from -0.1 to
4.4° C (10.5° C the first week of November was
75
TABLE XV
Seasonal succession. Dominant, subdominant and common species grouped according to 8 periods of succession. Periodicity, precipitation and temperatures given for each successional period. Periodicity given in weeks (1,2,3,4) and months (S,O,N,D,J.F,M,A,Ma,Ju,J1,Au); Precipitation in cm; Temperature in o C, Ma = maximum, Mi = minimum.
Species Periodicity Precipitation Temperature
Initiated:
Chroogomphus 1S-10 0.0-5.0 (1) Ma 22.0-25.0 vinicolor (2) 0.0-3.5 (2) Mi 4.2- 6.5 Collybia dryophila (1) Hygrophoropsis aurantiacus Lactarius rufus (2) Russula brevipes (2) R. pectinata (1) Suillus tomentosus (1)
Early-Abundant:
Amanita 20-3D 5.0-34.5 (1) Ma 16.9-21.2 muscaria 3.5-19.6 (2) Mi -0.9-10.5 Armillaria robusta Began Began Began Boletus edulis 20-2Ν 5.0-24.3 (1) Ma 17.9-21.2 Chroogomphus 3.5- 8.3 (2) Mi -0.1- 4.4 vinicolor (1) Clitopilus Ended Ended Ended prunulus Cortinarius 4Ν-3D 33.2-38.5(1) Ma 16.9-18.6 collinitus 10.9-19.6(2) Mi -0.9- 2.5 C. glandicolor C. vibratilis Hebeloma crustuliniforme Lactarius deliciosus (1) L. rufus (1) Leccinum manzanitae Mycena pura 76
Species Periodicity Precipitation Temperature
Paxillus involutus Russula raoultii R. rosacea (1) Suillus brevipes S. pseudobrevipes S. tomentosus (2) S. umbonatus Tricholoma albobrunneum T. cingulatum T. flavovirens T. saponaceum T vaccinum
Longer-Lasting:
Collybia 20-4J 5.0-51.5 (1) Ma 16.9-21.2 dryophila 3.5-36.7 (2) Mi 3.8-10.5 Russula Began Began Began brevipes 20-40 5.0-16.5 (1) Ma 19.9-21.2 Tricholoma 3.5- 4.7 (2) Mi 3.8- 4.4 flavobrunneum Ended Ended Ended 1J-4J 42.9-51.5(1) Ma 16.9-20.4 22.6-36.7(2) Mi -2.0- 4.0
Later-Abundant:
Armillaria ΙΝ-4J 24.3-51.5(1) Ma 16.9-20.4 ponderosa 5.6-36.7(2) Mi -2.0-10.5 A. zelleri Began Began Began Cortinarius ΙΝ-2Ν 24.3-24.3(1) Ma 17.9-20.4 pacificus 5.6- 8.3(2) Mi -0.1-10.5 C. phoeniceus Ended Ended Ended Lactarius 1J-4J 42.9-51.5(1) Ma 16.9-20.4 deliciosus (2) 22.6-36.7(2) Mi -2.0- 4.0 Russula pectinata (2) R. rosacea (2)
Cold-Stimualted:
Amanita Began Began Began pantherina 4Ν-1J 33.2-42.9(1) Ma 16.9-18.6 Russula 10.9-22.6(2) Mi -0.9- 2.5 'griseoviolaceus' Sarcosphaera ammophila Tricholoma atroviolaceus
77 Species Periodicity Precipitation Temperature
Spring:
Began winter, continued through spring:
Amanita 1Μ-2Ju 66.2-88.3(1) Ma 19.6-24.1 pantherina 46.8-63.4(2) Mi 2.4-10.1 Russula 'griseoviolaceus' Sarcosphaera ammophila
Began fall, continued through winter and spring:
Amanita Winter Winter Winter porphyria 3D-4F 38.5-63.7(1) Ma 16.9-21.9 Cortinarius 19.6-43.9(2) Mi -2.0- 4.8 phoeniceus Spring Spring Spring Hebeloma 1Μ-2Ju 66.2-88.3(1) Ma 19.1-24.1 crustuliniforme 43.9-63.4(2) Mi 0.9-10.1 Hebeloma sp. 1 Laccaria laccata Lactarius rufus Mycena pura Rhizopogon occidentalis Russula raoultii Tricholoma cingulatum
Began fall, and refruited in spring:
Cortinarius 1Μ-2Ju 66.2-88.3(1) Ma 19.1-24.1 pacificus 46.8-63.4(2) Mi 0.9-10.1 Lactarius deliciosus (2) Tricholoma flavobrunneum (2) 78 exceptionally high for the season). The season for these
26 species ended between the fourth week of November and the third week in December when precipitation varied between 33.2 and 38.5 cm the first year and between 10.9 and 19.6 cm the second year. It was probably the low minimum temperatures (-0.9 and 2.5° C) that disrupted the season for these species. The longer-lasting species,
Collybia dryophila (yr 2), Russula breviρes (yr 1), and
Tricholoma flavobrunneum, began fruiting early in the season during the latter three weeks of October when precipitation was between 5.0 and 16.5 cm the first year, and between 3.5 and 4.7 cm the second year, and when maximum temperatures were between 19.9 and 21.2° C and minimum temperatures ranged from 3.8 to 4.4° C.
Although these three species began fruiting early, they lasted through December until the first to fourth week in
January even when minimum temperatures stayed below 4°
C. Later-abundant species, Armillaria ponderosa, A. zelleri, Cortinarius pacificus, C. phoeniceus, Lactarius deliciosus (yr 2), Russula pectinata (yr 2), and R. rosacea (yr 2), appeared during the first two weeks in
November when precipitation had reached 24.3 cm the first year and was between 5.6 and 8.3 cm the second year (Table
XV). Maximum temperatures were between 17.9 and 20.4° C and minimum temperatures ranged from -0.1 to 10.5° C during this period. Sporocarps disappeared between the 79 third week in December and the fourth week of January due
to prolonged minimum temperatures between -2.0 and 4° C.
Four cold-stimulated species, Amanita pantherina,
Russula 'griseoviolaceus', Sarcosphaera ammophila, and
Tricholoma atroviolaceus, seemed to be stimulated to fruit by low minimum temperatures between -0.9 and 2.5° C from the last week in November to the first week in January
(Table XV).
Twenty-one of the 43 dominant, subdominant and common species fruited for more than one season. Fourteen species (Table XVI), began fruiting in fall (September - third week in December) and produced sporocarps during
winter (late December - February) and spring (March -
June). These fungi showed a wide range of tolerance to environmental factors. The winter species, Amanita pantherina and Sarcosphaera ammophila, continued to fruit through April and May. Tolerance of warmer temperatures in spring seemed to indicate one of two things: either these species did not fruit in fall due to intense competition by other fungi or they required a cold- temperature stimulus. Both factors may influence their
sporocarp development during the winter months.
Sporocarps of Russula 'griseoviolaceus' and Tricholoma atroviolaceus disappeared between the end of December and the third week in January. Since minimum temperatures
were low (-2.0 - 4.0° C) during the entire period, it
may have been the warmer maximum temperatures of 20.0 and 80
TABLE XVI Patterns of Seasonal Duration. Seasons in which species were present: fall (F); winter (W); spring (S); summer (U). Year One only (1); Year Two only (2).
Species F W S U
Amanita muscaria χ Α. pantherina χ χ χ χ χ Ā. porphyria (1) χ χ Ā. porphyria (2) Armillaria ponderosa χ χ Α. robusta χ χ χ Ā. zelleri Boletus edulis χ Cantharellus cibarius χ χ χ Chroogomphus vinicolor χ Clitopilus prunulus χ Collybia dryophila χ Cortinarius collinitus χ C. glandicolor χ C. pacificus (1) χ χ C. pacificus (2) χ χ χ C. phoeniceus (1) χ χ C. phoeniceus (2) χ χ χ C. vibratilis χ Hebeloma crustuliniforme (1) χ Hebeloma crustuliniforme (2) χ χ χ Hebeloma sp. 1 χ χ χ Hygrophoropsis aurantiacus χ Laccaria laccata χ χ χ Lactarius deliciosus (1) χ Lactarius deliciosus (2) χ χ L. rufus χ χ χ Leccinum manzanitae χ Mycena pura (1) χ χ Mycena pura (2) χ χ Paxillus involutus χ Rhizopogon occidentalis (1) χ χ Rhizopogon occidentalis (2) χ χ χ Russula brevipes χ χ R. 'griseoviolaceus' (1) χ χ R. 'griseoviolaceus' (2) χ χ R. pectinata χ R. raoultii (1) χ raoultii (2) χ χ χ R. rosacea χ 81
Species F W S U
Sarcosphaera ammophila χ χ Suillus brevipes χ χ χ S. pseudobrevipes (1) χ χ S. pseudobrevipes (2) χ χ χ S. tomentosus χ χ χ S. umbonatus (1) χ S. umbonatus (2) χ χ χ Tricholoma albobrunneum χ Τ. atroviolaceus χ Τ. cingulatum (1) χ χ Τ. cingulatum (2) χ χ χ Τ. flavobrunneum (1) χ χ T flavobrunneum (2) χ χ χ Τ flavovirens (1) χ Τ. flavovirens (2) χ χ Τ. saponaceum χ Τ. vaccinium χ 82
20.4° C during the second and third weeks of January that halted sporocarp production. R. 'griseoviolaceus'
did refruit in March and lasted from March to June the second year indicating a tolerance of warmer temperatures similar to that of A. pantherina and S. ammophila (Table
XVI). Three species, Cortinarius pacificus, Lactarius deliciosus (yr 2) and Tricholoma flavobrunneum (yr 2) which fruited abundantly in fall, produced a few sporocarps again in spring when maximum temperatures were between 19.1 and 24.1° C and when minimum temperatures ranged from 0.9 to 10.1° C. Cantharellus cibarius was noted for its occurrence during June and July. Cantharellus cibarius is known for its tolerance of warm, dry weather (Lange, 1948). Fog drip apparently supplied enough moisture during these months to allow sporocarp development in this species. All species from Table XVI, which are followed by a
(1) or a (2), were not consistent between years with regard to seasonal succession. Variation between years
may be attributed to a later peak in precipitation and sporocarp production for Collybia dryophila, Lactarius deliciosus, Russula pectinata, R. rosacea, and Suillus tomentosus the second year. However, it is not understood
what factors may have caused Chroogomphus vinicolor,
Lactarius rufus, and Russula brevipes to initiate
sporocarp production earlier the second year. For Lactarius deliciosus and Tricholoma flavobrunneum, which 83
fruited only in fall the first year but in fall and spring the second year, it would seem that conditions must be right for sporocarps to be produced in spring. 3. Seasonal Periodicity Dominant, subdominant and common species of fungi
were most easily grouped into five patterns of seasonal periodicity and are shown in relation to seasonal succession in Table XVII. Twelve species (Table XVII) fruited consistently each year for 4-6 weeks during the early-abundant successional period. Tricholoma atroviolaceus fruited for 4-6 weeks during the cold-stimulated period each year. Ten species (Table XVII) were consistent for their seasonal succession and for a 7-10 week periodicity. Sarcosphaera ammophila lasted between 12 and 16 weeks and was a cold-stimulated species both years. There were six species (Table XVII) that fruited for 18-24 weeks during the same season each year. Laccaria laccata began fruiting in the early-abundant period and lasted for 8 months both years. The remaining 13 species that are shown with a (1) or a (2) in Table XVII were inconsistent in fruiting in one of three ways during the two years. Eight species,
Amanita porphyria, Cortinarius pacificus, C. phoeniceus,
Hebeloma crustuliniforme, Suillus pseudobrevipes, S. umbonatus, Tricholoma cingulatum and T. flavobrunneum,
began during the same successional period each year, but had different seasonal periodicity. Periodicity remained 84
ΤΑΒLΕ XVII
Seasonal Periodicity. Dominant, subdominant and common species of fungi grouped according to five seasonal periods, and shown in relation to periods of seasonal succession: initiated (I); early-abundant (ΕΑ); longer-lasting (LL); later-abundant (LA); and cold-stimulated (CS), and in relation to abundance patterns: many & short (1); few & short (2); few & several months (3); most fall & few winter & spring (4); most fall & few spring (5); many & long (6).
Seasonal Succession Abun- Periodicity Ι ΕΑ LL LA CS dance 4-6 weeks: Armillaria robusta χ 2 Boletus edulis χ 2 Clitopilus prunulus χ 2 Cortinarius collinitus χ 2 C. glandicolor χ 1 C. vibratilis χ 2 Hebeloma crustuliniforme (1) χ 2 Leccinum manzanitae χ 2 Paxillus involutus χ 2 Russula pectinata χ 2,1 Suillus pseudobrevipes (1) χ 5 S. umbonatus (1) χ 1 Tricholoma albobrunneum χ 2 Τ. atroviolaceus χ 2 Τ. flavovirens χ 2,5 Τ-: saponaceum χ 2
7-10 weeks:
Amanita muscaria χ 1 Armillaria ponderosa χ 1 Α. zelleri χ 2 Cantharellus cibarius χ 1 Chroogomphus vinicolor χ(2) χ(1) 1 Collybia dryophila χ(1) χ(2) 2 Cortinarius pacificus (1) χ 1 Hygrophoropsis aurantiacus χ 1,2 Lactarius deliciosus χ 2,5 85
Seasonal Succession Abun- Periodicity Ι ΕΑ LL LA CS dance Mycena pura χ 4 Russula brevipes (1) χ 2 R. rosacea χ 1 Tricholoma cingulatum (1) χ 2 Τ. vaccinium χ 1
12-16 weeks: Amanita porphyria (2) χ 3 Cortinarius phoeniceus (1) χ 4 Russula brevipes (2) χ 3 Sarcosphaera_ ammophila χ 3 Tricholoma flavobrunneum (1) χ 2 18-24 weeks: Amanita pantherina χ 3 Amanita porphyria (1) χ 3 Cortinarius pacificus (2) χ 4 Hebeloma crustuliniforme (2) χ 4 Hebeloma Sp. 1 χ 4,6 Rhizopogon occidentalis χ 3 Russula 'griseoviolaceus' χ 5 R. raoultii χ 2,4 Suillus brevipes χ 4 S. pseudobrevipes (2) χ 4 S. umbonatus (2) χ 4 Tricholoma cingulatum (2) χ 4 Τ. flavobrunneum (2) χ 4
8 months: Cortinarius phoeniceus (2) χ 4 Laccaria laccata χ 6 Lactarius rufus χ(2) χ(1) 6 Suillus tomentosus χ(1) χ(2) 4 86
constant, but succession varied for Chroogomphus vinicolor, Collybia dryophila, Lactarius rufus, Suillus tomentosus. Both periodicity and succession were different the two years for Russula brevipes.
Sampling over a longer period would confirm the patterns of seasonal succession and periodicity for the species that were consistent between years and would further demonstrate that other non-consistent species remain flexible, fruiting at various times of the year when suitable environmental conditions arise.
4. Mid-dates of Fruiting
The overall mid-date for dominant, subdominant, and common species to fruit was one week later the second year, due to the later peak in precipitation that fall.
However, trends in mid-dates of fruiting differed from the overall pattern for individual species. 'Number mid-dates' for Amanita muscaria, Armillaria ponderosa, A. zelleri, Chroogomphuis vinicolor, Collybia dryophila, Cortinarius collinitus, Lactarius deliciosus, L. rufus,
Russula rosacea, Tricholoma albobrunneum, T. flavobrunneum and T. saponaceum were only 0-3 days apart between years
(Table XVIII). Armillaria ponderosa, A. robusta, A. zelleri, Chroogomphus vinicolor, Cortinarius collinitus,
C. phoeniceus, Leccinum manzanitae, Sarcosphaera ammophila
and Suillus pseudobrevipes had 'cluster mid-dates' 0-3 days apart (Table XVIII). These species clearly did not
respond to rainfall, which varied considerably between the 87
Ι ABLE XVII T
Mid-dates of Fruiting. Dominant, subdominant and common species of fungi grouped according to 4 patterns in mid-dates of fruiting, separated by 'number' and 'cluster' mid-dates, and by whether they were earlier (E) or later (L) the second year.
Number Cluster Patterns Ε L Ε L
0-3 days apart:
Amanita muscaria -- -- Armillaria ponderosa χ χ Α. robusta χ χ χ Ā. zelleri Chroogomphus vinicolor χ χ Collybia dryophila χ Cortinarius collinitus χ χ C. phoeniceus χ Lactarius deliciosus χ L. rufus χ Leccinum manzanitae χ Russula rosacea χ Sarcosphaera ammophila χ Suillus pseudobrevipes χ Tricholoma albobrunneum -- -- Τ. flavobrunneum χ Τ saponaceum χ
4-7 days apart:
Boletus edulis χ χ Cortinarius glandicolor χ χ Hygrophoropsis aurantiacus χ Lactarius deliciosus χ L. rufus χ Leccinum manzanitae χ Russula raoultii χ χ Suillus umbonatus χ Tricholoma albobrunneum χ Τ. flavobrunneum χ Τ. flavovirens χ Τ. saponaceum χ 88
Number Cluster Patterns Ε L Ε L
8-15 days apart:
Amanita muscaria χ Α. pantherina χ χ Armillaria robusta χ Clitopilus prunulus χ Collybia dryophila χ Hebeloma crustuliniforme χ χ Hygrophoropsis aurantiacus χ Laccaria laccata χ χ Paxillus involutus χ Russula brevipes χ χ R. rosacea χ Suillus brevipes χ χ S. pseudobrevipes χ Tricholoma flavovirens χ
Greater than 15 days apart:
Amanita porphyria χ χ Cantharellus cibarius χ χ Clitopilus prunulus χ Cortinarius pacificus χ χ C. vibratilis χ χ Hebeloma sp. 1 χ χ Mycena pura χ χ Paxillus involutus χ Rhizopogon occidentalis χ χ Russula 'griseoviolaceus' χ χ R. pectinata χ χ Sarcosphaera ammophila χ Suillus tomentosus χ χ S. umbonatus χ Tricholoma atroviolaceus χ χ Τ. cingulatum χ χ 89 two years. For these species, a stimulus such as a response to a particular photoperiod by the mycorrhizal host, triggered sporocarp development at the same time each year. However, most species had different mid-dates for the two years and 'number mid-dates' varied less than the 'cluster mid-dates' between years. An explanation for why 'number mid-dates' were more similar may be that fungi are adapted to producing the majority of sporocarps at approximately the same time each year. However, mycelia may continue to produce a few sporocarps for as long as conditions are favorable. Hence, the mid-date for clusters of a species would vary more over the years.
'Number mid-dates' for six species, Boletus edulis,
Cortinarius glandicolor, Leccinum manzanitae, Russula raoultii, Suillus umbonatus, and Tricholoma flavovirens, were between 4 and 7 days later the second year, due to the later peak in rainfall that year. Similarly, 'cluster mid-dates' for B. edulis, C. glandicolor, Hygrophoropsis aurantiacus, Lactarius deliciosus, L. rufus, R. raoultii and Tricholoma saponaceum were 4-7 days later the second year. But 'cluster mid-dates' for two species, Tricholoma albobrunneum and T. flavobrunneum, were 4-7 days earlier the second year. Since 'number mid-dates' for these two species were Ο and 3 days apart respectively, it is likely that these species fruited as a response to their mycorrhizal host (Table XVIII). 90
Cortinarius phoeniceus, Hebeloma crustuliniforme,
Hygrophoropsis aurantiacus, Paxillus involutus, Russula brevipes, Suillus brevipes and S. pseudobrevipes had
'number mid-dates' 8-15 days later the second year, and
'cluster mid-dates' for Clitopilus prunulus, Collybia dryoρhila, H. crustuliniforme, R. brevipes, R. rosacea, S. brevipes and Tricholoma flavovirens were also 8-15 days later. Amanita pantherina, Armillaria robusta and
Laccaria laccata and 'number mid-dates' 8-15 days earlier the second year and Amanita muscaria, A. pantherina and L. laccata had 'cluster mid-dates' 8-15 days earlier.
Species that fruited later the second year probably responded to the later peak in precipitation. Those that fruited earlier may have done so because of competitive relations with other fungi.
Those species whose number and cluster mid-dates were more than 15 days later the second year (Table XVIII) probably depended on a certain amount of accumulated rainfall to stimulate sporocarp development.
Number and cluster mid-dates for Cantharellus cibarius were perhaps earlier the second year because the peak in
October may have been an extension of the summer production and without a peak in February the second year, the summer production came sooner.
5. Abundance
Dominant, subdominant, and common species of fungi exhibited six basic patterns of fruiting. The first was 91
production of many sporocarps in a short time (4-10
weeks). Other species produced few sporocarps over a similar time period. Production of a few sporocarps over
12-24 weeks constituted a third pattern. Some species fruited abundantly during fall, but continued to produce a few sporocarps during winter and spring. Several fungi
produced most of their sporocarps in fall, but reappeared briefly in spring. Lastly, the sixth pattern comprised abundant sporocarp production for 8 months (Table XVI).
Abundance varied considerably within species between years (Table XIX). It is not understood why abundance for the 20 species listed in Table XIX was equal or was less the first year. Six of the 20 species were dominant species. Since total number of sporocarps was significantly higher for these six species, perhaps a difference of several hundred sporocarps between years is not significant. Abundance for the majority of species was probably greater the first year due to greater moisture that year. Factors such as competition between species certainly compound the direct effects of
precipitation and temperature on sporocarp production.
However, the later peak in precipitation the second year
produced a later peak in overall abundance which was reflected in the same overall trend for mid-dates of
fruiting.
6. Distribution
Thirty-nine of 43 dominant, subdominant and common 92
TABLE XIX
Variation in Abundance of sporocarps within a species. Abundance shown as Equal, Less or Greater the First Year. Equal abundance included a difference of 0-25 sporocarps between years.
Equal:
Amanita pantherina Hebeloma crustuliniforme Armillaria robusta Leccinum manzanitae Cortinarius glandicolor
Less:
Amanita muscaria Sarcosphaera ammophila Chroogomphus vinicolor Suillus brevipes Hebeloma sp. 1 S. pseudobrevipes Laccara laccata S. tomentosus Lactarius rufus S. umbonatus Paxillus involutus Tricholoma albobrunneum Rhizopogon occidentalis Τ. cinqulatum Russula pectinata
Greater:
Amanita porphyria Hygrophoropsis aurantiacus Armillaria ponderosa Lactarius deliciosus Α. zelleri Mycena pura Boletus edulis Russula brevipes Cantharellus cibarius R. 'griseoviolaceus'_ Clitopilus prunulus R. raoultii Collybia dryophila R. rosacea Cortinarius collinitus Tricholoma atroviolaceus C. pacificus Τ. flavobrunneum C. phoeniceus T. flavovirens C. vibratilis Τ. saponaceum 93 species were mycorrhizal. A 100 percent association between all fungus sporocarps collected and the potential host strongly suggests that the phanerogam species is the mycorrhizal host for that fungus. However, this correlation does not verify the association; pure culture synthesis in the laboratory is required for this. Thirty-seven mycorrhizal species were located within 5 m of Pinus contorta var. contorta in all instances recorded
(Table XII). Nine mycorrhizal species, Armillaria ponderosa, A. zelleri, Cortinarius collinitus, Lactarius deliciosus, Mycena pura, Russula 'griseoviolaceus', R. raoultii, Tricholoma atroviolaceus, T. flavobrunneum, and T. saponaceum, were situated near Vaccinium ovatum; 9 species, Armillaria zelleri, Cortinarius collinitus, C. glandicolor, Hebeloma crustuliniforme, Leccinum manzanitae, Mycena pura, Suillus pseudobrevipes, Tricholoma atroviolaceus, T. flavobrunneum, and T. saponaceum, were near Arctostaphylos uva-ursi var. coactilis in 100 percent of the records. Mycorrhizal relationships between Basidiomycetes and ericaceous shrubs are well-documented (Trappe, 1962; Largent et al., 1980).
Armillaria ponderosa and Lactarius deliciosus may be mycorrhizal with Vaccinium instead of pine since this phanerogam host was recorded 100 percent of the time with these species. Such a hypothesis is supported by the observation that in Douglas fir or true fir ecosystems in northwestern California, A. ponderosa and L. deliciosus 94 always fruit near Vaccinium ovatum (personal observation). Similarly, it is possible that Leccinum manzanitae and Suillus pseudobrevipes, which occurred in the presence of Arctostaphylos uva-ursi var. coactilis in 100 percent of the records, were mycorrhizal with this ericaceous species.
Distribution of some mycorrhizal species may be explained by age of the host plant. Amanita muscaria,
Armillaria robusta, Hebeloma crustuliniforme, Laccaria laccata, Paxillus involutus, Suillus umbonatus, and Tricholoma cingulatum formed sporocarps in the vegetated hollows where trees were younger (1-40 years old), and only rarely were found in the forest. Consequently these species may only be associated with pines, spruces and willows early in the phenology of the forest community as it begins in the dune hollows. Other fungi were completely absent from the vegetated hollow, and fruited only in the forest where trees have been aged to 90 years. These fungi, Amanita porphyria, Armillaria ponderosa, A. zelleri, Cantharellus cibarius, Cortinarius phoeniceus var. occidentalis, Lactarius rufus, Leccinum manzanitae, Russula 'griseoviolaceus', and R. raoultii, may represent fungi that colonize roots of these hosts as the forest matures.
Four of the subdominant and common species, Collybia dryophila, Hygrophoropsis aurantiacus, Μycena pura, and Sarcosphaera ammophila, were saprophytes. Of these, M. 95 pura occurred 100 percent in association with litter of Vaccinium and Arctostaphylos. H. aurantiacus and M. pura decomposed forest litter. C. dryophila grew in the vegetated hollows as well as in the forest. Sarcosphaera ammophila colonized plant debris in the foredunes, established dunes, and periphery of the vegetated hollows. 96
VI. Conclusion
A new method for recording sociability of fungi was developed based on a cluster, which represents an
individual mycelium for a species. The cluster is the closest approximation to an individual in the environment,
and the number of clusters provided more valuable information when used with measures of abundance or
productivity, but was not as meaningful alone. However, the number of clusters was underestimated for gregarious species.
Two hundred six species of Basidiomycetes and Ascomycetes were collected in a coastal sand dune/beach
pine ecosystem primarily composed of Pinus contorts var. contorts, Vaccinium ovatum, Gaultherin shallon,
Arctostaphylos uva-ursi and herbaceous dune species. A distinct succession of 43 dominant, subdominant and common species was observed from September to July and was correlated with precipitation and temperature. The succession could be divided into 3 general patterns: 1) the initiation of the flora occurred when precipitation was between 0.0 and 5.0 cm and when maximum temperatures were beteween 22 and 25° C and when minimum temperatures ranged from 4.2 to 6.5° C; the major portion of the flora appeared in the second week of October and lasted
until the end of January. Fruiting of these species began when rainfall totaled between 5.0 and 24.3 cm the first year, and between 3.5 and 19.6 cm the second year. Cooler 97 maximum and minimum temperatures, between 17.9 and 21.1° C and -0.1 and 4.4° C, respectively may also have stimulated fruiting during this time. Successive frosts from late November through the end of January ended the season for the majority of species; 3) low minimum temperatures between -0.9 and 2.5° C stimulated fruiting of a few cold-tolerant species during late November through the end of January. Lack of competition from other fungi may also explain the appearance of these species during the winter. Many species continued to fruit or refruited during spring when minimum temperatures ranged between 0.9 and 10.1° C. High maximum temperatures between 20.0 and 24.1° C brought the spring season to an end for all species except Cantharellus cibarius, which produced a flush of sporocarps in June and July. Of the 206 species, six were considered dominant (defined by 5 percent of the total number of sporocarps or total number of clusters), 17 were determined as subdominant (defined by 2-4.9 percent of the total number of sporocarps or clusters and by 40 or more occurrences), and 20 were defined as common species (based on 15 or more occurrences, but comprising less than 2 percent). These 43 dominant, subdominant and common species comprised 21 percent of the flora. This percentage of the flora was lower than in 4 previous studies due to greater number of species recorded. 98
Five patterns of seasonal periodicity and six patterns of abundance were observed for the dominant, subdominant, and common species. These patterns may be confirmed or may change with additional years of observations. Thirty-nine of the 43 species were mycorrhizal, and were found to be associated mostly with pine and to a lesser degree with huckleberry and bearberry. Four of the
43 species were saprophytic species and were distributed among five different plant communities.
From this ecological survey, numerous questions arose, some of the more important of which are: 1) Will careful observation and mapping of clusters in a smaller permanent plot yield information about the response of species to different sets of environmental conditions as in a larger
plot?; 2) Is cluster size the best method for correlating environmental factors with the formation of fungal sporocarps?; 3) Do the patterns observed for seasonal duration and abundance remain the same from year to year for individual species?; 4) Why do these patterns differ
among species?; 5) What are the specific adaptations that may explain these patterns?; 6) What environmental
evidence is available to explain speciation in the fungi? 99
APPENDIX A
Synecological terms. The following definitions were taken from Cain & Castro (1968).
1. Abundance: a quantitative measure of number of individuals derived from actual counts per unit area.
2. Dominance: the relative degree to which a kind of plant predominates in the community, usually expressed as cover classes. 3. Frequency: refers to the portion of units of a sample in which at least one plant of a species is present.
4. Sociability: the degree to which plants are normally aggregated in nature, usually expressed as classes.
5. Periodicity: refers to the periods of the year during which a plant is vegetatively or reproductively active.
6. Constancy: is the percentage occurrence of a species on samples of the same size in various units of the community type. 7. Fidelity: is the degree of exclusiveness that a species shows in a given community type.
8. Productivity: the rate at which plants assimilate energy, usually measured as biomass. 100
APPENDIX B
Plant communities. The dominant species found in each community according to Barker (1976).
Community Plant Group Dominant species
Littoral grasses 1. Ammophila arenaria (L.) Link
strip 2. Elymus mollis Trin. ex. Spreng.
3. Poa douglasii Nees ssp. douglasii
herbs 1. Abronia latifolia Eschs.
2. Ambrosia chamissonis (Less.)Greene 3. Cakile maritima Scop. 4. Calystegia soldanella (L.) R.Br.
5. Lathyrus littoralis
(Nutt.ex.T.& G.) Endl.
Established grasses 1. Poa confinis Vasey
dunes 2. Poa douglasii Nees.
var. macrantha (Vasey) Keck.
herbs 1. Abronia latifolia Eschs.
2. Achilles borealis Bong.
ssp. arenicola (Heller)Keck. 3. Ambrosia chamissonis (Less.)Greene 4. Calystegia soldanella (L.) R.Br.
5. Camissonia cheiranthifolia
(Hornem.) Raimann
6. Carpobrotus chilense N.E.Br.
7. Erigeron glaucus Ker.
8. Eriogonum latifolium Sm. in Rees. 101
9. Fragaria chiloensis (L.)Duckn.
10. Lathyrus littoralis
(Nutt. ex. T.& G.) Endl.
11. Polygonum paronychia
Cham. & Schlecht.
12. Solidago spathulata DC.
13. Tanacetum douglasii DC.
Vegetated sedges 1. Carex obnupta Bailey
hollows rushes 1. Juncus lesueurii 801.
herbs 1. Gnaphalium palustre Nutt.
2. Lotus micranthus Benth.
3. Lotus purshianus (Benth.)
Clem. & Clem.
4. Lupinus bicolor Lind. var. bicolor
5. Scrophularia californica Cham. &
Schlecht var. californica
shrubs 1. Baccharis pilularis DC.
ssp. consanquineus (DC.)C.B.Wolf.
2. Lupinus arboreus Sims.
3. Myrica californica Cham.& Schlecht
trees 1. Pinus contorts Doug. ex. Loud
var. contorts
Beach lichens 1. Cladina portentosa (Dufour) Follm. pine subsp. pacifica (Anti) Anti forest mosses 1. Dicranum scoparium Hedw.
2. Kindbergia oregana (Sull.)Ochyra '102
ferns 1. Pteridium aquilinum (L.)Kuhn.
var. pubescens Underw.
grasses 1. Hierchloe occidentalis Buckl.
herbs 1. Anaphalis margaritacea (L.) Benth. ex. Clarke
2. Satureja douglasii (Benth.)Briq.
3. Solidago spathulata DC.
shrubs 1. Arctostaphylos uva-ursi (L.)Spreng
var. coactilis Fern.&Macbr.
2. Garrya elliptica Dougl.
3. Gaultherin shallon Pursh.
4. Vaccinium ovatum Pursh.
trees 1. Pinus contorts Dougl. ex. Loud
var. contorts
Willow ferns 1. Polypodium glycyrrhiza DC. Eat.
swamps 2. Polypodium scouleri Hook.& Grey.
3. Pteridium aquilinum (L.) Kuhn
var. pubescens Underw.
sedges 1. Carex obnupta Bailey
herbs 1. Galium aparine L.
vines 1. Rubus ursinus Cham.& Schlecht.
shrubs 1. Lonicera involucrata
(Richards.) Banks
2. Myrica californica Cham.& Schlecht
3. Salix hookeriana Barr.
trees 1. Alnus oregana Nutt. 103
APPENDIX C
Sites and Site Codes. Site names and their alpha and numeric codes.
Site Name Alpha Numeric Site Name Alpha Numeric
Area I Backfield Bk 1 Myrica meadows Hr 18 Baler meadow Br 2 Owl meadows Om 19 Big Myrtles Mh 3 Perspiration ridge Pr 20 Big Myrtles meadow Hr 4 Piperic meadow Pm 21 Broken tree meadows Bt 5 Sea Turf St 22 Bunny rabbit hollow Bh 6 Sedge Swamp I Ss 23 Buried spruce meadow Bs 7 Sedge Swamp II Sw 24 Corral corners Cc 8 Site 1 51 25 Crooked Pine meadow Cp 9 Site 1 meadow SO 26 Driftwood meadows Dm 10 Site 2 S2 27 Jack rabbit hollow Jh 11 Site 3 53 28 Lone Pine meadow Lp 12 S fence meadow Sm,Se 29 Maze meadow Mm 13 South fence swamp Sf 30 Middle fence trail Mf 14 Southwest corner Sc 31 Mossy Nook Mk 15 Strawberry glutch Sg 32 Mouse mound meadow N Mn 16 Tallyho road Tr 33 Mouse mound meadow S Ms 17 Three corners 3c 34 Weott corral Wc,Gf 35
Area II Behind house Br 36 Site 6 56 43 Busy Forest Bf 37 Site 7 37 44 Fireweed ridge Fr 38 Site 8 38 45 Myrtle hollow 56 39 Tallyho trail Tt 46 New Trail Nt 40 Twin spruce E-W Te 47 North meadow Nm 41 Twin spruce N-S Tn 48 Roller coaster trail Re 42 Twin spruce Ts 49
Area III Back Dune Hollow Dh 50 N Site 5 hollow N5 58 Bee hollow Bd 51 North trail Nt 59 Beg. Forest trail Ft 52 Pampas hollow Ph 60 Foredunes Fd 53 Rifle range Rr 61 Middle dune trail Mt 54 Site 5 hollow 55 62 Middle hollow Md 55 South foredunes Sf 63 North foredunes Nf 56 S Site 5 hollow D5 64 North marsh Nd 57 Outback Ob 65 104
APPENDIX D
Species of Macrofungi at the Lanphere-Christensen Dunes Preserve, Arcata, California.
A total of 206 species of macrofungi were collected from the Dunes Preserve. Of these, 182 species, or 90 percent of the flora, were found the first year and 11 species (4 percent) were discovered the second year.
During the following season, 1981-1982, an additional 13 species, representing 6 percent of the flora, were collected.
Eighty-three of the 206 fungi were mycorrhizal species. Sixty-nine of the 99 saprophytic species grew terrestrially, 26 were lignicolous, and 4 species decomposed needles or leaves. There were 4 pathogenic species of macrofungi and 1 lichenized Basidiomycete.
The majority of fungi belonged to the class of
Basidiomycetes. There were only 6 species of Ascomycetes.
Information about individual species follows the format below:
1. Species name, author 2. Collection numbers 3. Authority who verified identification: David L. Largent (D.L.), Orson K. Miller (0.M.), Joseph Ammirati (J.A.), Harry Theirs (H.T.) 4. Growth habit 5. Ecological function 6. Habitat on preserve 7. Dates collected 8. Associated phanerogams 105
SPECIES LIST
1. Agaricus campestris Fries 2. 231 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. grassy area surrounding preserve parking lot 7. 10 Nov. '79 8. Rubus ursinus, Pteridium aquilinum, grasses 1. Agaricus cf. haemorrhoidarius Schulzer 2. 164 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. moist depression in forest 7. 1 Nov. '79 8. Pinus contorts, Myrica californica, Vaccinium ovatum, Rubus ursinus, Carex obnupta
1. Agaricus perraris Schulzer 2. 245 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest beneath spruce 7. 11 Nov. '79 8. Picea sitchensis
1. Agaricus cf. rodmani 2. 216 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. vegetated dune hollow 7. 7 Nov. '79 8. Pinus contorts, Baccharis pilularis, Juncus lesueurii 1. Agaricus cf. xanthodermis Genevier 2. 384, 493 3. D. Largent 4. solitary to scattered 5. saprophytic, terrestrial 6. foredunes, established dunes 7. 30 Dec. '79, 30 Nov. '80 8. Achilles borealis, Elymus mollis, Erigeron glaucus, Eriogonum latifolium, Fragaria chiloensis, Lathyrus littoralis, Juncus lesueurii, Solidago spathulata 106
1. Agaricus cf. xanthodermis Genevier 2. 384, 493 3. D. Largent 4. solitary to scattered 5. saprophytic, terrestrial 6. established dune ridges and littoral strip 7. 30 Dec. '79, 30 Nov. '80 8. Erigeron glaucus, Solidago spathulata, Eriogonum latifolium, Juncus lesueurii, Achilles borealis, Fragaria chiloensis, Lathyrus littoralis, Elymus mollis.
1. Aleuria rhenana Fuckel 2. 720 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest among pine duff. 7. 12 Jan. '82 8. Pinus contorts, Vaccinium ovatum
1. Amanita aspera Fr. 2. 183, 315 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 1 Nov. '79, 29 Nov. '79 8. Pinus contorts, Picea sitchensis, Myrica californica, Arctostaphylos uva-ursi, Garrya elliptica, Vaccinium ovatum, Solidago spathulata, Anaphalis margaritacea
1. Amanita gemmata (Fr.) Gill. 2. 253, 422d, 533 3. D. Largent 4. solitary 5. mycorrhizal 6. forest and vegetated dune hollows 7. 12 Nov. '79, 4 May, '80, 28 Mar. '81 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Juncus lesueurii, Pteridium aquilinum
1. Amanita muscaria (L. ex. Fr.) Hooker 2. 62, 197 3. D. Largent 4. solitary, scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 16 Oct. '79, 4 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultherin shallon, Garrya elliptica, Salix hookeriana, Solidago spathulata, Carex obnupta 107
1. Amanita pantherina (Dc. ex. Fr.) Seer. 2. 330, 390, 396, 398, 401, 403 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 6 Dec. '79, 3 Jan. '80, 14 Jan. '80, 21 Jan. '80, 4 Feb. '80, 8 Feb. 80 8. Pinus contorts, Picea sitchensis, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Lonicera involucrata, Gaultherin shallon, Rubus ursinus, Solidago spathulata, Fragaria chiloensis, Pteridium aquilinum, Carex obnupta
1. Amanita porphyria (Alb. & Schw. ex. Fr.) Seer. 2. 50, 187, 422f 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest, usually near Myrica californica 7. 15 Oct. '79, 1 Nov. '79, 25 May '80 8. Pinus contorts, Picea sitchensis, Myrica californica, Gaultherin shallon, Vaccinium ovatum, Rubus ursinus
1. Armillaria mellea (Vahl ex. Fr.) Karst. 2. 334, 705 3. D. Largent 4. caespitose clusters 5. parasitic - base of Salix hookeriana 6. willow swamps, edge of moving dunes and willow swamps 7. 6 Dec. '79, 20 Nov. '81 8. Salix hookeriana
1. Armillaria ponderosa (Pk.) Sacc. 2. 207, 210, 643, 684 3. D. Largent 4. solitary to gregarious 5. mycorrhizal 6. forest under Vaccinium 7. 5 Nov. '79, 5 Nov. '81 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultherin shallon, Solidago spathulata, Goodyera oblongifolia, Anaphalis margaritacea, Pteridium aquilinum 108
1. Armillaria robusta Fr. 2. 97, 106 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. vegetated hollows, forest 7. 23 Oct. '79, 25 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Solidago spathulata, Juncus lesueurii, Vaccinium ovatum, Cladina portentosa
1. Armillaria zelleri Stuntz & A.Η. Smith 2. 204, 278 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest 7. 4 Nov. '79, 26 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Solidago spathulata, Vaccinium ovatum, Cladina portentosa, Anaphalis margaritacea
1. Baeospora myosura (Fr.) Sing. 2. 91, 92 3. D. Largent 4. gregarious 5. saprophytic on spruce cones 6. forest on spruce cones 7. 8. Picea sitchensis
1. Bolbitius vitellinus (Pers. ex. Fr.) Fr. 2. 227, 242 3. D. Largent 4. solitary to scattered 5. saprophytic on manured grass 6. forest, formerly grazed meadows 7. 8 Nov. '79, 10 Nov. '79 8. Rumex acetosella, Juncus lesueurii, grasses
1. Boletus amyloideus Theirs 2. 58, 88 3. D. Largent, Η. Theirs 4. solitary to scattered 5. mycorrhizal 6. vegetated dune hollows 7. 16 Oct. '79, 23 Oct. '79 8. Pinus contorts, Salix hookeriana, Arctostaphylos uva-ursi, Vaccinium ovatum, Garrya elliptica, aquilliniumGautheria shallon, Solidago spathulata, Pteridium 109
1. Boletus chrysenteron Bull. ex. St-Amans 2. 145, 147, 148, 150 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. vegetated dune hollows 7. 30 Oct. '79 8. Pinus contorts, Salix hookeriana, Arctostaphylos uva-ursi, Lupinus arboreus, Solidago spathulata, Fragaria chiloensis, Anaphalis margaritacea, Achillea borealis, Carex obnupta, Juncus lesueurii
1. Boletus edulis Bull. ex. Fr. 2. 126, 135 3. D. Largent 4. solitary 5. mycorrhizal 6. forest and vegetated dune hollows 7. 28 Oct. '79, 29 Oct. '79 8. Pinus contorts, Vaccinium ovatum, Arctostaphylos uva-ursi, Cladina portentosa, Solidago spathulata, Picea sitchensis, Salix hookeriana, Myrica californica, Carex obnupta
1. Boletus piperatus Bull. ex. Fr. 2. 149 3. D. Largent 4. solitary 5. mycorrhizal 6. vegetated dune hollows and forest sedge swamps 7. 30 Oct. '79 8. Pinus contorts, Myrica californica, Salix hookeriana, Lonicera involucrata, Rubus ursinus
1. Boletus zelleri Murr. 2. 229, 462 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 8 Nov. '79, 12 Nov. '80 8. Pinus contorta, Picea sitchensis, Myrica californica, Vaccinium ovatum
1. Bovista pila Berk. & Curt. 2. 424, 429, 518 3. 0. Miller 4. solitary to scattered 5. saprophytic 6. established dune ridges 7. 28 July '80, 17 Aug. '80, 7 Feb. 81 (fresh) 8. Solidago spathulata, Erigeron glaucus, Eriogonum latifolium, Fragaria chiloensis, Daucus pusillus, Carpοbrotus chilense 110
1. Calocera viscosa (Pers. ex. Fr.) Fr. 2. 100 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 25 Oct. '79
1. Cantharellus cibarius Fr. 2. 1, 4, 5, 7, 425 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest 7. 2 Sept. '79, 28 July '80 8. Pinus contorts, Picea sitchensis, Myrica californica, Vaccinium ovatum, Arctostaphylos uva-ursi, Gaultherin shallon, Solidago spathulata
1. Chroogomphus vinicolor (Pk.) O.K. Miller 2. 63, 103 3. 0. Miller 4. gregarious 5. mycorrhizal 6. vegetated dune hollows and forest 7. 17 Oct. '79, 25 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Vaccinium ovatum, Carex obnupta, Cladina portentosa
1. Clavaria rugosa 2. 234, 255 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 10 Nov. '79, 14 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Garrya elliptica, Gaultherin shallon, Vaccinium ovatum, Pteridium aquilinum
1. Clavulina sp. Schroet. 2. 511 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 13 Jan. '81 8. Pinus contorts, Gaultherin shallon, Vaccinium ovatum 111
1. Clavulinopsis corniculate (Schaeff. ex. Fr.) 2. 346, 385 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 7 Dec. '79, 3 Jan. '80 8. Pinus contorts, Picea sitchensis, Abies grandis, Vaccinium ovatum, Garrya elliptica, Gaultheria shallon, Solidago spathulata, Rubus ursinus, Satureja douglasii, Polypodium glycyrrhiza
1. Clavulinopsis gracillimis (Pk.) Petersen 2. 325, 341 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 4 Dec. '79, 7 Dec. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium_ ovatum, Solidago spathulata, Fragaria chiloensis
1. Clavulinopsis laeticolor (Berk. & Curt.) Petersen 2. 289 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 28 Nov. '79 8. Pinus contorts, Vaccinium ovatum, Gaultherin shallon
1. Clitocybe albirhiza Bigelow & Smith 2. 400, 520, 523a 3. 0. Miller 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 4 Feb. '80, 8 Feb. '81, 15 Feb. '81 8. Pinus contorts, Picea sitchensis, Abies grandis, Myrica californica, Garrya elliptica, Vaccinium ovatum, Satureja douglasii, Pteridium aquilinum
1. Clitocybe clavipes (Pers. ex. Fr.) Kummer 2. 226 3. D. Largent 4. scattered 5. mycorrhizal 6. forest sedge swamp 7. 8 Nov. '79 8. Pinus contorts, Picea sitchensis, Carex obnupta 112
1. Clitocybe fragrans (Fr.) Kummer 2. 503 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. vegetated dune hollows 7. 7 Dec. '80 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Solidago spathulata, Fragaria chiloensis, Juncus lesueurii
1. Clitocybe inversa (Fr.) Quel. 2. 531, 621 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 3 Mar. '81, 1 Nov. '81 8. Pinus contorts, Picea sitchensis, Vaccinium ovatum, Solidago spathulata
1. Clitocybe smaragdina (Berk.) Bigelow & Smith 2. 719 3. Η. Theirs 4. scattered to gregarious 5. saprophytic, terrestrial 6. forest meadow 7. 15 Jan. '82 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Polygonum paronychia
1. Clitopilus prunulus (Scop. ex. Fr.) Kummer 2. 66, 84, 598, 692, 687 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest meadows and sedge swamps 7. 19 Oct. '79, 21 Oct. '79, 11 Oct. '81, 5 Nov. '81 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Garrya elliptica, Gaultherin shallon, Solidago spathulata, Rubus ursinus, Goodyera oblongifolia, Carex obnupta
1. Collybia cf. butyracea (Bull. ex. Fr.) Kummer 2. 509, 340 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 6 Jan. '81, 7 Dec. '79 8. Pinus contorts, Picea sitchensis, Myrica californica, Garrya elliptica, Vaccinium ovatum,Arctostaphylos uva-ursi, Solidago spathulata, Pteridium aquilinum 113
1. Collybia dryophila (Bull. ex. Fr.) Kummer 2. 25, 43, 164, 439 3. D. Largent 4. solitary to gregarious 5. saprophytic, terrestrial 6. forest 7. 5 Oct. '79, 15 Oct. '79, 31 Oct. '79, 19 Oct. '80 8. Pinus contorts, Picea sitchensis, Myrica californica, Salix hookeriana, Vaccinium ovatum, Garrya elliptica, Gaultherin shallon, Solidago spathulata
1. Collybia fuscopurpurea (Pers. ex. Fr.) Kummer 2. 98, 177 3. D. Largent 4. solitary to gregarious 5. saprophytic, terrestrial 6. forest 7. 23 Oct. '79, 1 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Garrya elliptica, Vaccinium ovatum, Gaultheria shallon, Solidago spathulata, Anaphalis margaritacea, Goodyera oblongifolia, Pteridium aquilinum
1. Collybia luteoolivaceum (Berk.) 2. 239, 298 3. D. Largent 4. scattered to gregarious 5. saprophytic, terrestrial 6. forest 7. 10 Nov. '79, 28 Nov. '79 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultherin shallon, Garrya elliptica, Pteridium aquilinum
1. Coltrichia perennis (L. ex. Fr.) Murr. 2. 538 3. D. Largent 4. scattered 5. parasitic 6. forest 7. 7 April '81 8. Pinus contorts, Arctostaphylos uva-ursi, Gaultherin shallon, Vaccinium ovatum
1. Conocybe sp. (Fr.) Staude 2. 295 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 28 Nov. '79 114
1. Conocybe sp. (Fr.) Staude 2. 391 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 3 Jan. '80 8. Myrica californica, Vaccinium ovatum, Gaultherin shallon
1. Conocybe sp. (Fr.) Staude 2. 402 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 4 Feb. '80 8. Pinus contorts, Myrica californica, Vaccinium ovatum, Solidago spathulata
1. Conocybe sp. Fayod 2. 4 14 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 8 Mar. '80 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Garrya elliptica, Solidago spathulata, Lotus micranthus
1. Conocybe cf. macrorrhiza (Speg.) Sing. 2. 411 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. dune hollow 7. 29 Feb. '80 8. Solidago spathulata, Achilles borealis, Eriogonum latifolium, Juncus lesueurii
1. Coprinus micaceus (Bull. ex. Fr.) Fr. 2. 465 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 14 Nov. '80 115
1. Coprinus sp. (Pers. ex. Fr.) S.F. Gray 2. 254 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest meadow 7. 14 Nov. '79 8. Garrya elliptica, Vaccinium ovatum, Solidago spathulata, Fragaria chiloensis, Rumex acetosella, Hierchloe occidentalis 1. Cortinarius aureofolius Peck var. hesperius Ammirati 2. 709 3. J. Ammirati 4. gregarious 5. mycorrhizal 6. open meadow in forest 7. 20 Nov. '81 8. Pinus contorts, Fragaria chiloensis 1. Cortinarius collinitus (Fr.) S.F. Gray 2. 222, 225 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 8 Nov. '79 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum 1. Cortinarius cf. glandicolor (Fr.) Fr. 2. 333 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 6 Dec. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Carex obnupta, Juncus lesueurii
1. Cortinarius humboldtensis Ammirati 2. 612, 702, 714 3. J. Ammirati 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 24 Oct. '81, 20 Nov. '81, 29 Nov. '81 8. Pinus contorts, Arctostaphylos uva-ursi, Garrya elliptica, Gaultherin shallon, Vaccinium ovatum, Solidago spathulata 116
1. Cortinarius oregonensis Ammirati 2. 274 3. J. Ammirati 4. gregarious 5. mycorrhizal 6. forest 7. 26 Nov. '79 8. Pinus contorts, Vaccinium ovatum
1. Cortinarius pacificus Ammirati 2. 221, 228, 249, 279, 286, 317, 392, 498, 663 3. J. Ammirati 4. scattered to gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 8 Nov. '79, 12 Nov. '79, 26 Nov. '79, 27 Nov. '79, 14 Jan. '80, 4 Dec. '80, 6 Dec. 80, 5 Nov. '81 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Garrya elliptica, Gaultherin shallon, Vaccinium ovatum, Salix hookeriana, Solidago spathulata, Fragaria chiloensis 1. Cortinarius phoeniceus Bull. ex. Maire var. occidentalis A.Η. Smith 2. 246, 273, 276, 310, 387, 389, 634, 681, 685 3. J. Ammirati 4. gregarious 5. mycorrhizal 6. forest 7. 10 Nov. '79, 26 Nov. '79, 29 Nov. '79, 3 Jan. '80, 5 Nov. '81 8. Pinus contorts, Myrica californica, Arctostaphylos_ uva-ursi, Garrya elliptica, Gaultherin shallon, Vaccinium ovatum, Solidago spathulata, Pteridium_ aquilinum 1. Cortinarius uliginosus Berkeley 2. 703 3. J. Ammirati 4. solitary 5. mycorrhizal 6. vegetated dune hollow 7. 20 Nov. '81 8. Salix hookeriana, Juncus lesueurii 1. Cortinarius vibratilis (Fr.) Fr. 2. 104, 107, 137 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 25 Oct. '79, 29 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum 117
1. Cortinarius sp. Fr. 2. 113, 294 3. D. Largent 4. scattered 5. mycorrhizal 6. vegetated dune hollow 7. 26 Oct. '79, 28 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Gaultherin shallon, Vaccinium ovatum, Lupinus arboreus, Solidago spathulata
1. Cortinarius sp. Fr. 2. 143 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest 7. 29 Oct. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Fragaria chiloensis, Pteridium aquilinum
1. Cortinarius sp. Fr. 2. 194 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 1 Nov. '79 8. Pinus contorts, Myrica californica
1. Cortinarius sp. Fr. 2. 245b 3. D. Largent 4. scattered 5. mycorrhizal 6. sedge swamp in forest 7. 11 Nov. '79 8. Pinus contorts, Myrica californica, Rubus ursinus, Carex obnupta
1. Cortinarius sp. Fr. 2. 294 3. D. Largent 4. scattered 5. mycorrhizal 6. forest ridge 7. 28 Nov. '79 8. Abies grandis 118
1. Cortinarius sp. Fr. 2. 328, 332 3. D. Largent 4. scattered 5. mycorrhizal 6. vegetated dune hollow 7. 6 Nov. '79 8. Pinus contorts, Salix hookeriana, Arctostaphylos uva-ursi, Lupinus arboreus, Carex obnupta, Juncus lesueurii
1. Cortinarius sp. Fr. 2. 369 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 13 Dec. '79 8. Pinus contorts, Garrya elliptica, Vaccinium ovatum
1.371 Cortinarius sp. Fr. 2. 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 13 Dec. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata
1. Cortinarius sp. Fr. 2. 373 3. D. Largent 4. caespitose 5. mycorrhizal 6. forest 7. 13 Dec. '79 8. Pinus contorts, Myrica californica, Vaccinium ovatum
1. Cortinarius sp. Fr. 2. 540 3. D. Largent 4. caespitose 5. mycorrhizal 6. vegetated dune hollow 7. 22 April '81 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Solidago spathulata, Juncus lesueurii, Lotus micranthus, Eriogonum latifolium 119
1. Crepidotus haustellaris (Fr. ex. Fr.) Kummer 2. 39, 93, 252, 463, 707 3. D. Largent 4. gregarious 5. saprophytic, lignicolous on Rubus 6. forest and vegetated dune hollows 7. 13 Oct. '79, 23 Oct. '79, 12 Nov. '79, 12 Nov. '80, 20 Nov. '81
1. Crinipellis piceae Sing. 2. 9 3. D. Largent 4. gregarious 5. saprophytic, on Picea needles 6. forest 7. 2 Sept. '79 1. Cudonia circinans Fr. 2. 512 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 13 Jan. '81 8. Pinus contorts, Picea sitchensis, Garrya elliptica, Gaultherin shallon, Vaccinium ovatum
1. Cystoderma amianthinum (Scop. ex. Fr.) Fayod 2. 313 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 29 Nov. '79 8. Pinus contorts, Picea sitchensis, Arctostaphylos uva-ursi, Garrya elliptica, Vaccinium ovatum, Solidago spathulata
1. Cystoderma amianthinum (Fr.) Fayod var. rugoscrobiculatum (Lorinser) Smith & Singer 2. 144, 1461, 468 3. D. Largent 4. solitary to gregarious 5. saprophytic, terrestrial 6. forest 7. 29 Oct. '79, 10 Nov. '80, 19 Nov. '80 8. Pinus contorts, Picea sitchensis, Arctostaphylos uva-ursi, Solidago spathulata, Rubus ursinus, Juncus lesueurii 120
1. Cystoderma cinnabarinum (Alb. & Schw. ex. Seer.) Fayod 2. 111, 129, 175a, 212 3. D. Largent 4. scattered to gregarious 5. saprophytic, terrestrial 6. forest meadows 7. 25 Oct. '79, 29 Oct. '79, 1 Nov. '79, 7 Nov. '79 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Garrya elliptica, Baccharis pilularis, Vaccinium ovatum, Solidago spathulata, Rumex acetosella, Anaphalis margaritacea, grasses
1. Cystoderma granulosum (Batsch ex. Fr.) Fayod 2. 383 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest meadow 7. 3 Jan. '80 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Gaultherin shallon, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum
1. Dacrymyces palmatus (Schw.) Bres. 2. 312, 386 3. D. Largent 4. gregarious 5. saprophytic, lignicolous on Pinus 6. forest 7. 29 Nov. '79, 3 Jan. '80
1. Disciseda subterranea (Pk.) Coker & Couch 2. 431, 593 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. established foredunes 7. 17 Aug. '80, 4 Oct. '81 8. Erigeron glaucus, Gnaphalium purpureum, Solidago spathulata, Abronia latifolia, Eriogonum latifolium, Fragaria chiloensis
1. Entoloma rhodopolium (Fr.) Kummer 2. 466 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. sedge swamp in forest 7. 14 Nov. '80 8. Pinus contorts, Picea sitchensis, Myrica californica, Carex obnupta 121
1. Exidia glandulosa Fries 2. 422b, 530 3. D. Largent 4. gregarious 5. saprophytic, lignicolous on Pinus 6. forest 7. 22 April '80, 3 Mar. '81 1. Fistulina hepatica Schaeff. ex. Fr. 2. 189 3. D. Largent 4. solitary to scattered 5. saprophytic, lignicolous on Myrica 6. forest 7. 1 Nov. '79
1. Flammulina velutipes (Fr.) Karst. 2. 213 3. D. Largent 4. gregarious 5. saprophytic, lignicolous on Lupinus arboreus_ 6. vegetated dune hollows 7. 7 Nov. '79 1. Galerina sp. Earle 2. 128 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 29 Oct. '79 1. Galerina sp. Earle 2. 251 3. D. Largent 4. gregarious 5. saprophytic, terrestrial in moss 6. vegetated dune hollow 7. 12 Nov. '79 8. Pinus contorts, Salix hookeriana, Carex obnupta, Juncus lesueurii, Fragaria chiloensis
1. Galerina sp. Earle 2. 335 3. D. Largent 4. gregarious 5. saprophytic, terrestrial in mosses 6. forest 7. 6 Dec. '79 8. Pinus contorts, Picea sitchensis, Myrica californica, Vaccinium ovatum, Gaultheria shallon 122
1. Galerina sp. Earle 2. 209 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 5 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Garrya elliptica, Gaultheria shallon
1. Galerina sp. Earle 2. 399 3. D. Largent 4. solitary 5. saprophytic, terrestrial in moss 6. forest meadow 7. 21 Jan. '80 8. Arctostaphylos uva-ursi, Juncus lesueurii, Rumex acetosella, grasses
1. Geastrum floriforme Vitt. 2. 430, 527 3. 0. Miller 4. scattered 5. saprophytic, terrestrial in sand 6. vegetated dune hollows and established dune ridges 7. 17 Aug. '80, 1 Mar. '81 8. Erigeron glaucus, Solidago spathulata, Eriogonum latifolium
1. Gomphideus glutinosus (Schaeff. ex. Fr.) Fr. 2. 244 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 11 Nov. '79 8. Pinus contorts, Picea sitchensis, Abies grandis, Garrya elliptica, Vaccinium ovatum
1. Gomphideus subroseus Kauff. 2. 460, 648 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 10 Nov. '80, 5 Nov. '81 8. Pinus contorts, Picea sitchensis, Pseudotsuga menziesii, Arctostaphylos uva-ursi, Garrya elliptica, Vaccinium ovatum, Gaultheria shallon 123
1. Guepiniopsis alpinus (Tracy & Earle) Bres. 2. 2 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 2 Sept. '79
1.Gymnopilus penetrans (Fr. ex. Fr.) Murr. 2. 208,23 6 3. D. Largent 4. scattered to gregarious 5. saprophytic, lignicolous 6. forest 7. 5 Nov. '79, 1 Nov. '81
1. Gyromitra infula (Schaeff. ex. Fr.) Quel. 2. 715 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. forest 7. 19 Nov. '81 8. Pinus contorts, Vaccinium ovatum, Garrya elliptica, Gaultheria shallon
1. Hebeloma crustuliniforme (Bull. ex. St. Amans) Quel. 2. 118, 159 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. vegetated dune hollows 7. 26 Oct. '79, 30 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Solidago spathulata, Carex obnupta, Juncus lesueurii, Fragaria chiloensis
1. Hebeloma sp. (Fr.) Kummer 2. 170 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 31 Oct. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum 124
1. Hebeloma sp. (Fr.) Kummer 2. 248, 458, 500 3. D. Largent 4. gregarious 5. mycorrhizal 6. vegetated dune hollows 7. 12 Nov. '79, 9 Nov. '80, 7 Dec. '80 8. Pinus contorts, Salix hookeriana, Lupinus arboreus, Anaphalis margaritacea, Carex obnupta, Juncus lesueurii, Fragaria chiloensis
1. Hebeloma sp. (Fr.) Kummer 2. 292 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 28 Nov. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Pteridium aquilinum
1. Hebeloma sp. (Fr.) Kummer 2. 523 3. D. Largent 4. scattered 5. mycorrhizal 6. vegetated dune hollow 7. 15 Feb. '81 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Juncus lesueurii, Fragaria chiloensis
1. Helvella lacunosa Afzelius ex. Fr. 2. 366 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 10 Dec. '79 8. Pinus contorts, Picea sitchensis, Garrya elliptica, Vaccinium ovatum
1. H dnellum scrobiculatum (Fr. ex. Secr.) Karst. 2. 1 30,y 444 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest 7. 29 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Anaphalis margaritacea, Cladina portentosa 125
1. Hydnum scabrosum Fr. 2. 138, 224, 370 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 29 Oct. '79, 8 Nov. '79, 13 Dec. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum
1. Hygrophoropsis aurantiacus (Wulfen ex. Fr.) Maire 2. 15, 18, 40, 89, 127 3. D. Largent 4. scattered to gregarious 5. saprophytic, terrestrial 6. forest 7. 21 Sept. '79, 28 Sept. '79, 13 Oct. '79, 23 Oct. '79, 29 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Rubus ursinus, Pteridium aquilinum
1. Hygrophorus agathosmus (Fr.) Fr. 2. 406, 524 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 23 Feb. '80, 22 Feb. '81 8. Pinus contorts, Picea sitchensis, Vaccinium ovatum
1. Hygrophorus conicus (Fr.) Fr. 2. 309 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 29 Nov. '79 8. Pinus contorts, Garrya elliptica, Vaccinium ovatum, Solidago spathulata, Satureja douglasii, Pteridium aquilinum
1. Hygrophorus psittacinus (Fr.) Fr. 2. 318, 504 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 4 Dec. '79, 8 Dec. '80 8. Pinus contorts, Garrya elliptica, Vaccinium ovatum 126
1. Hygrophorus sp. Fr. 2.3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 8 Dec. '79 8. Picea sitchensis, Garrya elliptica, Vaccinium ovatum, Lupinus arboreus, Anaρhalis margaritacea, Satureja douglasii, Pteridium aquilinum 1. Inocybe geophylla var. lilacina Pk. 2. 375, 528 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 13 Dec. '79, 3 Mar. '81 8. Picea sitchensis, Gaultheria shallon, Vaccinium ovatum 1. Inocybe lacers (Fr.) Kummer 2. 78, 144, 171, 415, 422e 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 19 Oct. '79, 29 Oct. '79, 31 Oct. '79, 8 Mar. '80, 4 May '80 8. Picea sitchensis, Arctostaphylos uva-ursi, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Salix hookeriana, Solidago spathulata, Fragaria chiloensis, Carex obnupta, Juncus lesueurii, Pteridium aquilinum 1. Inocybe sp. (Fr.) Fr. 2. 290, 351, 487 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest 7. 28 Nov. '79, 8 Dec. '79, 24 Nov. '80 8. Pinus contorts, Arctostaphylos uva-ursi, Garrya elliptica, Vaccinium ovatum, Gaultheria shallon, Ribes sanguineum, Solidago spathulata, Pteridium aquilinum
1. Inocybe sp. (Fr.) Fr. 2. 83 3. D. Largent 4. solitary 5. mycorrhizal 6. forest meadow 7. 15 Oct. '79 8. Pinus contorts, Vaccinium ovatum 127
1. Laccaria bicolor (R. Mre.) Orton 2. 374 3. 0. Miller 4. gregarious 5. mycorrhizal 6. forest 7. 15 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Baccharis pilularis, Vaccinium ovatum, Solidago spathulata, Anaphalis margaritacea
1. Laccaria laccata (Fr.) Berk. & Br. 2. 34, 186, 241, 247, 288, 372 3. 0. Miller 4. gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 13 Oct. '79, 1 Nov. '79, 10 Nov. 1 79, 12 Nov. '79, 28 Nov. '79, 13 Dec. '79 8. Pinus contorts, Picea sitchensis, Myrica californica, Vaccinium ovatum, Gaultheria shallon, Salix hookeriana, Lupinus arboreus, Achilles borealis, Juncus lesueurii
1. Laccaria proxima (Boud.) Pat. 2. 191, 499, 516 3. 0. Miller 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 1 Nov. '79, 6 Dec. '80, 25 Jan. '81 8. Pinus contorts, Myrica californica, Garrya elliptica, Vaccinium ovatum, Carex obnupta
1. Lactarius deliciosus (L. ex. Fr.) S.F. Gray 2. 69, 76, 110, 142, 543, 691 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 19 Oct. '79, 25 Oct. '79, 29 Oct. '79, 22 Apr. '81, 5 Nov. '81 8. Pinus contorts, Myrica californica, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Anaphalis margaritacea 128
1. Lactarius pseudomucidus Resler and Smith 2. 299, 321, 359 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 28 Nov. '79, 4 Dec. '79, 8 Dec. '79 8. Pinus contorts, Picea sitchensis, Abies grandis, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum 1. Lactarius rufus (Scop. ex. Fr.) Fr. 2. 192, 245a, 419 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 1 Nov. '79, 11 Nov. '79, 6 Apr. '80 8. Pinus contorts, Myrica californica, Vaccinium ovatum, Rubus ursinus, Carex obnupta 1. Lactarius subflammeus sp. nov. A.H. Smith 2. 323, 451, 486, 497 3. 0. Miller 4. gregarious 5. mycorrhizal 6. forest 7. 4 Dec. '79, 27 Oct. '80, 24 Nov. '80, 6 Dec. '80 8. Pinus contorts, Picea sitchensis, Myrica californica, Garrya elliptica, Vaccinium ovatum, Lonicera_ involucrata, Carex obnupta
1. Leccinum largentii Theirs 2. 599 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 11 Oct. '81 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Anaphalis margaritacea, Cladina portentosa 1. Leccinum manzanitae Theirs 2. 168, 169, 344, 457 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest 7. 31 Oct. '79, 7 Dec. '79, 3 Nov. '80 8. Pinus contorts, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Anaphalis margaritacea, Pteridium aquilinum 129
1. Lepiota castanea Quel. 2. 287, 336 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 27 Nov. '79, 6 Dec. '79 8. Picea sitchensis, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Goodyera oblongifolia
1. Lepiota sp. Pers. ex. S.F. Gray 2. 120, 121, 153, 156, 162, 217, 438, 459 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. established dune ridges 7. 26 Oct. '79, 30 Oct. '79, 7 Nov. '79, 18 Oct. '80, 9 Nov. '80 8. Erigeron glaucus, Eriogonum latifolium, Solidago spathulata, Fragaria chiloensis, Carpobrotus chilense, Cakile maritima, Lathyrus littoralis, Ambrosia chamissonis, Elymus mollus
1. Lepiota sp. Pers. ex. S.F. Gray 2. 505, 510, 514 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. established dune ridges 7. 4 Jan. '81, 11 Jan. '81, 25 Jan. '81 8. Erigeron glaucus, Eriogonum latifolium, Solidago spathulata, Polygonum paronychia
1. Lepista nuda (Bull. ex. Fr.) Cke. 2. 243 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 11 Nov. '79 8. Pinus contorts, Picea sitchensis, Abies grandis, Garrya elliptica, Vaccinium ovatum, Pteridium aquilinum
1. Leptonia nigra (Murr.) Murr. 2. 181, 338 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 1 Nov. '79, 7 Dec. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium_ ovatum, Solidago spathulata, Pteridium aquilinum+ 130
1. Leptonia sarcinula Kuhner & Romagn. 2. 301 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 28 Nov. '79 8. Pinus contorts, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum
1. Lycoperdon cf. foetidum Bonorden 2. 305 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 29 Nov. '79 8. Pinus contorts, Abies grandis, Garrya elliptica, Solidago spathulata
1. Lycoperdon perlatum Pers. 2. 350, 620 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 7 Dec. '79 8. Pinus contorta, Myrica californica, Vaccinium ovatum, grasses
1. Lycoperdon cf. umbrinum Pers. 2. 534 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. established dune ridges 7. 29 Mar. '81 8. Erigeron glaucus, Eriogonum latifolium, Fragaria chiloensis, Achilles borealis, Claytonia perfoliata
Lycoperdon sp. Pers. 2. 96, 115 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. vegetated dune hollows 7. 23 Oct. '79, 26 Oct. '79 8. Pinus contorts, Arctostaphylos uva-ursi, Salix hookeriana, Lupinus arboreus, Solidago spathulata, Juncus lesueurii 131
1. Macowanites sp. Kalchb. 2. 545 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 16 May '81 8. Pinus contorta, Picea sitchensis, Myrica californica, Vaccinium ovatum
1. Marasmius magnisporus Murrill 2. 71 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 19 Oct. '79
1. Marasmius plicatulus Pk. 2. 304, 501 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest and established dune ridges 7. 29 Nov. '79, 7 Dec. '80 8. Pinus contorta, Abies grandis, Garrya elliptica, Solidago spathulata, Erigeron glaucus, Eriogonum latifolium, Fragaria chiloensis, Achilles borealis
1. Marasmius cf. scorodonius (Fr.) Fr. 2. 48 3. D. Largent 4. gregarious 5. saprophytic, on pine needles 6. forest 7. 15 Oct. '79
1. Melanoleuca melaleuca (Pers. ex. Fr.) Murr. 2. 306 3. D. Largent 4. scattered 5. mycorrhizal 6. forest meadow 7. 29 Nov. '79 8. Pinus contorta, Abies grandis, Garrya elliptica 132
1. Montagnea arenarius (DC.) Zeller 2. 753 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. established dune ridges 7. 3 Jun. '84 8. Erigeron glaucus, Eriogonum latifolium, Lathyrus littoralis, Fragaria chiloensis, Carpobrotus chilense
1. Mycena atromarginata (Lasch) Kummer 2. 30 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 13 Oct. '79 8. Pinus contorta, Myrica californica, Vaccinium ovatum, Rebus ursinus
1. Mycena aurantiomarginata (Fr.) Quel. 2. 260 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 14 Nov. '79 8. Pinus contorta, Picea sitchensis, Garrya elliptica, Vaccinium ovatum, Goodyera oblongifolia
1. Mycena capillaripes Peck 2. 616 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 25 Oct. '81 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum
1. Mycena clavicularis (Fr.) Gill. 2. 42 ,200, 594 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 15 Oct. '79, 4 Nov. '79, 11 Oct. '81 8. Pinus contorta, Abies grandis, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Calypso bulbosa, Goodyera oblongifolia, Pteridium aquilinum 133
1. Mycena epipterygia (Fr.) S.F. Gray var. lignicola Smith 2. 492 3. D. Largent 4, gregarious 5. saprophytic, lignicolous 6. forest 7. 26 Nov. '80 8. Picea sitchensis, Vaccinium ovatum
1. Mycena haematopus (Pers. ex. Fr.) Kummer 2. 708 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 20 Nov. '81 8. Pinus contorta, Myrica californica, Vaccinium ovatum
1. Mycena leptocephaly (Pers. ex. Fr.) Gill. 2. 31, 37 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest and vegetated dune hollows 7. 13 Oct. '79 8. Pinus contorta, Myrica californica, Vaccinium ovatum, Rubus ursinus
Μycena pura_ (Pers. ex. Fr.) Kummer 1. 2. 1 40 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 29 Oct. '79 8. Pinus contorta, Picea sitchensis, Arctostaphylos uva-ursi, Vaccinium ovatum, Anaphalis margaritacea
1. Mycena rorida (Scop. ex. Fr.) Quel. 2. 710 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. forest 7. 20 Nov. '81 134
1. Mycena subincarnata (Pk.) Sacc. 2. 45 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 15 Oct. '79 8. Pinus contorta, Garrya elliptica, Vaccinium ovatum, Gaultheria shallon
1. Naemataloma fasiculare (Fr.) Karst. 2. 108, 165, 201 3. D. Largent 4. caespitose 5. saprophytic, lignicolous 6. forest 7. 25 Oct. '79, 31 Oct. '79, 4 Nov. '79 1. Naucoria sp. (Fr.) Kummer 2. 25 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 19 Oct. '79 8. Pinus contorta, Myrica californica, Garrya elliptica, Vaccinium ovatum, Erechtites arguta, Maianthemum dilatatum
1. Nolanea cetrata (Fr. ex. Fr.) Kummer 2. 388, 615 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 3 Jan. '80, 25 Oct. '81 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Anaphalis margaritacea, Pteridium aquilinum
1. Nolanea sp. (Fr.) Kummer 2. 132, 378, 397 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest meadows 7. 29 Oct. '79, 30 Dec. '79, 14 Jan. '80 8. Pinus contorta, Arbutus menziesii, Garrya elliptica, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata 135
1. Nolanea sp. (Fr.) Kummer 2. 502 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest meadow 7. 7 Dec. '80 8. Pinus contorta, Baccharis pilularis, Lupinus arboreus, grasses
1. Omphalina ericetorum (Fr.) M. Lange 2. 36, 82, 105, 445, 529 3. D. Largent 4. gregarious 5. parasitic, lichenized 6. forest logs 7. 13 Oct. '79, 21 Oct. '79, 25 Oct. '79, 21 Oct. '80, 3 Mar. '81
1. Omphalina fibula Fr. 2. 232 3. D. Largent 4. gregarious 5. saprophytic, terrestrial in moss 6. forest 7. 10 Nov. '79 8. Pinus contorta, Vaccinium ovatum, Garrya elliptica, Arctostaphylos uva-ursi
1. Omphalina schwartzii Fr. 2. 220 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 8 Nov. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Gaultheria shallon, Pteridium aquilinum
1. Omphalina sp. Quel. 2. 174, 196, 198 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 31 Oct. '79, 4 Nov. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Pteridium aquilinum 136
1. Otidea leporina (Batsch ex. Fr.) Fuckel 2. 297, 322, 379, 407 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 28 Nov. '79, 4 Dec. '79, 30 Dec. '79, 23 Feb. '80 8. Pinus contorta, Picea sitchensis, Abies grandis, Gaultheria shallon, Vaccinium ovatum, Ribes sanguineum
1. Otidea sp. Fuckel 2. 303, 711 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. forest 7. 29 Nov. '79, 20 Nov. '81 8. Pinus contorta, Abies grandis, Garrya elliptica, Solidago spathulata
1. Paxillus involutus (Batsch ex. Fr.) Fr. 2. 151, 154, 470 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. vegetated dune hollows 7. 30 Oct. '79, 18 Nov. '81 8. Pinus contorta, Salix hookeriana, Lupinus arboreus, Carex obnupta, Fragaria chiloensis
1. Paxillus panuoides (Fr. ex. Fr.) Fr. 2. 316, 410 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. edge of forest 7. 4 Dec. '79, 23 Feb. '80 1. Phaeolus schweinitzii (Fr.) Patouillard 2. 8, 180 3. D. Largent 4. solitary 5. saprophytic, lignicolous 6. forest 7. 2 Sept. '79, 1 Nov. '79 137
1. Phallus hadriani (Venturi) Pers. 2. 146, 546 3. D. Largent 4. solitary to scattered 5. saprophytic, terrestrial 6. vegetated dune hollows, established dune ridges, littoral strip 7. 30 Oct. '79, 31 May '81 8. Salix hookeriana, Lupinus Arboreus, Solidago spathulata, Achilles borealis, Eriogonum latifolium, Erigeron glaucus, Fragaria chiloensis, Elymus mollis, Juncus lesueurii
1. Phellodon stratus K. Harrison 2. 706 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 20 Nov. '81 8. Pinus contorta, Myrica californica, Vaccinium ovatum, Carex obnupta
1. Phellodon melaleucus (Sw. spud Fr. ex. Fr.) Karst. 2. 230, 262 3. D. Largent 4. solitary to scattered 5. saprophytic, terrestrial 6. forest 7. 8 Nov. '79, 14 Nov. '79 8. Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Pteridium aquilinum
1. Pholiota velaglutinosa Smith and Resler 2. 347, 488, 495 4.3. D. Largent gregarious 5. saprophytic, lignicolous 6. forest 7. 7 Dec. '79, 24 Nov. '80, 1 Dec. '80 1. Phyllophorus rhodoxanthus (Schw.) Bres. 2. 5ΰ8, 536 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 6 Jan. '81, 5 Apr. '81 8. Pinus contorta, Picea sitchensis, Abies grandis, Pseudotsuga menziesii, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum 138
1. Plectania melastoma (Sow. ex. Fr.) Fuckel 2. 422c 3. D. Largent 4, scattered 5. saprophytic, lignicolous 6. forest 7. 22 Apr. '80
1. Pleurotus ostreatus (Jacq. ex. Fr.) Kummer 2. 739, 740 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. littoral strip 7. 27 May '82, 6 Jun. '82
1. Pluteus atromarginatus (Konrad) Kuhner 2. 101 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. forest 7. 25 Oct. '79
1. Podostroma alutacearum (Pers. ex. Fr.) Atkinson 2. 90 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 23 Oct. '79 8. Pinus contorta, Picea sitchensis, Vaccinium ovatum, Gaultheria shallon, Ribes sanguineum
1. Polyporus abietinus Dickson ex. Fr. 2. 6 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 2 Sept. '79
1. Polyporus caesius Schrad. ex. Fr. 2. 464 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. forest 7. 12 Nov. '80 139
1. Polyporus cf. elegans Bull. ex. Fr. 2. 11, 19 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. forest 7. 3 Sept. '79, 28 Sep. '79
1. Polyporus cf. osseus Kalchbr. 2. 436 3. D. Largent 4. scattered 5. saprophytic, lignicolous 6. forest 7. 3 Oct. '80
1. Psathyrella arenulina (Pk.) Sacc. 2. 326, 409, 412, 413, 521 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. established dune ridges, vegetated dune hollows, littoral strip 7. 6 Dec. '79, 23 Feb. '80, 29 Feb. '80, 8 Feb. '81
1. Psathyrella cf. hydrophila (Bull ex. Merat) Maire 2. 342, 364 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 7 Dec. '79, 10 Dec. '79 8. Pinus contorta, Picea sitchensis, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Sοlidagο spathulata
1. Psathyrella cf. longistriata (Murr.) Α.Η. Smith 2. 358 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest 7. 8 Dec. '79 8. Picea sitchensis, Vaccinium ovatum, Pteridium aquilinum 140
1. Rhizopogon occidentalis Zeller & Dodge 2. 68, 166, 380, 393, 395 3. D. Largent 4. solitary 5. mycorrhizal 6. forest and vegetated dune hollows 7. 19 Oct. '79, 31 Oct. '79, 30 Dec. '79, 14 Jan. '80 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Solidago spathulata, Eriogonum latifolium, Fragaria chiloensis, Carex_ obnupta, Juncus lesueurii, Pteridium aquilinum
1. Rhizopogon subcaerulescens Smith 2. 74, 254a 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 19 Oct. '79, 14 Nov. '79 8. Pinus contorta, Myrica californica, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Maianthemum dilatatum
1.Ripartites tricholoma (Fr.) Karsten 2. 704 3. J. Ammirati 4. gregarious 5. mycorrhizal 6. forest 7. 20 Nov. '81 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium_ ovatum
1. Russula strata Shaffer 2. 139, 237 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 29 Oct. '79, 10 Nov. '79 8. Pinus contorta, Picea sitchensis, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Solidago spathulat~, Pteridium aquilinum
1. Russula brevipes Pk. 2. 109, 133, 176, 236 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest 7. 25 Oct. '79, 29 Oct. '79, 1 Nov. '79, 10 Nov. '79 8. Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Garrya elliptica, Vaccinium ovatum, Gaultheria shallon 141
1. Russula brevipes var. acrior Shaffer 2. 427, 611 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 17 Aug. '80, 24 Oct. '81 8. Pinus contorta, Myrica californica, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum
1. Russula cf. emetics (Schaeff. ex. Fr.) S.F. Gray 2. 408 3. D. Largent 4. solitary 5. mycorrhizal 6. vegetated dune hollow 7. 23 Feb. '80 8. Pinus contorta, Salix hookeriana, Solidago spathulata, Polygonum paronychia, Carex obnupta, Juncus lesueurii
1. Russula fragilis (Pers. ex. Fr.) Fr. 2. 491, 6§7 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 26 Nov. '80, 5 Nov. '81 8. Pinus contorta, Myrica californica, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Rubus ursinus
1. Russula 'griseoviolaceus' Pseudo-name 2. 296, 506, 712 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest 7. 28 Nov. '79, 5 Jan. '81, 30 Nov. '81 8. Pinus contorta, Picea sitchensis, Abies grandis, Arbutus menziesii, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum
1. Russula pectinata (Bull. ex. St. Amans) Fr. 2. 20, 21, 23 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 5 Oct. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Myrica californica, Vaccinium ovatum, Solidago spathulata 142
1. Russula pelargonla Niolle 2. 46, 47, 193 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 15 Oct. '79, 1 Nov. '79 8. Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Rubus ursinus
1. Russula raoultii Quel. 2. 494, 526, 637 3. D. Largent 4. gregarious 5. mycorrhizal 6. forest 7. 1 Dec. '80, 28 Feb. '81, 5 Nov. '81 8. Pinus contorta, Picea sitchensis, Abies grandis, Pseudotsuga menziesii, Arbutus menziesii, Vaccinium ovatum, Gaultheria shallon
1. Russula rosacea (Pers. ex. Secr.) Fr. 2. 49, 55, 71 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 15 Oct. '79, 19 Oct. '79 8. Pinus contorta, Picea sitchensis, Myrica californica, Rhamnus purshiana,-Vāccinium ovatum, Solidago spathulata, Pteridium aquilinum
1. Russula xerampelina (Schaeff. ex. Secr.) Fr. 2. 300 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 28 Nov. '79 8. Pinus contorta, Picea sitchensis, Myrica californica, Gaultheria shallon, Vaccinium ovatum
1. Sarcosphaera ammophila (Dur. & Mont.) Moesz 2. 331 3. D. Largent 4. gregarious 5. saprophytic, terrestrial 6. vegetated dune hollows, established dune ridges, littoral strip 7. 6 Dec. '79 8. Eriogonum latifolium, Fragaria chiloensis, Solidago spathulata, Juncus lesueurii, Elymus mollis, Poa douglasii 143
1. Scleroderma arenicola Zeller 2. 423, 426, 544 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. forest and vegetated dune hollows 7. 24 Jul. '80, 10 Aug. '80, 3 May '81 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Salix hookeriana, Solidago spathulata
1. Scleroderma lycoperdoides Schweinitz 2. 163 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. established dune ridge 7. 30 Oct. '79 8. Pinus contorta, Salix hookeriana, Achilles borealis, Anaphalis margaritacea, Lathyrus littoralis, Eriogonum latifolium
1. Scleroderma macrorhizon Chev. 2. 12, 1:, 515, 59 3. D. Largent 4. clustered 5. saprophytic, terrestrial 6. established dune ridge interface with moving dune 7. 4 Sept. '79, 25 Jan. '81, 29 Sept. '81 8. Pinus contorta, Garrya elliptica, Lupinus arboreus, Solidago spathulata, Achilles borealis, Anaphalis margaritacea, Fragaria chiloensis
1. Stereum sp. Pers. ex. S.F. Gray 2. 405 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 8 Feb. '80
1. Stropharia ambigua (Pk.) Zeller 2. 337 3. D. Largent 4. scattered 5. saprophytic, terrestrial 6. forest 7. 7 Dec. '79 8. Pinus contorta, Baccharis pilularis, Garr a elliptica, Vaccinium ovatum, Solidago spathulata 144
1. Suillus brevipes (Pk.) Kuntze 2. 60, 81, 99, 161, 443, 447 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 16 Oct. '79, 21 Oct. '79, 25 Oct. '79, 30 Oct. '79, 21 Oct. '80, 23 Oct. '80 8. Pinus contorta, Picea sitchensis, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Solidago spathulata, Anaphalis margaritacea, Pteridium aquilinum, Juncus lesueurii
1. Suillus granulatus (Fries) Kuntze 2. 38, 59, 72, 343 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 13 Oct. '79, 16 Oct. '79, 19 Oct. '79, 7 Dec. '79 8. Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Salix hookeriana, Solidago spathulata, Fragaria chiloensis, Juncus lesueurii, Carex obnupta
1. Suillus lakei (Murr.) A.Η. Smith & Theirs 2. 649 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 5 Nov. '81 8. Pinus contorta, Pseudotsuga menziesii, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Pteridium aquilinum
1. Suillus ponderosus A.Η. Smith & Theirs 2. 381, 617 3. D. Largent 4. scattered 5. mycorrhizal 6. vegetated dune hollows 7. 30 Dee. '79, 25 Oct. '81 8. Pinus contorta, Arctostaphylos uva-ursi, Salix hookeriana, Carex obnupta, Juncus lesueurii 145
1. Suillus pseudobrevipes A.Η. Smith & Theirs 2. 35, 41, 57, 79 3. D. Largent 4. scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 13 Oct. '79, 16 Oct. '79, 21 Oct. '79 8. Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Gaultheria shallon, Salix hookeriana, Vaccinium ovatum, Solidago spathulata, Fragaria chiloensis, Carex obnupta, Juncus lesueurii
1. Suillus tomentosus (Kauff.) Sing., Snell & Dick 2. 22, 24, 51, 52, 67, 167, 442 3. D. Largent 4. solitary to scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 5 Oct. '79, 15 Oct. '79, 19 Oct. '79, 31 Oct. '79, 21 Oct. '80 8. Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Garrya elliptica, Solidago spathulata, Pteridium aquilinum
1. Suillus umbonatus Dick & Snell 2. 61, 64, 683 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 16 Oct. '79, 5 Nov. '81 8. Pinus contorta, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Salix hookeriana, Solidago spathulata, Carex obnupta, Fragaria chiloensis, Pteridium aquilinum
1. Thele hors terrestris Ehrhart ex. Fr. 2. 87, 9 3. D. Largent 4. scattered 5. mycorrhizal 6. vegetated dune hollows 7. 23 Oct. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Baccharis pilularis, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Rubus ursinus, Anaphalis margaritacea, Fragaria chiloensis, Juncus lesueurii 146
1. Tricholoma albobrunneum (Fr.) Kummer 2. 235, 329, 449, 661 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. forest and vegetated dune hollows 7. 10 Nov. '79, 4 Dec. '79, 25 Oct. '80, 5 Nov. '81 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Garrya elliptica, Gaultheria shallon, Solidago spathulata, Juncus lesueurii, Carex obnupta, Pteridium aquilinum
1. Tricholoma atroviolaceum Smith 2. 293, 302 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 28 Nov. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Pteridium aquilinum
1. Tricholoma aurantium (Schaeff. ex. Fr.) Ricken 2. 307, 339, 361 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 29 Nov. '79, 7 Dec. '79, 8 Dec. '79 8. Pinus contorta, Arctostaphylos uva-ursi, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum
1. Tricholoma cingulatum (Almfelt spud Fr.) Jacobasch 2. 157, 215 3. D. Largent 4. scattered to gregarious 5. mycorrhizal 6. vegetated dune hollows 7. 30 Oct. '79, 7 Nov. '79 8. Pinus contorta, Arctostaphylos uva-ursi_, Salix hookeriana, Lupinus arboreus, Solidago spathulata, Carex obnupta, Juncus lesueurii
1. Tricholoma flavobrunneum (Fr.) Kummer 2. 275, 291, 308, 489 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 26 Nov. '79, 28 Nov. '79, 29 Nov. '79, 26 Nov. '80 8. Pinus contorta, Arctostaphylos uva-ursi, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum 147
1. Tricholoma flavovirens (Pers. ex. Fr.) Lundell 2. 53, 114, 656 3. D. Largent 4. scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 15 Oct. '79, 26 Oct. '79, 5 Nov. '81 8. Pinus contorta, Abies randis, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Solidago spathulata, Juncus lesueurii
1. Tricholoma saponaceum (Fr.) Kummer 2. 219, 646 3. D. Largent 4. scattered 5. mycorrhizal 6. forest 7. 8 Nov. '79, 5 Nov. '81 8. Pinus contorta, Pseudotsuga menziesii, Arctostaphylos uva-ursi, Garrya elliptica, Gaultheria shallon, Vaccinium ovatum, Solidago spathulata, Pteridium aquilinum
1. Tricholoma vaccinium (Pers. ex. Fr.) Kummer 2. 117, 125, 182, 223, 283, 324, 355, 365 3. D. Largent 4. scattered 5. mycorrhizal 6. forest and vegetated dune hollows 7. 26 Oct. '79, 1 Nov. '79, 8 Nov. '79, 26 Nov. '79, 4 Dec. '79, 8 Dec. '79, 10 Dec. '79 8. Pinus contorta, Picea sitchensis, Arctostaphylos uva-ursi, Vaccinium ovatum, Salix hookeriana, Garrya elliptica, Gaultheria shallon, Solidago spathulata, Rubus ursinus, Carex obnupta, Juncus lesueurii
1. Tricholomopsis rutilans (Schaeff. ex. Fr.) Sing. 2. 281, 311 3. D. Largent 4. solitary to scattered 5. saprophytic, lignicolous 6. forest 7. 26 Nov. '79, 29 Nov. '79 148
1. Tilopilus porphyrosporus (Fr. & Hok) A.H. Smith & Theirs 2. 268 3. D. Largent 4. solitary 5. mycorrhizal 6. forest 7. 21 Nov. '79 8: Pinus contorta, Myrica californica, Arctostaphylos uva-ursi, Vaccinium ovatum, Gaultheria shallon, Pteridium aquilinum
1. Typhula juncae Fr. 2. 354, 450 3. D. Largent 4. gregarious 5. saprophytic on Garrya leaves 6. forest 7. 8 Dec. '79, 27 Oct. '80 1. Volvariella speciosa (Fr.) Sing. 2. 716, 717 3. D. Largent 4. solitary 5. saprophytic, terrestrial 6. margin willow swamp and moving dune 7. 30 Dec. '81, 5 Jan. '82 8. Alnus rubra, Salix hookeriana, Ribes sanguineum, Lupinus arboreus, Fragaria chiloensis
1. Xylaria hypoxylon (L. ex. Fr.) Grey. 2. 218 3. D. Largent 4. gregarious 5. saprophytic, lignicolous 6. forest 7. 7 Nov. '79 149
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