,\ECOLOGICAL AND TAXONOMIC STUDIES OF '1'IE RUSSULACEAE ‘ AND OTIER ECTOMYCORRHIZAL BASIDIOMYCETE} IN TIE

HIGIPELEVATION FORESTS OF TIE SOUTIERN APPALACHIANS bv ‘ Gerald F. Bills

dissertation submitted to the Faculty of the — ‘ Virginia Polytechnic Institute and State University

in partial fulfillment of the requirements for the degree of

DOCTOR OF PHILOSOPHY

in .

- Botany

‘ APPROVED:

//5*” /4//./ß’”)., '/’ ’ ’— 4 ¤»6 · * \·—L«

Orson K. Miller, Jr. , Chairman Khidir W. Hilu

6 . A rx · Ä' M/u-} Goldee.I. Häzman 2 W. CarterV Jvnson

I

I K 1RobertA. Peterson Jzy Stiges

May 24, 1985

Blackburg, Virginia ECOLOGICAL AND TAXONOMIC STUDIES OF THE RUSSULACEAE AND OTHR ECTOMYCORRHIZAL BASIDIOMYCETES IN THE

HIGH-ELEVATION FORESTS OF TH SOUTHERN APPALACHIANS

n by éä Gerald F. Bills

Orson K. Miller, Jr., Chairman

Botany A (ABSTRACT)

Temperate and boreal fungal floras indicate that species of the

Russulaceae (the genera Rgggglg and Lggtßgjgg) are among the dominant

ectomycorrhizal fungi in forest communities. The contribution of the

Russulaceae to the communities of ectomycorrhizal Basidiomycetes fruiting

in red spruce and adjacent northern hardwood forests in.West Virginia was

evaluated and compared with other ectomycorrhizal Basidiomycetes

occupying the same habitats. The Russulaceae exhibited the greatest

species diversity of any family of ectomycorrhizal fungi fruiting in the

stands studied (44 X of the species in spruce, 39 X of the species in

hardwoods). Species of Lagtagigg and Ryggylg were among the most

productive in both forests.

Species diversity, productivity, and fruiting phenology of all

ectomycorrhizal Basidiomycetes were compared between red spruce and

northern hardwood stands for a 3-year period. Sporocarp numbers and

sporocarp frequency in 384 four mz quadrats in each forest type was used

to estimate productivity. Species richness was greater in hardwoods (36 species) than in spruce (27 species). Nine species were common to both

forests. Most productivity was coneentrated in a few species, while most

species were rare. Speeies·area curves were constructed for both forests.

Fungal species and tree species eomposition in both forests were compared

by principal component analysis.

Fungi in spruce forests were more produetive than in hardwood forests.

Productivity was highly variable among the three seasons studied because V of climatie variability. Sporocarp abundance and frequency were

positively correlated with basal area and density of mycorrhizal trees

and were negatively correlated with fern cover in hardwood forersts. .

Fruiting seasons extended from early July to late September or early .

October.

_ Numbers of species fruiting from the same four mz quadrats ranged from

0 to 7 in spruce forests and O to 5 in hardwood forests. Spatial patterns

of sporocarps of major species were eharaeterized by the variance·to-mean

ratio, mean erowding, patehiness, and spatial autocorrelation and were

found to exhiblt highly aggregated, contagious patterns. Interspecific

associations between pairs of major species were measured by 2 ¤ 2

eontingency tables and Cole's index of association.

A taxonomie and geographie survey of Rggsylg and Lggtggjygloeeurring

in both the quantitative study areas and in similar habitats in the

Southern Appalaehians was presented. I would like to thank Dr. Orson K. Miller, Jr. for his teaching,

guidance, logistic support, and patience during the course of my graduates

studies. Drs. Golde I. Holtzman and W. Carter Johnson provided

indespensible assistance and guidance in the design, execution, and

analysis of the quantitative aspects of this study. The teaching, ' professional guidance, and personal advice of the other members of my

advisory comittee, Drs. Khidir W. Hilu, Robert A. Peterson, and R. Jay _ ”_ Stipes is greetly appreciated. The friendship, assistance, suggestions,

and logistic support of fellow graduate students, technical staff, and

faculty, past and present, at VPI & SU is also appreciated.

Financial support was provided by a graduate teaching assistantship

from the Department of Biology, VPI & SU. The Jeffress Memorial Trust

provided me with summer support and with a unique opportunity to learn

the higher fungi of Virginia. A Gertrude S. Burlingham Fellowship from

the New York Botanical Garden supported me during the spring of 1984 and

allowed me to study first-hand many of the Russulas and Lactarii described

by G. S. Burlingham, C. H. Peck, H. C. Beardslee, C. H. Keuffman, W. A.

Murrill, and R. Singer.

Finally, my education could not have been completed without moral and

financial support from my family and my wife, Nancy. Many thanks to all.

Acknowledgements iv TABLE QB SQNIENIS

INTRODUCTION ...... 1

CHAPTER 1. COMPARISON OF FRUITING OF ECTOMYCORRHIZAL BASIDIOMYCETES

BETWEEN RED SPRUCE AND NORTHERN HARDWOOD FORESTS ...... 4

Introduction ...... 4

The study areas ...... 8

Methods and materials ...... 11

Results ...... 14

Species diversity ...... 14

Inter·plot relationships ...... 25

Sporocarp frequency and density ...... 30

Fruiting Phenology ...... 33

Length of sampling period ...... 36

Discussion ...... 37

CHAPTER 2. SPATIAL PATTERNS AND INTERSPECIFIC ASSOCIATIONS OF

ECTOMYCORRHIZAL BASIDIOMYCETES IN RED SPRUCE AND HARDWOOD FORESTS 47

Introduction ...... 47

Methods ...... 49

Results ...... 52

Sporocarp patterns ...... 52

Interspecific associations ...... 60

Discussion ...... 70

Table of Contents V CHAPTER 3. SYNOPSIS OF RUSSULA IN THE HIGH-ELEVATION FORESTS OF THE

SOUTHRN APPALACHIANS ...... 76

Introduction ...... 76

Synopsis of Russula in the high·elevation forests ...... 79

CHAPTER 4. NOTES ON LACTARIUS IN THE HIGH-ELEVATION FORESTS OF THE

SOUTHRN APPALACHIANS ...... 97

Introduction ...... 97

Descriptions of taxa ...... 98

CHAPTER 5. DISTRIBUTION OF LACTARIUS IN THE HIGH-ELEVATION FORESTS

OF THE SOUTHRN APPALACHIANS ...... 118

Introduction ...... 118

Habitats and distribution ...... 121

Discussion ...... 124

APPENDIX A. LOCATION OF SPRUCE AND HARDWOOD PLOTS, POCAHONTAS CO.,

WEST VIRGINIA...... 128

APPENDIX B. FREQUENCY MAPS OF MAJOR SPRUCE AND HARDWOOD

BASIDIOMYCETES...... 133

APPENDIX C. TREE (> 2 CM DBH) LOCATIONS IN HARDWOOD AND SPRUCE PLOTS. 149

APPENDIX D. RAW DATA 1981-83...... 162

Table of Contents Vi BIBLIOGRAPHY ...... 163

BIBLIOGRAPHY ...... 164

VITA 173

Table of Contents v;|_;|_ LI§IQEFigure1. Red spruce and northern hardwood transition zone on the south end of Gauley Mt., Webster Co., West Virginia. . 6

Figure 2. Dominance·diversity curve for spruce ( ¤ ) and hardwood ( u ) plots...... 17

Figure 3. Species-area curve for spruce ( A ) and hardwood ( 0 ) plots...... 23 • Figure 4. Ordination of spruce ( 0 ) and hardwood ( ) plots by fungal frequency...... 29 • Figure S. Ordination of spruce ( 0 ) and hardwood ( ) plots by tree basal areas...... 31

Figure 6. Sporocarp phenology of all species in hardwood and spruce plots...... 39

Figure 7. Sporocarp phenology of some major species in spruce and hardwood plots...... 40

Figure 8. Log10 V/m ratio of major species in spruce forests. . . S4

Figure 9. Mean crowding of major species in spruce forests. . . . 55

Figure 10. Patchiness of major species in spruce forests. .... 56

Figure 11. Log10 V/m ratio of major species in hardwood forests. . 57

Figure 12. Mean crowding of major species in hardwood forests. .. S8

Figure 13. Patchiness of major species in hardwood forests. ... 59

Figure 14. Lactarius sporocarps...... 99

Figure 15. L. lignyotellus microscopic features (GB 161). . .. 101

Figure 16. L. oculatus microscopic features(GB S10)...... 107

Figure 17. L. fragilis microscopic features (GB 438)...... 113

Figure 18. Locations of plots S1 and S2, Woodrow quadrangle, West Virginia...... 129 l Figure 19. Locations of plots H3 and H4, Hillsboro quadrangle, West Virginia...... 130

List of Illustrations viii Figure 20. Locations of plots S7, S8, H5, and H6, Lobelia quadrangle, West Virginia...... 131 · 1 Figure 21. Locations ef plots S9, S10, H11, and H12, Lobelia quadrangle, West Virginia...... 132

Figure 22. Frequency of Lactarius oculatus in spruce plots. . . 134

Figure 23. Frequency of Clavulina cristata in spruce plots. . . 135

Figure 24. Frequency of Lactarius vinaceorufescens in spruce plots. 136

Figure 25. Frequency of Boletus badius in spruce plots. .... 137 · Figure 26. Frequency of Amanita flavaconia in spruce plots. .. 138

Figure 27. Frequency of Lactarius lignyotellus in spruce plots. 139

Figure 28. Frequency of Inocybe umbrina in spruce plots. .... 140

Figure 29. Frequency of Russula granulata in spruce plots. ... 141

Figure 30. Frequency of Amanita inaurata in spruce plots. . .. 142

Figure 31. Frequency of Lactarus camphoratus in spruce plots. . 143

Figure 32. Frequency of Lactarus camphoratus in hardwood plets. 144

Figure 33. Frequency of Russula granulata in hardwood plots. 4. . 145

Figure 34. Frequency of Boletinellus merulioides in hardwood plots. 146

Figure 35. Frequency of Scleroderma citrinum in hardwood plots. 147 .

Figure 36. Frequency of Laccaria laccata in hardwood plots. . . 148

Figure 37. Tree locations in plot S1...... 150

Figure 38. Tree locations in plot S2...... 151

Figure 39. Tree locations in plot H3...... 152

Figure 40. Tree locations in plot H4...... 153 _

Figure 41. Tree locations in plot H5...... 154

Figure 42. Tree locations in plot H6...... 155 .

Figure 43. Tree locations in plot S7...... 156

Figure 44. Tree locations in plot S8...... 157

List of Illustrations ix Figure 45. Tree locations in plot S9...... 158

Figure 46. Tree locations in plot S10...... 159

Figure 47. Tree locations in plot H11...... 160

Figure 48. Tree locations in plot H12...... —...... 161

List of Illustrations x The main objective of this study was to evaluate the participation of

Russulaceae in the communities of ectomycorrhizal fungi in the high-elevation spruce and northern hardwood forests of the Southern

Appalachians. The Russulaceae is one of the largest and most common families of mushrooms with hundreds of species occurring in boreal and temperate forests. Species of the Russulaceae and other ectomycorrhizal fungi depend on the composition of the vegetation because they are nutritionally dependent upon trees of the Pinaceae, Fagaceae, Betulaceae,

Salicaceae, and possibly the Juglandaceae (Trappe, 1962; Miller, 1982).

The Russulaceae are presumed to be among the most important mycorrhizal symbionts of forest trees because of the abundance and ubiquity of their sporocarps in forests.

Most knowledge about the composition, productivity, and fruiting phenology of Basidiomycetes is based on studies of European forests, grasslands, and tundra. Most of the European studies have been floristic, however, simply listing species in different vegetation types. Another group of predominately continental European investigators has applied association analysis and the Braun-Blanquet method of community sampling, data reduction, and association nomenclature to macrofungal communities.

American, British, and some Scandanavian researchers have relied on random sampling methods to obtain quantitative estimates of species diversity, sporocarp densities, biomass, and sporocarp longevity for

Introduction 1 deseribing and comparing macrofungal eommunities. The last group of investigations has served as the basic model for my community studies.

In my studies, a descriptive approach has been taken to determine the components of ecosystems rather than a modelling approach to elucidate ecosystem processes. An inventory and understanding of the basic components of an ecosystem are required before the components can be integrated into a working model.

The higher fungi of the Southern Appalachians have never been the focus of a community·oriented study. Nearly all the literature on the

Basidiomycetes of this geographie province has been taxonomic. Many studies have scrutinized taxonomic groups across a broad spectrum of plant eommunities but rarely presented anything more than cursory data on fungal-vegetation relationships.

This dissertation is divided into five chapters. The first chapter describes and compares the ectomycorrhizal Basidiomycete eommunities of red spruce (including the Russulaceae) and northern hardwood forests in

West Virginia. The fungal species composition, richness, and diversity are related to the diversity of the dominant trees of both forests types. Species•area relationships between the forests are compared.

Relative dominance of the fungi is compared by the sporocarp frequencies and densities. Finally, fruiting phenology is compared between the two forest types and among the major species.

Introduction 2 In the second chapter, pattern analysis based on quadrat sampling and the recently developed techniques of spatial autocorrelation are employed to describe and compare the spatial patterns of sporocarps of the major

species. Also, two questions commonly asked in plant ecological studies

are addressed: "are species distributed randomly" and "are species

associated?"

One of the main reasons fungal ecology remains an underdeveloped branch

of ecology is the taxonomic complexity of fungi. Systematic

specialization and extensive research are often needed to accurately U determine and document the taxa involved. A large effort is directed to

a rlgorous taxonomic survey of the genera Rggsglg (Chapter 3; Bills and

Miller, 1984; Bills, 1984) and Lggtggjgg (Chapter 4) in the quantitative

study areas and in comparable habitats throughout the region.

Lgggggjgg is a relatively well-known genus taxonomically. In Chapter

S, a comparison of the Lgggggigg flora of the high-elevations of the

Southern Appalachians with those known to occur in boreal forests of

northeastern North America is presented to demonstrate how geographic

distributions and community structure of mycorrhizal fungi in the

Southern Appalachian spruce-fir forests might differ from those in a true

boreal conifer forest.

Introduction3 QHAHIERL. QEEBLlIIIN§Q£BEIHEENREDSERLLQEANDNQRHIERNHARRWQQDEQREEIS

Community studies of higher fungi have relied largely on quantitative descriptions of fruiting. The limitations of studying fungal communities based upon observation of sporocarps have been discussed by Hueck (1953),

Arnolds (1981), and Fogel (1981). Sporocarp productivity has been estimated in one type of higher plant community (Richardson, 1970; Fogel,

1976) or has been compared among different plant communities (Lange, 1948;

Hering, 1966; Petersen, 1977; Wasterlund and Inglelog, 1981; Arnolds,

1981). Sporocarp presence, frequency, or productivity have been used alone or in conjunction with the higher plants to compare vegetation samples (Lange, 1948; Hering, 1966; Petersen, 1977; Arnolds, 1981). The

influence of precipitation and temperature on the phenology and productivity of fruiting has been investigated to understand the environmental conditions influencing the physiology of the fruiting

(wilkins and Patrick, 1940; Wilkins and Harris, 1946; Fogel, 1976).

Monitoring sporocarp productivity of different fungal species in a forest dominated by a single ectomycorrhizal host has indicated possible

ectomycorrhizal associates of the dominant tree (Trappe, 1962;

Richardson, 1970; Fogel, 1976). Observation of the spatial pattern and

relative numbers of sporocarps in permanent reference areas over extended

time periods has elucidated the spatial pattern and relative abundance Chapter 1 4 of vegetative mycelia in the forest rhizosphere (Last gg gl., 1983;

Newell, 1984; Cotter and Bills, in press).

_ Quantitative studies of macrofungal communities have usually focused

on sporocarp numbers or sporocarp biomass in standard reference areas.

The critical assumption of many of these studies was that the relative

productivity of sporocarps among fungal species in some way reflected

their relative dominance, mycelial biomass, or resource utilization.

Naturally occurring mycelia cannot be delimited in a direct manner, except

in rare cases. Presently there is no basis for correlating sporocarp

biomass or sporocarp numbers with mycelia biomass. However, sporocarp

density is a useful parameter because it can be applied objectively in

any study, and if sampling intervals and methods are comparable, sporocarp

densities can be compared among communities. This parameter is included

in this study to provide continuity with previous investigations. Rather

than emphasizing sporocarp numbers as a measure of mycelial activity, we

estimated the frequency with which sporocarps were present in relatively

small contiguous quadrats. lf the quadrats are small enough, frequency ' can show that sporocarps in widely separated quadrats at one sampling _

time may be part of a single zone of contiguous fruiting when sporocarps

are sampled in quadrats in intermediate positions. Therefore, frequency

provides an estimate of the spatial extent or ubiquity of the fungal

mycelium.

Species concepts of Basidiomycetes are based upon sporocarp

morphology, but rarely on habitat preferences. Statistical relationships

Chapter 1 5

between the distribution of fungal species and their associated higher

Vegetation may refine these concepts. In many "modern" taxonomic

treatments of Basidiomycetes, little attention is devoted to habitat

descriptions. Often fungi are described simply as fruiting "under A conifers" or "in mixed woods". A species that appears to be "rare" or

is "poorly known" may be common locally in specialized habitats. This

local abundance may not be recognized by conventional collecting methods

but can be detected by periodic, systematic sampling of local communities_

(Arnolds, 1982; Fogel, 1982).

The ectomycorrhizal fungal community of pure red spruce forests was

chosen for study because the fungi could be assumed to form

ectomycorrhizae with only a single host and that niches for

ectomycorrhizal fungi would be limited by the availability of a single A tree species. Variations in tree age, root age, root size, litter and

soil depth, etc. could contribute, however, to niche diversification

among fungal species that occupied a similar substrate. To identify

properties of the community and species unique to the red spruce stands

we compared them to surrounding northern hardwood stands (Braun, 1950;

Whittaker, 1956; Core, 1966) of mixed ectomycorrhizal and endomycorrhizal

trees.

No comprehensive source of information on Basidiomycetes associated

with the red spruce forests of eastern North America exists, and few

mycorrhizal associates of red spruce are known (Treppe, 1962; Homola and

Mistretta, 1977). The stands of red spruce in this study are part of the

Chapter 1 7 disjunct southern extension of the boreal coniferous forest. The

structure and floristic composition of the Southern Appalachian and

Allegheny red spruce-Fraser fir-balsam fir forests are similar to those

in New York and New England (Oosting and Billings, 1951; Mclntosh and

Hurley, 1964; Stephenson and Clovis, 1983). Are the Basidiomycetes

associated with red spruce similar throughout its geographical range?

Presently, the only way to answer this question is by examining the usually incomplete habitat descriptions of Basidiomycetes to determine whether they were collected in the vicinity of red spruce.

The main objectives of this study were (1) to determine how species composition, diversity, and density of ectomycorrhizal Basidiomycetes fruiting in forest stands dominated by a single ectomycorrhizal host tree, red spruce, differed from those of nearby forest stands of mixed ectomycorrhizal and endomycorrhizal hardwood trees, (2) to describe the inter-year variation in fruiting and sporocarp density in both forest types and to characterize the fruiting phenology of major Basidiomycete species of both forest types, and (3) to provide a basis for comparing

Basidiomycete communities in similar coniferous and hardwood forests in eastern North America.

IHE.S§I1ZD.YAREA§

The study areas were located near the eastern edge of the unglaciated

Allegheny Plateau in Pocahontas Co., WV, within the Monongahela National

Chapter 1 8 Forest (boundaries 38° l7° N latitude, 38° 07° N latitude, 80° 22° W

longitude, 80° 12° W longitude). Sites were located on ridge crests

(elev. 1200-1350 m) on shallow, rocky well-drained, sandy-loam or

clay-loam soils. Within the spruce stands, soil pH (n = 23) ranged from

3.3 to 3.8 with an average of 3.5, and soil organic matter content was

12.3 % i 3.6 % S. D. Within the hardwood stands (n = 20), soil pH ranged

from 3.3 to 4.5 with an average of 3.6, and soil organic matter content

was 10.6 % i 3.4 % S. D.

Annual precipitation in the vicinity of the study sites during a

five-year period in 1967-72 ranged from 144 cm/yr to 160 cm/yr (Edens,

1973). Precipitation patterns for each growing season were estimated from

data collected by the U. S. Forest Service, Marlinton, WV, (elev. 650 m)

about 15-20 km west of the study areas (Table 1). Extended periods of

high humidity and fog are partially responsible for the persistance of

red spruce at these southern latitudes (Core, 1966). Frost-free periods

range from 88 to 145 days (Edens, 1973). A

Both spruce and hardwood stands were second-growth (55-75 years old).

Floristically and structurally the spruce stands were similar to stands

described by Stephenson and Clovis (1983). Spruce stands were nearly pure

red spruce with a sparse to dense shrub layer of ygggjgjgg

ggythgggggpgg, suppressed spruce seedlings, and ferns. An extensive

ground cover of bryophytes, especially the leafy liverwort ßgzzggjg

ggilgbgtg (L.) S. F. Gray, was often present. Composition of the hardwood

Chapter 1 9 Table 1. Biweekly summary of average (cm/day) and total (cm) rainfall for growing seasons of 1981-3 based on U. S. Forest Service data, Marlinton, WV.

1981 1982 1983 weeks average total average total average total

Jun 1-15 0.53 7.85 0.79 12.01 0.25 3.71

Jun 16-30 0.20 3.00 0.10 1.55 0.23 3.58

Jul 1-15 0.97 14.27 0.54 8.05 0.27 4.04

Jul 16-31 0.48 7.54 0.28 4.45 0.25 3.81

Aug 1-15 0.28 4.29 0.25 3.76 0.12 1.80

Aug 16-31 0.03 0.51 0.25 3.76 0.01 0.10

Sep 1-15 0.71 10.52 0.08 1.09 0.02 0.25

Sep 16-30 0.09 1.40 0.16 2.42 0.20 3.07

total (cm/season) 49.38 40.82 20.37

Chapter 1 1Q stands are listed in Table 3. Herbaceous plants, including ferns, and

young Age; stems were abundant in the understories of the hardwood stands.

MEIHQQSANDHAIERIALS

This study was conducted during the growing seasons of l98l•3. Twelve

permanent 16 X 16 m (256 mz) plots were established on ridge crests of

three different mountains. On each mountain, two plots were located in

V a spruce forest and two in a hardwood forest (Table 2). Each plot was

subdivided into 64 2 X 2 m (4mz) quadrats.

Spruce plots were selected to exclude as many other woody species as

possible. Hardwood plots were selected to be as physically close to the

spruce plots on the same ridge crest, without including any red spruce.

In each plot, DBH (diameter at breast height) of all stems greater than

two cm was estimated and mapped, and fern, bryophyte, and spruce seedling

cover was estimated. Hardwood plot H3 was destroyed by a survey crew in

the spring of 1983. Sporocarp density and frequency estimates for 1983

were based only ou the five remaining hardwood plots.

Only sporocarps of fungi of families known to form ectomycorrhizae

(Watling, 1982; Miller, 1983) and some whose ecological role is uncertain

me;g1jgjgee, Eggglgge spp., and some Hyg;gphg;ee spp.) were counted.

Litter decomposers (e.g. Qgllyhie, Me;eemig;, or Mygege spp.) or

Chapter 1 11 Table 2. Location, forest type, and plot label.

forest type location spruce hardwood

Black Mountain Sl & S2 H3 & H4

Kennison Mountain S7 & S8 H5 8: H6

Rocky Knob S9 & S10 H11 & H12

Chapter 1 12 bryophyte-associated species (e.g. as Qglering spp.) were not counted.

Sporocarps of agarics and boletes were easily defined, but the coral

fungus, Qlgygliga ggistgtg, forms multiple, fused stems. A single

sporocarp of this fungus was defined as a separate stem completely

surrounded by the surface of the litter or the bryophyte layer, although

the stems may have been fused below ground. From examination of records

of sporocarp positions from previous sampling times, it was evident that

nearly all sporocarps deteriorated between sampling periods. Only

sporocarps that were long-lived and could have been counted twice (e.g.

ßglggggggmg gitringm, large Lggggrigg spp.), those needed for

identification, and those needed for Voucher specimens were removed from

the plots. Sporocarp numbers were prcbably underestimated because

sporocarps of fleshy fungi are usually short-lived (Richardson, 1970;

Lacy, 1984) and could have fruited and deteriorated between sampling

times. Representative Voucher specimens of all fungi are deposited at

VPI. Vascular plant nomenclature follows that of Strausbaugh and Core

(1978). Fungal nomenclature is listed in Table 4.

Individual sporocarps were counted within each 2 ¤ 2 quadrat at each

visit. Plots were visited eight to ten times per growing season at 7-

to 17-day intervals with the average time interval between visits 13.1 i

3.2 days S. D. (n = 23).

_ Density is the number of sporocarps per unit of area (either in all

plots of a forest type or in each plot). Frequency is the number of

quadrats in which a species occurred summed over the entire the study.

Chapter 1 13 Percent frequency is the percentage of the total number of quadrats (384) l in each forest type in which a species is present. The total number of quadrats in which a species occurs in any given year is yearly frequency.

Frequency of a species is not the sum of its yearly frequencies.

BE§LJLI§

S.2EQIE§ IZIYERSJIY

Forests with several possible mycorrhizal hosts might have a greater diversity of fungal associates than a forest composed of a single mycorrhizal host. Two elements of species diversity are usually considered, (1) species richness, the number of species, and (2) equitability, the evenness of the contributions of different species to the community. Fungal species richness was comparable between both forest types (Tables 4, 5), but the two forest types exhibited little overlap in fungal species composition (Tables 4, 7). A total of 54 species was encountered over the three years, 27 species in the spruce plots, 36 in the hardwood plots (Table 5). Nine (17 Z) species occurred in both forest types. The family Russulaceae accounted for more species than any other family in both forest types, 39 Z of the species in the hardwoods and 44 ‘

Z of the species in spruce. The total number of species occurring only in one forest type or the other (19 only in spruce, 26 only in hardwoods) _ were not significantly different from what would be expected assuming the two forest types were equally likely habitats (22.5 in each forest type),

Chapter 1 14 tr•• sp•ci•s Tab1• 3. Basal ar••s lmz7ha) of in spruco and ord•r•d hardnood plots. Plots ar• from laft to ridut by ascanding _ ‘ $p•ci•s ar• first principal couponant scoras. ordorad by dascanding first principal couponant scoros from a sp•ci•s ordination. Plot labals ar• indicatad in Tabla 2.

” plot

spacias $7 $2 S8 S10 $9 $1 H6 H5 H11 H12 H4 H3

Picea nbens 53 43 57 46 43 42

Acor saccharuu 4 >1 16 13 35 20 >1 >1 Fagus grandifolia 3 _>1 6 7 15 37 Prusus sarotina 16 24 5 5 Quarcus nbra 37 Fraxinus amoricana 11 2 Batula alladwaniussis >1 1 2 >1 · Acar nbnn 3 >1

Acar pansylvanicuu >1 >1 Pruxus pensylvanicmn >1 Sorbus amaricana >1 ’ I1•x montana >1

Total 58 45 57 46 43 42 47 40 48 36 54 38

Chapter 1 15 (Table 5). Likewise, the mean number of species per plot was not

. significantly different between spruce and hardwood plots (Table 5). The

similarity in slopes of the species-area curves, (Fig. 2) also indicated

comparability in species richness, although the initial slope of the

hardwood curve was slightly steeper because of the encounter of more

species in hardwood plots than in spruce plots.

Species equitability was greater in the hardwood plots than the spruce V plots. The Shannon-Wiener index (Shannon and Weaver, 1949), H'= -2 Pi

log2 Pi, was used to compare both species richness and equitability

between both forest types (Table 5). Where Pi is the probability of

sampling the ith species among all species. Shannon-Wiener diversity (H')

was greater for hardwood plots than spruce plots using both sporocarp

frequency and density (Table 5) because of the greater number of species

and greater equitability among species. Equitability or eveness of a

community (J') (Pielou, 1974, p. 300) can be measured directly from the

ratio of the observed species diversity (H') to the maximum value of H'

(H°max) in a completely equal community with the same number of species

(S). Therefore, equitability (J') is given by H'/ log 2 S. As expected,

higher (H') values were obtained using sporocarp density because density

measurements were more equally distributed among species. The

dominance-diversity curve (Greig-Smith, 1983), (Fig. 1) indicated that a

high proportion of the frequency in both forest types was concentrated

in relatively few species, that a high proportion of the species appear

rarely, and equitability was greater among the hardwood species. Greater

Chapter 1 16 III 1• El ' " • oUI S :1 § = I5 No QIIOE- = A In 5 Q u • Q E Ü l „„ a •• C10 I 3 S cx lu; 2 nn g ¤¢"‘ "' ¤ nun CI II » ¤¤ = ¤¤¤¤¤¤¤¤Ü¤¤KIIIIII1:1:1 IIIIIIII 0 • 0 no II ao zu ao es SPICIIS Stoutuct

Figure 2. Dominance-diversity curve for spruce ( ¤ ) and hardwood ( • ) plots. : Most frequent species to least frequent species are ordered from left to right.

Chapter 1 17 T•b1• 4. ßasidianyeatss balonging ta •ata«•yearrhiz•1 f•••il.i•s aeaurring in spruea ls) and harchaad Ih! faust typ•s.

F••i1y• typ• sp•ci•s faust

IMANITACEAE • Amnib ilmuszmin Atk. •, muib hlx: |$ah••ff.l •¤•«·•. In S•¢•-. Ällüil s, h ßemuib sasiliu ts. a •«·.1¤••1 mmib s.t¤n¤u1.•b Fr.• smsu sm. h gi}; gig}; lBu11.:Fr.) Vitt. h .

BOLETACEAE uisjimiisa mmlinidu «s¤+ui•n.1 Hurrill. h Bzlßm !.f.f.i¤sa1 PK- 1 h •„ l klein Indian rr. h Mina szbmumbuzn re-. h Ehxlimamm daszäaznilyg ssen.) am. h • Ixisniisn 1111191 u=•·.1x•«~•1.

cm‘n•ARELLAcEAE • Qmilmmlhm ishniemb re-.

CLAVARIACEAE • Sbxuiim snisbb m~.r s¢•«~¤•e.

Chapter 1 18 Teble 4. Con'!.

Feeily. •p•¤i•e plct

CORTINARIACEAE Gsdimnige alnsgn Fr- • •¤• §.¤Lti¤e:iyg 1 • •¤- §2Lti¤:¤1;i.u.; 3 • ¤•·••• Immda sadzcim • Ingame so- h ENTOLGIATACEAE Eniszlm limisaujig N••l· •¤~ h Enielsm 2 •

ggü IB. I C.) Seec. I1 Enjglg IPR.) Seee. I1 Enisism •¤- h

HYGROPHORACEAE

|$cI·••ien.I Fr. I1 Iknuff.) Smith I Heel. I1 (Fr.) Fr. I1 (Fr.) Fr. I1 Uamnanbzug sp- b TRICHOLGIATACEAE §.¥!$&r:gmi1¤§1im¤¤Fr•)F•y¤d •.h I lgggjg lggg I$o¤p.ZFr.) Berk. I Bree. e, I1

SCLERODERHATICEAE Sslncszsang P••·•• süzimn h

19. Chapter 1 T•b1• 4. Cant.

l •p•ci•• Fnnily, plot

PAXILLACEAE • ggging; imgigg Ißatseh.) Fr.

RUSSULACEAE Fr. uh • Lasimim magnlixm Pk- . Lasizcius simusn Pk- H •• L1s$.u;iy1 m.¤.¤üi Pk- H Lgsigdm ndmm Pk- H l M••1- • Lasiuzisn limxsztnllus Smith I • Lgjggigg gglgjgg IPR.) Burl. • Imizciua mcdistn Pk- ],;g_g;_;ig; ghgjgggig; IBu11.) Fr. h • Lasxgcisa Smith L•C1•i•· • ßgggß; gg; S•·¤v• • t · Bsasula slemtlma ßmub sanmm Pk- H $•¤r-• Sh•ff••· • B!-!.t§.ul.! mi.f.21il 3;:;;; •• ßggggi; IPR.) PR. h ßggggi; Fr. ? h SH•f*••· Basale Iszzmäxzkii H ¤•-•r1- auml: mm H ¤¤•·1- mmala mlm h Biggi]; ;j_],yj,gg_],; $h•ff•r ag h e Byssyla sgtatm Smith 1 H &;;_;g_],; gigggg I$eh••ff.) Fr. In

Chapter 1 20 species diversity, species Table 5. Comparison of species richness,

equitability, and species density of ectomycorrhizal

fungi between hardwood amd spruce plots z (H') and (n=6 plots/forest type). Shannon-Wiener index density and frequency. equitability (J') calculated from sporocarp

spruce hardwood

36 number of species 27 27a number of species 188

unique to each A

mean i S. D. of mz) 13.2b i 4.3 species/plot (256 15.7b i 2.2

' maximum number of A A mz) 19 species/plot (256 19

minimum number of I mz) 7 species/plot (256 13

4.11 H'(frequency) 2.39

4.52 H'(density) 3.59

0.76 J'(frequency) 0.50

0.87 · J'(density) 0.79

xz 1.8, 1 d.f. a Not significantly dlfferent, P>0.10, = T 1.0, 10 d.f. b Not significantly different, P>O.10, =

Chapter 1 21 equitabilty in the hardwood forests was indicated by direct measurement

of J!. .

The total number of species fruiting in a quadrat estimates the minimal ' number of ectomycorrhizal species that may be occupying the rhizosphere

within these small areas (Table 6). Species density was much higher in

the spruce plots than in the hardwoods. The proportion of the quadrats

with no sporocarps was much higher than those with sporocarps, 225 (59 '%) in the hardwoods compared to 59 (15 %) in the spruce. Two species

fruited in most (85 %) of the occupied spruce quadrats, and as many as

seven species fruited in a single quadrat. One species fruited in most

(25 X) of the occupied hardwood quadrats, and as many as five species

fruited in a single quadrat.

The concept of minimal area (Cain, 1938), or the smallest area in which

the species composition of a community is adequately represented, is

impossible to apply prior to sampling macrofungi because sporocarp

densities are in a state of continual flux. However, the possibility of

achieving a minimal sampling area was examined in retrospect by

constructing the species·area curve (Greig·Smith, 1983) (Fig. 2) from

accumulated data on sporocarp locations. One criterion for minimal area

(Mueller-Dombois and Ellenberg, 1974) is a sample size that contains 90

to 95 % of the maximum number of species encountered in the largest sample

unit. This criterion is impractical for fungi because the maximum number

· of species cannot be determined. Cain (1938) considered the minimal area

Chapter 1 22 ' 35 30 m A Lu {3 25 ,9 E th 20 ‘° U- • AO O_ 15 • O • ‘ z 10 O‘A 5

0 zoo 600 600 000 1000 1200 v•0¤ 16¤¤ AREA MZ

• Figure 3. Species-area curve for spruce ( A ) and hardwood ( ) plots.: The total number of species in the sample areas was plotted against increasing areas of contiguous quadrats. When 256 mz, the size of one contiguous plot, was reached additional plots were chosen randomly and their species added.

Chapter l 23 sp•ci•• Tabla 6. Distribution af nuebars af in 384 quadrafs af spruea and hardnaod plats.

na. af na. af quadrafa spaciaa/qeadraf spruaa harcbaaad

0 54 226 1 83 97 2 105 44 3 84 14 4 42 3 5 ll 1 6 4 0 7 l 0

fatal aeenpiad qnndrafs 330 158 · uaan 2 S. 0. 2.4 rt 1.2 1.5 t 0.8 nurbar af sp•ei••/cpadraf

Chapter 1 24 to be the point on the species·area curve at which an increase of 10 %

in the sample area yields only 10 % more species of the total number

recorded. Using this criterion, the minimal area for the spruce forests

is approximately 250 mz and approximately 150 mz for the hardwood forests.

However, Rice and Kelting (1955) demonstrated that this 10 X point wlll

shift continually to the right as greater total areas are sampled.

To compare and to summarize variation in fungal and tree species

composition among the plots, principal component analysis (PCA) (Gauch,

1982, chap. 4) was used to order the plots x species data matrices for

fungi and trees separately. The hardwood plots appeared to be distinctly

different from the spruce plots because of the taxonomic discontinuity

in fungal species between the two forest types (Fig. 4). Ordination

indicated a closer relationship among the fungal samples in spruce forests than among hardwood forests. The spruce plots were relatively uniform

in fungal composition having nine fungal species in common. The wide separation among hardwood samples along all three axes indicated a high degree of taxonomic discontinuity. Hardwood plots H6 and H5 were the most different from the rest, probably due to the high frequency of

ßglgtjggllgg mgggljgjggg in H6 and Lgg;g;1gs gigggggg in H5. In addition,

Hygggphgggg spp. and Engglgmg spp. were largely restricted to these two plots (Table 7).

Chapter 1 25 T•b1• 7. Pareant frnqaucy of basidionyuata sp•¤i•s in spruea and Inrénood plots in southnastnrn Hast Virginia. Plots ars ordnrad fron 1•ft to n·i¢1t by asoanding first principal cannvonnnts scoros. Spncix · arn ordarnd by dasacnding first principal couponmt scoras fron a spncias ordirctiun. Plot labais are indieatad in Tsbla 2.

p1¤t S¤•¤ix $9 S2 S8 S7 $1 $10 H11 H12 H3 N4 M6 M5

Lactarius oculatus 23 66 61 56 47 27

Clavulina cristata 20 36 75 31 53 16

Bolatus badiua 25 14 50 17 3 17

Lactarius vinncaonafascuns 33 4 23 3 1 19 Annnita flarvaconia 33 37 4 4 13 08

Laetarius 1ig·ny¤t•11us 23 33 5 3 19 6

Inocyba ubrina 22 11 5 8 8 6 Lactarius sordidn 8 1 11 Annusita incurata 5 1 1 22 3 1 3 3 3 Rusaula granulata 3 5 14 9 14 1 6 9 9 3 8 i Mlnita fulva 5 9 3 3 3 Russula eilvioola 8 3 1 3 3

Iussula aqnsa 1 5 3 Cortinarius sp. 1 3 1 5 1 Lactarius dacnptivus 9 Ccrtinariu painacus 3 1 5 1 5 1 Lactarius canuhoratus 6 6 17 3 1 11 0 1 5 8 Cystodarnnna anianthinuu 6 u 1 1 Cantharallus txbaafornnis 6 1 ' Entolonna sp. 1 5 _ laceata Laccaria 6 6 3 1 8 1 6* 8 1 Tylopilua fallnus _ 5 '

Russula dansifolia 1

Chapter 1 26 Tab].! 7. cent.

plot speeiu S9 $2 $8 S7 $1 $10 H11 M12 M3 M4 H6 HS

LIC*!Pi\ß gererdii 1 1 Pexillue involutue 1 1 Certinerius ne. 3 1 Russule clerofleve 1 Ruesole heterophylla? 1 I Leeteriue griseus Russel! ubfoetens? 1 Aunnite strengulate 3 2 loletue nffinie? 1 Entolenn so.1 Hygreehorus sp. 1 Russula crueteee 1 1 Nygroohems enneeius 1 3 Russule virosctu 1 Russo]! eperta 1 1 1 Entolene ¤u•—•-eü 3 Älütl vegirnte 3 1 S 1 Nygrepherus flavoscrse _ Iussul! r•de1•1s S 3 3 Nyuronhorue psitticirh 6 1 Laetsrius thejogelue 1 S 3 Enteloun sellunenn ‘ 8 Phyllooerun r+¤d¤xu•thu• R 3 S 1 Inecybe sp. 1 1 _ 3 Entoleun lsgmicystis S Boletus d*••·ys•nt•r¤s 8 S 3 1 8 Russuls krcnbholzii 1 11 Sclerodenn citrinn 3 S 31 Mygrophorus centherollue 1 Lactarius cinerous 9 9 1 3 Boletinellus nerulioidee 11 36

Chapter 1 27 Ordination of plots by tree basal areas (Fig. 4) also indicated

discontinuity between spruce and hardwood plots. A strong disjunction

among the hardwood plots was evident along both the first and second axis.

This disjunction was probably caused by the high basal area of Qge;eye

and Iggy; in H4 and the high basal area of Iegye in H3 (Teble 3). Plots

H5, H6, H11, and H12 were strongly dominated by £;ggge and Age; and were U not widely separated along any principal component.

The main similarities between the ordinations of plots by fungal

species and tree species were (1) the polarization of the spruce stands

at the negative end of the first axis because of the taxonomic

discontinuity between spruce and hardwood plots, (2) the limited

separation of spruce stands by additional axes because of their relative

homogeneity, and (3) the wide separation of hardwood plots on all three

axes because of their relative heterogeneity. However, the most

dissimilar hardwood plots based upon fungal species (H5 and H6) did not

correspond with the most dissimilar plots based on tree species (H3 and

H4). Principal component analysis is usually applied to continuous

vegetation samples but can estimate the number and distinctness of the

discontinuities in the data matrix (Noy-Meir, 1973). If the

discontinuities are not absolute, as are those between spruce and hardwood

plots, the discontinuities can be detected by asymmetry of the positive

or negative values of the various components.

The fungal communities were more similiar to each other than the tree

communities because nine fungal species occurred in both forests types.

Chapter 1 28 ? ’ § os { ä Os, I °ss { E Osa Os") ' °Hs •• • _27 E yu { H11 H12 .3 I I I I wa I ° —a o , S 7 l PRINCIPAL COMPONENH

‘ I O I us 5 E I 9 css E E J I ä I é °$° a E <> S2 I G °*··———·Og1 I ······— •H‘ OS1 I Mg! i ••43 Nr! •"‘ 'J {' ”0 ·J J [ 1 •·¤•6c•m couwuusnn

• Figure /6. Ordination of spruce ( 0 ) and hardwood ( ) plots by fungal frequency.: Fist three components extracted by PCA of frequency values of fungal species. The labels beside the plots are indicated in Table 2. The first axis accounted for 26 74 of the variation in the data matrix, the second axis 1/6 %, and the third axis 13 %.

Chapter 1 29 Some of the species common to both forests were among the most ubiquitous (a-s-Las1:.a1:.1.u.1. Bsmsnlaxramuasa, Amanisalnmarasa, and

Lgggggjg lggggtg). These species are widely distributed in many forests

types and thought to have a broad ectomycorrhizal host ranges.

AND11EN.S.I1'X

The frequency of major species is a measure of their relative ubiquity

within the plots (Tables 8, 9). These major species accounted for 88 Z

of the total frequency values, 96 Z of the total sporocarps in the spruce

plots, 66 Z of the total frequency values, and 75 Z of total sporocarps

in the hardwood plots during the three years.

Sporocarp frequency and density were highly variable among years.

Sporocarp frequencies and densities were consistently higher during the

second year for nearly all species, while the lowest values usually

occurred during the third year (Tables 8, 9; Fig. 5). Some species did

not fruit every year, especially the third year (Tables 8, 9; Fig. 6).

Species with the greatest sporocarp frequency or density were not the same

every year. Comparison of sporocarp frequency and density indicates that

the list of major species would be somewhat different if species were

ranked by density. For example, the sporocarp density of Qjgyyling

ggglgtgs during the study, but L. ggistgtg was greater than Lggtgrjgs

Chapter 1 30

i E < § 0 E OHL ä I E äbsr %• I S7 "° I °°" „ °¤s2 I

·2 0 3 5 mmcnm commsurw

· I I OH4 I · I ·; I E •HI2 S20 S‘@°:$‘ I ••Ms ä S7 Sßsg I _, ° •"‘ un} E I Eä' -a OH)

I ~s ‘ -2 0 a s r¤••cum0¤u•=0••s»m

• Figure S. Ordination of spruce ( 0 ) and hardwood ( ) plots by tree basal areas.: The first three components extracted by PCA of basal area values of tree species. The labels beside the plots are indicated in Table 2. The first axis accounted for 29 % of the variation in the data matrix, I the second axis 22 %, and the third axis 16 %.

Chapter 1 31 ggglgtys was more frequently encountered and therefore was more

ubiquitous throughout the spruce plots.

Sporocarp frequency and density were much higher in both the spruce

and hardwood plots during 1981 and 1982. (Tables 8, 9; Fig. 5). Fruiting

was severely suppressed by the extreme drought conditions during July and

August of 1983 (Table 1). July and August of 1983 were the driest in

eastern North America in nearly 50 years (Wagner, 1984). Sporocarp

frequency and density was greater in spruce plots than hardwood plots

during the first two seasons but not the third.

The higher sporocarp frequency and density in the spruce plots may be

due partially to the greater number of trees providing a substrate for

ectomycorrhizal fungi. Spearman°s rank correlation coefficient (p) was

. used to measure the association of vegetation parameters with sporocarp

density and frequency. When all twelve plots were considered, the density

of ectomycorrhizal canopy trees (£;gxing; included to account for

frequency and density of ßglgtjngljgg mgrgljgjggg) was positively

correlated with sporocarp density (p = 0.69; P = 0.01), the basal area

of ectomycorrhizal canopy trees (lggxingg included) was positively —

correlated with sporocarp density (p = 0.75; P = 0.005), the density of

ectomycorrhizal canopy trees was positively correlated with sporocarp

frequency (p = 0.63; P = 0.006), and the basal area of ectomycorrhizal

canopy trees with sporocarp frequency (p = 0.73; P = 0.007). When the

six spruce plots or the six hardwood plots were analyzed separately,

Chapter 1 32

I neither tree basal area nor tree density was correlated with sporocarp ‘ density or sporocarp frequency.

. Several investigators have observed an inhibitory effect of ferns and

herbaceous vegetation on sporocarp productivity (Wilkins and Harris,

1946; Wasterlund and Ingelog, 1981). This effect was observed in some

of the hardwood plots where fern cover ranged from 1.5-58 %. Within the

hardwood plots, there was a negative correlation between fern cover and

sporocarp density (p = -0.87; P = 0.05) and fern cover and sporocarp

frequency (p = -0.90; P = 0.04).

ERHIIINQThe

fruiting season for ectomycorrhizal fungi begins in early July and

extends into late September or early October for both forest types (Fig.

5). Sporocarps were observed for 89, 91, and 75 days for the three years,

respectively. The end of the first two fruiting seasons coincided with ·the advent of heavy frosts or snowfall. In the third year, prolonged drought in combination with cold temperature prematurely ended the

fruiting season. In both forests during the first and third years,

sporocarp density declined sharply after the first of August because of

late summer drought (Fig. 5). Both forests exhibited a strong peak of

sporocarp density in early September of the second season. During the

second half of August, 1982, rainfall was much greater than the same time

Chapter 1 33 T•b1• so•¤i•• 8. Tan nat faqnnt Ifr•q| •¤t¤¤yarrhi¤1 in sorua f¤r••ts y••r1y in 386 6 6* ande-ots, dung with areunt fr•¤u•rn¤=y» •o•r0¤•ro •o•ci•• fN•I¤'¤¤¥» und dunsity. 1n¤1ud•• oaunt in nt last SZ of Hu qndntn.

2:::*** '··* ¥::;‘=' zrzvm .· •„:1ix:*··‘* Lactnria o¤u1•tu• 179 67 81 65 127 827 82 155 677 3105 83 6 9 59 C1•vu1in• crishta 168 39 81 60 166 1081 -82 138 1739 11322 I 83 0 0 0 8o1•Qu• bndius 81 21 81 10 13 85 82 76 107 697 83 3 7 66 vir•¢•¤n.•f••an• L•ct•riu• 67 17 81 21 33 215 82 57 165 966 83 0 0 0 Ianih f1•v¤¢«ni• 66 17 81 25 63 280 82 56 99 966 83 13 23 150 Loctarius 1ig••y¤t•11u• 57 15 81 26 36 221 82 61 61 397 83 13 23 150 !n¤cyb• sérirl 38 10 81 26 36 221 _ 82 28 78 508 83 3 6 26 |N••u1• g•·uu1¤t• 30 8 81 9 9 59 82 25 37 261 83 0 0 0 lanih i•n•ur•t• 21 5 81 8 10 65 82 13 16 106

83 2 3 zu Lactnrim 19 5 81 0 0 0 82 19 65 293 83 0 0 0 other 95 26 81 13 15 98 82 81 113 736 l 83 2 1 7

82 687 2935 19108 83 29 69 319

Chapter 1 34 ‘I’•b1• T•n •¤•t ••>•ei•• 9. fr•v.·n•nt lfr•qI •¤to¤y¤¤rrhiz·a1 in Iurdvood for••h ••‘ fr•q.••1cy, in 304 4 mndr•h• •1¤ng nüh p•r¢•nt y•••·1y fr·•qu•ncy• md sooroarp •p•ci•• 1••st dnnsity. In¤1ud•• in at · ZZ of Hu qndrnh.

„= ä2‘f$§."""

uehrius a¤¤h¤•·•tu• 23 6 01 0 17 111 02 10 M 206 03 0 0 0 luuub grnnubh 23 6 01 7 7 46 - 02 17 33 215 03 4 4 31 0o1•tin•11u• n•ru1i0id•• 23 6 01 16 x• 55 365 02 64 zao 03 3 7 55 - Sc1•ro&r—, cifrinn 21 5 01 2 2 13 02 20 33 215 03 3 3 23 L•¤ari• boah 17 4 01 4 6 39 02 13 45 292 03 0 0 0 Runsula kr¤••t|·••lzii 12 3 01 0 0 53 02 2 2 -13 03 6 10 70

Hygruohonee a·•{h•r•11u• 12 3 01 0 0 52 02 5 0 52 03 1 2 7 !h•jop1u• Lsohrius 10 3 01 3 14 169 02 10 26 91 - 03 0 0 - 0 _ cin••·•u• Lachrius 10 3 01 0 0 0 02 9 10 117 sx 1 1 7 d·••·ys•·•hr¤·• loloius 9 2 01 0 0 0 az 1 10 65 as 2 s Z3 . oh, 02 37 01 Z5 49 319 . sz sa 72 469 03 9 15 90

02 169 327 2129 03 29 45 293

Chapter 1 35 period in 1981 and 1983 (Table 1). Heavy rainfall in late August appears to be necessary for dense late summer fruiting.

Fruiting phenology was comparable among major species in spruce and hardwood plots (Fig. 6). Based on 1982, most sporocarps of most species fruited in late August and early September. However, some species tended to have their peak density earlier in the season, e.g. Amggjgg

.flmm.¢.¤n.La„ Bnlsmshadixu, Kusaylazzamllßta. a¤dRus.mzl.a-

LENQIH Q2 SAMBLING BERIQD

More than one year was needed to observe all the species included in this study, and the number of species sampled varied among years.

Seventeen of the total species in the hardwood plots (47 Z) and 15 of the total species in the spruce (56 Z) plots were found in 1981. If only 1982, the year with the greatest density and frequency, had been observed, 30

(89 Z) of the hardwood species and 24 (89 Z) of the spruce species would have been found. Only 12 (33 Z) of the total species of the hardwood plots and seven (26 Z) of the total species in the spruce plots were found in

1983. No additional species were found during 1983. This may be partially because of the low sporocarps density during this year. Limited observations of the plots during a fourth year (1984) yielded one sporocarp of an indetermined Rgssglg that had not been found in 1981-83.

Chapter 1 36 The fruiting habitats of both well known and poorly known Basidiomycete

species were analyzed in this study. Several of the species found in our

spruce plots had been described earlier in the mycological literature as

fruiting under "spruce", presumably red spruce, in eastern North America.

These species include Amgnitg flgygggßjg (Hesler, 1960), ßglgtyg hggig;

(Snell and Dick, 1970), Iylgpilgg fgllgug (Snell and Dick, 1970),

Lggtggjgg dgggptiygg (Hesler, 1945; Hesler and Smith, 1979), L. ggg1g;gg'

(Burlingham, 1908), L. sgrdijgs (Burlingham, 1908; as Lgg;g;;g ggggig

(Weinm.) Fr.), L. yigggggggjggggng (Burlingham,1908; as Lggyggjg

ghgjggglg (Bull.) Fr.), Kggsylg gggnglgtg (Singer, 1957; Bills, 1984),

Lg_g_t_g;_j,g_5_wasand Kgggyjg (Bills and Miller, 1984).

described from a red spruce forest on Clingman°s Dome, Tennessee

(Hesler and Smith, 1979) and has not been reported elsewhere. The

remainder of the species in the spruce plots apparently never have been

reported fruiting in association with red spruce.

A high proportion of the species diversity in both the spruce and

hardwoods was attributable to species of the Russulaceae. This would

probably be the case in most boreal or temperate forests dominated by

ectomycorrhizal trees. Certain genera of ectomycorrhizal Basidiomycetes

commonly found in other coniferous forests (especially Ring; or Lggig) were absent from the red spruce forests. Among these are species in the . genera Irishclnma. Hxzmuhnmla (S¤<=ti¤¤¤

Hxzmnhoma.Chapter1 37 Qgmphjgjgg, Qhgggggmphgg, and Sgjllgs. There are no reports of hypogeous

Basidiomycetes in red spruce forests.

Most of the species of the hardwood plots are co«~on in the deciduous forests of northeastern North America. Entglgmg lgggnigygtjg is known only from North Carolina and Tennessee (Hesler, 1967). Prior to this study, Kgsgglg gggglgns was known only from the type locality in Vermont

(Bills, 1984). Kg5;g1g.gpg;;5 is a poorly known species described from northern hardwood forests of Vermont and may be synonymous with Buggy]; pggillg Peck (Singer, 1957). A few of the species from the hardwoods have been reported to be associated with specific woody hosts; e.g.

ßglggjggllgg mgggligjggg with Egggjngs spp. (Snell and Dick, 1970),

Lastaxiuasinezmawith F.az¤1szz.a¤s1i.f.o.1i¤„ and Lastariusthsrinaalus with

ßgtyla spp. (Hesler and Smith, 1979). These associations were evident in this study because plots with high density of these species either had these trees in the plots or near the edge of the plots (Table 7).

§glg;ggg;mg giggjggm which fruits in association with many woody plants, consistently produced sporocarps near Qggggyg gghgg in plot H4. Lgggggig lgggggg, Lgggggigg ggmphgrgggs, and ßggsglg ggggylgtg had the widest amplitudes in habitat of any of the major species, and both were found fruiting in most plots.

The number of species found in spruce (27) and hardwoods (36) compares favorably with numbers of species reported by other investigators. Hora

Chapter 1 38 120 HARD? · 100

I0

[Q . Q so E ’° ä g 0. V1 IL O ‘$° g svnucs ID g 12so Z 1000

7

500 250 ...... \ 0 7/1 8/1 9/1 ” 10/1 DATE

Figure 6. Sporocarp phenology of all species in hardwood and spruce • • plots.: 1981 (-4--+-), 1982 ( ), 1983 (...•....•. ). Note differences in scales on y-axis.

Chapter 1 39 II g N/Ax utunus •cu1•un /’ ucuruusW'“¢•1••11v•t•nu• · Ilvuco \, \ 1•• 1

1/ 1‘ ’, A Ä ’,A\\ z x I "\ ·“ \\ -•·¤--„....,,,• -

‘ QClavulllucnsuu •¤vuc•\NSlutl "'°‘V°' '“"°'"" 1°-

{/ ".\ / f \1 n \‘N-- "* -.:1,

¢•rn01•¤•‘•¢•ß /*-1 „ ugunus //' x 1 GUNS! 1 fx \ 9 /1 ‘/ \~ \1

‘• /f \‘ N A1"

1 LEKINIIS VHI¢OWIHI§¢O|‘\$ ߤ\ |u§|y|; °yg-luga gg, _ 1¤r¤•100¤ ! / \ N IO _/ I, ‘,

,/ \•

hnctnnmus 1, I1 1N 1/ • \\ Anunnu Havacoma \N :” I \N / \ \ I "~·· II I] ‘¥ ‘x AN \N

I/‘ A U, vg U1 1011 111 II' 1011 uu MY!

Figure 7. Sporocarp phenology of some major species in spruce and hardwood plots.: 1981 (··l•--A--) , 1982 ( , , ), 1983 .•. •. (. .. ). Note differences in scale on y-axis.

Chapter 1 40 (1959) reported 25 species and Richardson (1970) 12 species belonging to

ectomycorrhizal families fruiting in Scots pine plantations. Fogel

(1976) found 24 putatively ectomycorrhizal hypogeous fungi in a Douglas

fir stand. A Swedish stand of Norway spruce had 25 species fruiting

andin(Wasterlund and Ingelog, 1981). Stands of Qggrggg

Great Britain produced 4 to 11 species (Hering, 1966). In addition,

the numbers of epigeous species fruiting in various spruce stands is

comparable to the number of types of ectomycorrhizae observed on spruce.

Worjciechowska (1960) described 16 "form genera" of ectomycorrhizae on

Norway spruce within its northern range in Poland. Thirty·seven ”form

genera" of ectomycorrhizae were observed throughout the range of Norway

spruce in Poland (Dominik, 1961). Thomas gg gl. (1983) observed 25 types

of ectomycorrhizae in English Sitka spruce plantations.

_ We believe the stands described here are representative of other red

spruce and northern hardwood stands in the Southern Appalachians. Most

of the major species reported here have been observed fruiting abundantly

in other red spruce and northern hardwood stands in West Virginia,

Virginia, and North Carolina. However, many other Basidiomycetes

belonging to ectomycorrhizal families have been collected in spruce

andnorthernhardwood forests throughout the Southern Appalachians.

Therefore, my results do not represent the entire mycorrhizal fungal

flora.

Some doubt exists as to whether the concept of minimal area can be

applied to fungal communities (Christensen, 1981). An adequate

Chapter 1 41 species·area curve for macrofungi cannot be determined at a single point ’ in time. The species·area curves derived from retrospective examination

of the frequency data resembles Christiansen°s (1981) species-isolates

curves for soil fungi. In her studies, repeated isolations of soil fungi

within one plant community continued to yield additional species with no

tendency for the species·area curve to level off. Fogel (1976) determined

the minimal sampling area for hypogeous fungi in a Douglas fir stand

during peak sporocarp production to be 100 mz. Arnolds (1981) concluded

that fungal species numbers continued to increase in grasslands at plot

sizes up to 400 mz and that a plot size of 1000 mz may be preferable to

ensure an adequate sample size.

The number of ectomycorrhizal species fruiting in a small area may

reflect the minimal number of available niches in the rhizosphere. But

few estimates of ectomycorrhizal species density in small (<10 mz) areas

are available. Several species of ectomycorrhizal fungi can occupy a very

small root surface area (Zak and Marx, 1964). Up to seven ectomycorrhizal

fungi have been isolated from a single four-year old Ring; glljggtji tree

(Zak and Marx, 1964). Deacon gt gl. (1983) reported at least five types

of ectomycorrhizae occurring within a seven m radius of a young birch.

These estimates of species density are within the range of the maximum

of seven species of fungi fruiting in a single 2 x 2 m quadrat reported

here.

Direct and indirect gradient analysis, and other multivariate

techniques could be adapted readily to the description of macrofungal

Chapter 1 42 communities. Ordination has been used to compare communities of yeasts

(Bow1es and Lachance, 1983) and soil fungi (Christensen, 1982). In this

study, PCA ordination clearly differentiated between the fungal

communities of spruce and hardwood forests, emphasized the similarities

in species composition among spruce plots and the dissimilarities in

fungal species composition among hardwood plots. In addition, it

demonstrated how strongly fungal species composition of the plots was

influenced by tree species composition.

Frequency estimates must be interpreted with caution. Frequency

depends upon the pattern and density of the individuals (or in this case

sporocarps), the size and shape of the quadrats (Greig-Smith, 1983), and

in this study the duration of the observations of the quadrats. The

resolution achieved by frequency can be increased or decreased by varying

the quadrat size. Sporocarp frequency cannot measure the true extent of

a mycelium or distinguish between sporocarps produced by a large

continuous mycelium or many cluster of small localized mycelia. Sporocarp

frequency does not provide information on fungi that do not produce

epigeous sporocarps. Frequency may give an impression of the horizontal

distribution of mycelia but indicates nothing about their vertical distribution.

Sporocarp densities of the spruce plots were within the range for

sporocarp densities of ectomycorrhizal fungi in other coniferous forests.

Sporocarp density in the spruce plots ranged from 319 to 19,180 sporocarps ha" yr". Richardson°s (1970) estimates ranged from 8750 to 20250

Chapter 1 43 sporocarps ha" yr°‘, and Fogel°s (1976) estimates ranged from 11052 to yr'“ 16753 sporocarps ha" The sporocarp densities of the hardwood plots

in this study were lower than for other coniferous forests. Sporocarp

density in the hardwood plot ranged from 352 to 1081 sporocarps ha°‘yr°‘.

An impression of the overall pattern of fruiting phenology is difficult

to gain from comparison of the three years. The late summer drought of

1981 and 1983 contributed to the high Variation in phenology. The 1982

season had adequate rainfall during July and August and presented a

phenology pattern similar to those described for other temperate and

boreal plant communities (Wilkins and Harris, 1946; Lange, 1948;

Richardson, 1970; Petersen, 1977). All these studies show the typical

strong peak of productivity near the end of the season.

l _ The length of the fruiting season for ectomycorrhizal fungi at

high-elevations in West Virginia appears to be relatively short compared

to fruiting seasons of ectomycorrhizal fungi at lower elevations or in A maritime climates but is commensurate with those in high-latitude,

low-elevation communities. Sporocarps of ectomycorrhizal fungi were

observed in both spruce and hardwood plots from July to early October

(75-91 days). Wilkins and Harris (1946) reported a fruiting season

lasting from August to November in an English pinewood and from June to

November in a beechwood. In Scotland, ectomycorrhizal fungi fruited from

June to December in a Scots pine plantation (Richardson, 1970). In.R1ggg

yiggigiggg stands in southwestern Virginia (elev. 450-750 m),

ectomycorrhizal fungi fruit from June to December (author°s unpublished

Chapter 1 44 data). The fruiting season is much shorter in Greenland tundra (Petersen,

1977), where ectomycorrhizal fungi fruit from July to mid-September.

Although only one additional species was found after the third year, it is difficult to judge how many additional species would be found if more years had been sampled. Fogel (1976) sampled 98 % of the hypothetical number of hypogeous species of a Douglas fir forest in a three year period. Arnolds (1981) observed most of his grassland sites for three years but observed some selected sites up to six years. Based upon these six year observation periods, he concluded that three years of sampling yielded 75-92 % of the total species. However, during these extended sampling periods, the Vegetation of these grasslands changed significantly, which may have contributed to subsequent additions to the mycoflora. Lange (1978) recorded 266 mycorrhizal species in a series of

Vegetation types over a ten-year period in the Beech Wood District of

Denmark. During any given season, 21-S9 % of these species were observed to fruit.

In summary, this study provides baseline data on the diversity, density, frequency, and phenology of ectomycorrhizal fungi fruiting in red spruce and northern hardwood forest types. The fungal species composition and sporocarp density of the two forest types were distinctly different because of a strong dependence of fungi species composition on the composition of tree present. Inter-plot similarities and differences in fungal species composition and the dependence of fungal species composition on the tree composition was emphasized by PCA, one of many

Chapter 1 45 ordination techniques which may be adapted to future studies of fungal

communities in response to vegetational or environmental gradients or

identifying fungal community structure in complex Vegetation landscapes.

Fungal species composition and relative abundance of fungal species in

other red spruce or northern hardwood communities may shift with changing

latitude, or with slight changes in tree composition.

Species species richness was not significantly different between ‘ spruce and hardwood plots, the hardwood plots were more diverse because

of greater equitability among species. Evidence for this greater

diversity included the Shannon-Wiener index, the dominance diversity

curve, species area curve (indicating more species among hardwood plots,

and PCA ordination.

The results suggest that estimating sporocarp frequency in small

contiguous quadrats may be a more appropriate method than estimating

sporocarp density for comparing the the relative activity or ubiquity of

fungal species in plot studies. Fruiting phenology was comparable between

the two forest types, and the fruiting season was relatively short. The

fruiting season at this southern latitude was compressed because of the

short growing season at high-elevation. Variation in fruiting season is

assumed to occur in response gradients of altitude or latitude, but

quantitative, comparative studies are needed to measure the extent of this Variation. _

Chapter 1 46 QHAHIEBL. S.BAI1AL2AIII":1RhJ§ANDQE IHKE12$.BB§l£EAhIDIüBHW9QDEQRE§IS

Basidiomycetes usually fruit in spatially and temporally aggregated

patterns. Spatial aggregation is caused by multiple sporocarp production

by a single or a series of localized mycelia. Heterogeneity of habitat,

inoculum density, and non—random occurrence of host plants and substrates

contribute to the non-random patterns of mycelial occurrence. Temporal

aggregation is caused by repeated sporocarp production by perennating

mycelia. Basidiomycete fruiting is often described in the literature as

gregarious, clustered, in groups, or caespitose. The perennial nature

· of fruiting is best exemplified by the mushroom collector who returns

year-after-year to the same location for edible fungi. Perennial fruiting

has been expressed quantitatively as "constancy" (Lange, 1948) or as an

"index of specific fluctuation" (Arnolds, 1982). The spatially

aggregated, perennial, fairy ring fruiting pattern of certain grassland

fungi has been thoroughly investigated (Ingold, 1974; Smith, 1980;

Edwards, 1984). However, relatively little is known about the spatial

patterns of terrestrial sporocarps in forests, and variation of these

patterns within or among species (Fogel, 1981)

Observation of spatial patterns of sporocarps in reference plots

yields several kinds of information related to the biology of the fungus

Chapter 2 47 and its associated organisms. Sporocarps verify the presence of the vegetative mycelium, and sometimes sporocarps numbers and their patterns have indicated the spatial pattern and relative abundance of the vegetative mycelia (Laiho, 1970; Thompson and Rayner, 1982; Last gg gl.,

1983; Edwards, 1984; Newell, 1984; Cotter and Bills, in press). In addition, the spatial patterns of sporocarps may influence the spatial patterns of animals that utilize sporocarps as feeding or breeding sites

(Shorrocks and Charlesworth, 1982; Ashe, 1984; Lacy, 1984). Knowledge of spatial patterns of sporocarps is essential to the understanding inoculum density and dispersal, and of the establishment of °

Basidiomycetes in forests.

How should fungal communities be sampled to yield maximum information?

Ideally, as in plant or animal communities, quadrat sizes and placement would be determined by the species-area relationships within the community, the relative sizes of the organisms, and the Variation in the spatial patterns of the organisms. These critera are difficult to apply to fungi because the composition of the community and its spatial pattern based on sporocarps is continually varying (Fogel, 1981). Repeated and prolonged observations of fungal communities are needed because of the temporal Variation in their compositions. In addition, the lack of knowledge of variation in sporocarp density and the unpredictable nature of species density has prevented the application of consistent sampling methods with subsequent difficulties in comparing and interpreting results of different investigators (Hueck, 1953). Knowledge of spatial

Chapter 2 48 variation of fruiting patterns can be applied to further improve sampling

· methods for fungal communities.

Quantitative methods for describing spatial patterns of plants or

animals have rarely been applied to higher fungi (Fogel, 1981). Recently,

the spatial patterns of plant pathogenic fungi and diseased plants have

been investigated in several agricultural systems (Campell and

Pennypacker, 1980; Taylor gt al., 1981; Nicot gt gl., 1984; Shew gt al.,

1984). The objectives of this study were to describe and compare

quantitatively the spatial patterns of sporocarps of terrestrial,

presumably ectomycorrhizal, basidiomycetes commonly found in

high-elevation red spruce and northern hardwood forests of West Virginia

and to determine whether interspecific associations or antagonisms among

fungi occupying a similar trophic level could be detected based on ” sporocarp observations alone.

MEIHDDS

Both temporal and spatial aggregation were combined by summing the

cumulative sporocarp densities and frequencies for 1981-1983. From

cumulative sporocarp densities in each 2 x 2 quadrat in each plot, the

variance-to-mean ratio (V/m) (Pielou, 1974) was calculated for each major

(frequency Z 5 %) species in each plot and for each major species in each

forest type. .The biological null hypothesis that sporocarps fruit at

random among the quadrats of a plot is tested as the statistical null

Chapter 2 49 hypothesis that sporocarps are equally likely to occur in each quadrt.

Greig-Smith (1983, p. 62) explains that under this null hypothesis the

V/m is 1 and its statistical significance is tested by a

goodness-of·fit-test of the form: l x= = (n-1)(V/m)

where v is the unbiased estimate of variance, m is the sample mean, and n—1 is the degrees of freedom. The number of quadrats, n, is the sample

size. A V/m not significantly different from one is expected if the ' sporocarps are distributed at random among the quadrats.

Mean crowding (m* m V/m 1) measures the mean number of neighbors = + ·

of the same species per individual sporocarp in a single quadrat.

Lloyd°s index of patchiness (m*/m) also characterizes sporocarp

aggregation by measuring the intensity of the pattern. In other words,

patchiness is high when the pattern shows strong density contrasts and l is nearly one or less if density contrasts are low (Pielou, 1974). The

patchiness index is independent of density under certain conditions.

Random removal of individuals from sampling units does not effect its

value, therefore patchiness is recommended for comparing the intensity

of fruiting patterns among sample sites or among species.

Spatial autocorrelation (Cliff and Ord, 1973; Sokal and Oden, 1978)

determines whether at one location is correlated with the value of a

variate depends on the values of the variate at neighboring locations.

This technique was employed to determine whether the presence of each ·

major species depended on its location in neighboring quadrats. Expected

Chapter 2 50 join counts of occupied quadrats were calculated under the null hypothesis that occupied quadrats were randomly distributed and that they could be joined as queen moves in chess or weighted by the inverse of the distance squared between centers of occupied quadrats. lf the observed joins of occupied quadrats significantly exceeds the expected, then they are positively autocorrelated indicating aggregation or clumping. If the observed joined are significantly less than the expected joins, then the occupied quadrats are negatively autocorrelated indicating uniform or regular patterning.

When a large number of species occupy a similar trophic level of a comunity, a natural question is, are they associated or do they occur independently of each other? When the number of species (k) is large the number of possible interspecific interactions increases geometrically. 2k Conceivably these interactions could be tested by constructing a contingency table, but this is impractical. An alternative test was developed by Barton and David (1959) and illustrated by Pielou (1974).

If species are distributed independently, the frequency distribution of species densities should approximate a binomial distribution. The deviation of the observed frequency distribution can be tested for significance by a X2 goodness·of-fit test.

To test for possible interspecific associations or antagonisms, the statistical significance for all co-occurrences for major species the X2 values using Yate°s correction for continuity was calulated for the 2 x I 2 contingency table (Pielou, 1974). In addition, Cole°s index of

Chapter 2 51 interspecific association (Cole, 1949) was calculated for pairs of major species. Values for Cole°s index range from +1 when two species are always associated to -1 when they never co-occur. A value of 0 indicates random association.

KE§llLI§

BAIIERNS ·

Sporocarps of all major species in both forest types were aggregated

(Figs. 8, 9, 11, 12; Table 11). The degree of aggregation varied among species (Figs. 8, 9, 11, 12) and among plots for a single species (Tables

10, ll) as indicated by the V/m, mean crowding, and patchiness. Low, insignificant V/m°s occurred only when sporcarp density was low

(<7/plot). Also as expected, when enough frequency classes were available, the frequencies of sporocarp densities/quadrat exhibited poor fits to expected values derived from a Poisson distribution.

In most cases, deviations from the expected values occurred because more quadrats had no sporocarps and more had high densities than expected indicating aggregation within quadrats.

The degree of aggregation of sporocarps based on the entire forest samples differed among the major species as indicated by the V/m, mean crowding, and patchiness indices (Figs. 8, 9, 11, 12). The V/m and mean crowding were highest in the coral fungus, Qlaygligg ggistggg, because

Chapter 2 52 it formed dense clusters of sporocarps (up to 261/quadrat/3 yr). Some

species. e- s· Russula xramlla:1. Lastarius lizn1o.ts.l1us„ Balems b.¤su.us,

and Amggjgg igggrggg usually fruited singly or in small groups. High

patchiness was characteristic of species producing localized, dense

clusters of sporocarps with the clusters separated by large intervening

areas where either sporocarps were absent (e.g. Lgggggigg ggmphgggggg,

lgggyhg gghgjng) or uniformly present at low densities (e.g. Qlgggljgg

g;i;;g;g) (Figs. ll, 13). Species with a low uniform sporocarp density

(e.g. Amggjgg jggggggg) or high uniform density (e.g. Lgggggigg

ggglgggg) were characterized by low patchiness indices.

With the exception of Qlgggligg grjggggg, the V/m and mean crowding

of the major hardwood and spruce species were comparable. However, the

patchiness indices were much higher for all hardwood species because the

gregarious sporocarp production of Lgggggjg lgggggg, Rg;;g1g ggggglggg, l Lgggggjgg gggphg;g;g;, and Sglgrgggrmg gigrjggm was restricted to the

rhizospheres of the few {ggg; and Qgggggg trees and ßglggjggllg;

mgggligigg; around the {rggiggg trees. Conversely, exclusion of

ectomycorrhizae from the areas dominated by endomycorrhizal tree species

and ferns may have contributed to the patchy distribution of sporocarps

in the hardwood forests.

Frequency maps (Appendix B) indicate that species often occured in

contagious patterns. The contagious sporocarp frequency of ßglggjggllgg ggggljgiggg (plot H6) was caused by two individual mycelial patches

producing sporocarps (Cotter and Bills, in press). Sporocarp density of

Chapter 2 53 1.6

1.2 O E

0.6 >o 6 O .J

0.•

Oro cc LV nu LO AF LC LL HG 88 ^*

SPECIES

Figure 8. Logl0 V/m ratio of major species in spruce forests.: Species initials are from Table 10. A11 log10 V/m ratios were significantly greater than 0.

Chapter 2 S4 00

es

O E O 2 O so 1 U Z < LU 2 es

V 0 ‘ cc LO LV uu AF LC LL as nc; AI

SPECIES

Figure 9. Mean crowding of major species in spruce forests.: Species initials are from Table 10.

Chapter 2 55 II

10 U7 V7 . UJ Z I _ U I- g | .

0 LC nu cc LV AF R AI LL es LO

SPECIES

Figure 10. Patchiness of major species in spruce forests.: Species ixxitials are from Table 10. .

Chapter 2 56 1.0

\‘

°_ aésséaäsLw 0 ¤M LC nc Sc

SP ECIES

Figure 11. Logl0 V/m ratio of major species in hardwood forests.: Species initials are from Table 10. All log10 V/m ratios were significantly greater than 0.

W Chapter 2 57 s

z ’-,;;:;L-

Lg am L c n 6 s c

S PE C I ES

Figure 12. Mean crowding of major species in hardwood forests. : Species initials are from Table 10.

Chapter 2 58 •c

E man‘°0•-¤

Lg EiägigäL c =E=i;E;E3En 6 =E=E=E=E=·a M :i=i=i=E=s 6 SPECI ES

Figure 13. Patchiness of major species in hardwood forests.: Species initials are from Table 10.

Chapter 2 59 ß. mgggljgiggs in one quadrat was clearly dependent on sporocarp density

in nearby quadrats. Although the mycelial pattern was not be directly

mapped for other species, the hypothesis that the occurrence of sporocarps

in a quadrat depended on the presence of sporocarps in nearby quadrats

rather than being randomly or regularly dispersed was tested using spatial

autocorrelation. Each major species, except ßglggggggm giggjggm, in both

hardwood and spruce forests was positively autocorrelated in at least one

of the plots in which it occurred (Tables 13, 15).

Most pairs of major species in the spruce forest were distributed

independently of each other as indicated by the X2 of the 2 ¤ 2

contingency tables and Cole's index (Table 16). However, :ignificantly pairsA.more co·occurrences than expected were detected in three species

flgyggggig). Significantly fewer co·occurrences than expected were

detected in three other species (A. flgyggggjg ggjstggg, pairs - Q. A. tlmzsaszuia · E- b.¤dJ.u.s. A- flaysmnla · K- z:.¤11ula:.a>- Significant x'

for association were always accompanied by Cole's indices equal to or

X2 greater than 0.18. Insignificant or low values indicating random or negative association were not always matched by low values of Cole's

indices. Low values for Cole's index were often the result of species being so rare that their co-occurrences were improbable.

Chapter 2 60 cf_th• spatial patterns gf ferrastrial Tabl•_10. Characferization by var1anc•—·to-mean sporocarps xn sxx spr-uce ploßs basgdxomyceta lm-!). and pafchxnass _ raho (Y/nl. nean_cs-owdxng ln!/ml xn 64 conhguous 2 X 2 n quadrafs.

pIo¥ ¥¤¥al uu! vu!/a specxas spororcarps V/an

46.79c 53.61 6.86 crisfata $1 500 5.74 Clavulina $2 163 13.07c 14.62 Clavulgna crrstaia 141.51c 148.56 18.46 crgstata $7 515 3.08 Clavulgna $8 605 20.68c 29.13 Clavulgna crgstafa 9.00c 8.94 9.53 $9 60 13.62 Clavulgna crgstata 62 13.23c 13.21 Clavulxna crxstata S10 1.72c 1.75 1.75 cculatus $1 64 1.84 Lactarjus $2 159 3.09c 4.58 Lactarrus oculafus 6.31c 7.97 3.00 oculafus $7 170 2.60 Lacfaqus $8 140 4.49c 5.68 Lacfargus oculafus 4.11c 3.79 5.63 $9 43 3.90 Lacfaqus oculatus 37 2.68c 2.26 Lacfarxus oculatus $10 0.49 $1 2 0.98 0.02 Bclefus badjus 1.66c 0.88 4.03 S2 14 6.53 Bolefus badgus $7 20 2.73c 2.04 Boletus badgus 1.44b 1.30 1.51 S8 55 2.18 Boleius bad;us 22 1.41b 0.75 badgus S9 2.71 Boletus $10 14 1.37b 0.59 Bolatus badxus — 0.02 1.00 vjnaceorufescens $1 1 0.52 8.36 Lactarjus $2 4 - Laciargus vgnaceorufescens 6.38: 1.89 4.33 vgnaceorufescens S7 28 6.94 Lacfargus $8 42 18.95c 4.55 Lactaqus vgnaceorufescens 37.75c 5.01 4.72 vgnaceorufescens S9 68 16.89 Lacfargus 35 17.55c 9.24 Lactarxus vxnaceorufescens S10 0.72 2.74 S1 17 0.75 Lactarius ljgvyofellus 7.35b 1.52 2.49 lrgsyofellus S2 39 0.69 14.77 Lac{ar;us S7 3 - Lactargus lggnyotellus 0.55 8.37 $8 4 - 4.00 Lacfargus lrgnyofellus 28 6.42: 1.75 lggvyotellus S9 0.69 7.37 Lactargus $10 _ _ 17 0.75 Lactarxus lzgsyoiellus 6 7.93 24 3.60c 2.97 flavoconia S1 5.55 3.91 Amanjta S2 91 5.13c Amangta flavoconga 1.46b 0.52 8.36 flavoconya $7 4 0.33 Amangfa $8 3 0.98 0.02 Amanyk: flavoconp 35 1.68c 1.22 2.24 flavocong.: $9 0.77 6.20 Amangia $1 8 1.65c Amanzfa flavoconxa 1.90c 1.03 8.23 $1 8 9.88 Inocyb• uubrjna 15 3.08c 2.31 $2 2.67 13.14 Inocyb• unbrgna $7 13 3.46c Inocybe uvbrgna 1.75c 0.83 10.59 S8 5 7.89 Inocybe urbrgna 54 6.82c 6.66 unbrgna $9 23.72 Inocybe $10 11 4.90c 4.08 Inocybe uubrxna 4.67 15 1.86c 1.09 ' granulata $1 2.04 18.71 Russula S2 7 2.93c Russula granulata 1.19 0.30 2.78 S7 7 3.91 Russula granulafa 13 1.59b 0.79 granulaf: S8 0.69 14.77 Russula $9 3 1.64b granulaf: 0.02 1.00 Russula S10 1 1.00 Russula granulat: 1.31: 0.62 1.99 Qnaurafa $1 20 1.00 Amanjia $7 1 1.00 0.02 Amangfa xnaurafa 1.00 0.02 1.00 $8 1 8.36 Amangta Qnaurat: S9 4 1.46b 0.52 Amangia gnaurata 3 1.64c 0.69 14.77 Amamta znaurafa S10 0.76 $7 24 2.41c 1.78 Lacfarjus eanphoratus 3.62c 2.83 13.91 S8 13 10.27 Lactargus camphoratus 8 2.16b 1.28 Lactarxus cauphoratus S9 Ü,Ö§ 3 5,ÖI < F < =< . b 0.001 < P =< 0.01 c P =< 0.001 for test enoudu frequency classes · nof

Chapter 2 61 i Tab1•_11. Charachrizafiop of_fh0 spafial Yntfems of_hrr•s·!ri•1 ·_ bas;d1omyce‘l:• gporocarps xn sgx harduoogl p ots_b¥ yar~1;n0•-fo-mean raho IV/nl, X ?o0

specxas pIo¥ ¥o¥aI spororcarps V/n nl nl/n

L•0hrjus canphorafus H3 11 1.750 0.94 5.45 Lachrgus canphorafus H4 27 9.020 8.44 19.99 Lachrgus camphorafus H5 12 2.680 1.87 Lachrgus 10.00 camphorafus H6 7 2.940 2.04 18.70 Lachrgus ¢8lI!h0|‘I*U$ H11 3 1.650 0.69 14.77 Lach:-aus caunhoratus H12 1 1.00 0.02 1.00 Russuln granulah H4 17 5.530 4.79 18.04 Russula granulah H5 6 1.26 0.35 3.76 Russula granubta H6 4 2.480 1.54 24.62 Russub granubh H11 11 3.060 2.23 12.97 Russula granulah H12 - 6 0.92 0.01 0.15 _ Bo-1•·tin•11us ¤••ru1ioid•• H6 106 5.040 5.70 3.44 _ I Sclsroderma cifrirun H4 37 2.400 1.98 3.43 Sclaroderma citrinun H6 1 1.00 0.02 1.00 La00•ria bccah H3 1 1.00 0.02 1.00 Laccaria . 1•00af• H4 8 2.410 1.50 12.30 Laccaria bccah H5 1 1.00 0.02 1.00 « Laccaria laccafa H6 32 11.430 10.93 21.86 Laccaria bccah H11 1 1.00 0.02 1.00 Laccaria bccah H12 8 1.650 0.78 6.21 a 0.01 < F =< 0.05 b 0.001 < P =< 0.01 0 P =< 0.001

Chapter 2 62 Table 12. Significance tests for spatial autocorrelation of - major basidiomycete species in spruce plots. Frequency (freq) is the number of quadrats a species occupies. S.N.D. is the standard normal deviate. Occupied quadrats (BB), occupied and empty quadrats (BW), and empty quadrats (WW) were joined by queen°s moves.

species freq no. of S.N.D. plots BB BW WW

Lggtggigg ggglgggg 179 6 2.92c -3.19c 1.64a

Qlgygligg ggiggggg 148 6 4.46c -6.52c 4.81c

ßglgggg hggjgg 79 6 1.90a -4.32c 3.93c

Las:.a:.um 67 6 7-35<= ·S•21<= 1.916

Aggjtg flgyggggjg 64 6 2.44b -4.31c 3.64c

ljggygtgllgg 57 6 5.80c -2.51b -0.17

lgggygg gmbgjgg 38 6 4.28c -1.12 -0.36 l Rgggglg gggglgtg 30 6 4.54c -5.56c 4.36c

1.89a Amggita jgggggtg 21 5 1.90a -2.48a

Lggtggigg ggmphgggggg 19 3 2.15b -2.39b 1.78a

a 0.05 < P S 0.10 · b 0.01 < P S 0.05 c P S 0.01

Chapter 2 63 Table 13. Significance tests for spatial autocorrelation of major basidiomycete species in hardwood plots. Frequency is the number of quadrats a species occupies. S.N.D. is the standard normal deviate. Occupied quadrats (BB), occupied and empty quadrats (BW), and empty quadrats (WW) were joined by queen°s moves.

species freq no. of S.N.D. plots BB BW WW

Lggtggiggyggmphgggggg 23 6 4.69c -2.72c 1.57

Rgggglg ggggglggg 23 6 3.49c -0.91 0.01

ßglggjggllgg mg;g1;gi§g5 23 1 6.47c -7.92c 5.08c Sslsmdema szitxinum 21 2 1- 47 -1- 20 0- 46 ° Lgggggjg lggggtg 17 6 4.13c -2.51b 1.54 a 0.05 < P S 0.10 b 0.01 < PS 0.05 c P S 0.01

’ Chapter 2 64 Table 14. Significance tests for spatial autocorrelation of major basidiomycete species in spruce plots. _Frequency (freq) is the number of quadrats a species occupies. S.N.D. is the standard normal deviate. Occupied quadrats (BB), occupied and empty quadrats (BW), and empty quadrats (WW) were joined by the inverse of the distance squared between their centers.

species freq no. of S.N.D. plots BB BW WW

· Lggtggigg ggglgtgg 179 6 2.37b -2.17b -0.07

Qlgygligg ggjggggg 148 6 3.33c -4.86c 3.72c

ßglgtgg hgjigg 79 6 0.99 -3.03b 3.04b

Lasxarius 67 6 4-95<= -3- 33c 1- 10 -

Amggjgg f1gygggn1g 64 6 1.86a -3.69c 3.29c

Lgggggjgg ljgnggtgjlgg 57 6 4.84c -2.17b -0.07

Lggggbg gmhgigg 38 6 3.70c -0.19 -1.19

Rgggglg ggggglggg 30 6 3.64c -5.14c 4.24c

Amggjgg jngygggg 21 5 1.16 -2.26b 2.00b K Lggtggjgg ggmphgggtgg 19 3 1.72a -2.16b 1.70a

a 0.05 < PS 0.10 b 0.01 < P S 0.05 c P S 0.01

Chapter 2 65 Table 15. Significance tests for spatial autocorrelation of major basidiomycete species in hardwood plots. Frequency is the number of quadrats a species occupies. S.N.D. is the standard normal deviate. Occupied quadrats (BB), occupied and empty quadrats (BW), and empty quadrats (WW) were joined by the inverse of the , . distance squared between their centers.

species freq no. of S.N.D. plots BB BW WW

Lagtggigg ggmghgrgggs 23 6 3.75c -2.51c 1.58

Ryggylg ggggglgtg 23 6 2.71c -1.05 0.34

ßglgtjggllgg mgggligjjgg 23 1 6.47c •5.88c 3.76c

ßglgggggggg gjtgjggm 21 2 » 1.66a -0.64 -0.33

Lgggagjg lggggtg 17 6 3.19c 3.19c 1.34

a 0.05 < P S 0.10 b 0.01 < P S 0.05 c P S 0.01

Chapter 2 66 one species (L. Only pair gamphgrgtus - R. grgnylgtg) in the hardwood

forests was positively associated (Table 17). However, quadrats occupied

by L- samvlxuamu. E- z:.¤m1l¤.ta„ L- lasaeata. and S- sitrinum after:

occurred near each other in plots H4, H11, and H12 probably because their

sporocarps arose from mycelia associated with the same [ggg; and Qgggggg

trees. The lack of co-occurrence among these species might have been

because the quadrats were too small include more than one or two species.

When 2 x 2 quadrats were combined into 4 x 4 quadrats, the co-occurrence

of these species still did not deviate appreciably from expected values.

The statistical significance for co-occurrence could not be tested ‘ because of low expected values obtained when the quadrats were combined

(1 to 3).

The hypotheses that all species of the spruce forest type (27) and all

the species of the hardwood forests type (36) were distributed

independently among the quadrats were rejected (Tables 18, 19). In the

spruce forest, fungi were absent in more quadrats (54) than expected under

the null hypothesis and fewer quadrats had four or more species than

expected (Table 18). The species of the hardwood forest type appeared

to be aggregated in certain locations and absent in others because no

fungi occurred in more quadrats than expected and two or more species

occurred more quadrats than expected (Table 19).

Chapter 2 67 • • r · S'. '6 S• ä ° , • ,,‘ Q ° Ü 6 H 9 4 II T ° ° _

N III ¢ Q 2,0 In -1 ‘ Q , , . 1 6 => Q J ° 9 9

. . Q NI Ü Q Q 2• S••2 2 Q Q . ' Q··· ··· 6 Ü 44 I 9 9 9 9 ¤

A 0 ¤ . • S 'a•• ¤ , S a' 2 Q Q Q . 9 6 pl Ü 9 ° ° ~• . 2 0 ~0 _ ä, S••2 $ Q Q ' 6 0 0 ° N Q -2N I ¤ '^

•'* { zu . Ih ~¤ 0 , ••In G• S, Ä••Z In In In Q' • . 9 QQ u • ° ° ••• . an Q. 0 QHg S :2 °; . II ggx_•_ 4-4 c , g g =· 0 III ¢ Q °§,•_ ,3 • 2 -1 *2 Q ää•-• •••¤•O 6 6 J Q Q N .·• ° Ö 0:}+ ° ••S '. -2:*58 ·•{ °¤_Q•« U' A M *2 ° *4 gg -^H ä Q-I3 , *0·•• "‘Ä• 3, M• °f Q fg ' rl QQ ,4 rl Egtä · .-I ¤ ·‘°§ $—•Q°6· _ 0 N ga ~ $0 ä ~e g 5 S vg - Q Q ‘ '6 Q *2° S.; MSE: ° N · 6 6 ¤ g.-I R ° ..3-°• ' gß > ° ° ° ° ° > «.••ss·g -·:....§ ° Ü„ an I g ÜN "°C.c·•-.2 ,2 •-I 4-• ••—°"'?‘·- ‘°0 Ü Q "'·•Q •‘ [ .~Ä .5 0% T-l: L 'V U Ü ••• QIä 3 ' un ·••U "‘E § .;•° E 2 ^^ *‘¤II. > L Ü 00 +--0-** 1 > ·Ü I-I = 8 ••••°U L ,.4 0-H „: .•• ‘: ...• •-·• -·• Ü ,,- +• 4, "I L -***7 ¤.• L 7 ...• • 2: N —•08 • ·•·*n * ·Q -•« ¢> g•• ··• ·¢ 0 C X JW "Q „0 "•U eo · }.·••¤•Xnum g H 7 E Q I U -¤ ,4 HÜ < .;° xx

Chapter 2 68 _- Table};. Hafrix of $2 l•f$) •n•i|1Co1q'• igdex afl _ assocxapxXmn ¤.pp•r rx va ues or a aus o mga s us ig hardnood {lots wifgia frequency of at Ieasf values ca culated usmg Yate's correchon factor.

E. c. E. 9. E. nn. S. c. C. I.

Lachrius cauphoratus 0.21 -1.00 -1.00 -1.00

Russula grnnulata I3.96! -1.00 -1.00 0.02

Bolefimllus narulioides 2.90 2.90 -1.00 -1.00

_ $cl•r0der¤•• eifrimn 2.76 2.76 2.76 0.13

Laccaria laccata 2.52 0.29 2.52 2.94 ••·• I•v•I ·l Rz vaIu•s > 6.04 $1§11¥l¢3h{ a¥ TF: 0.0I

Chapter 2 69 Mosaics are patterns resulting from different areas of a plane having

different properties (Pielou, 1974). Continuous colonies of vegetatively

propagating plants are often considered as vegetation mosaics. The

spatial patterns of sporocarps can also be envisioned as species forming

overlapping mosaics of different sizes with different densities of

sporocarps within the mosaics. The presence and absence of a mushroom

species forms a two-phase mosaic while the combined presence and absence

of several species forms a multi-phase mosaic. The boundaries of the

mosaic phases are not well·defined because of the variation in sporocarp

densities and, as with the vegetative clones of higher plants, mosaics

of mushrooms were variable in size and shape. The clonal nature of

adjacent sporocarps was not obvious, however, because scattered, widely

separated individual sporocarps may be connected by a continuous

widespread mycelium or they may arise from small widely scattered mycelia.

Mosaic patterns may form in response to either the heterogeneity of

the biotic or abiotic environment. Abiotic factors might include

subsurface rock, soil depth, soil moisture, or canopy cover. Biotic

factors might include the availability of colonizable host tree roots,

physical exclusion by large tree roots or rhizomes of herbaceous plants, 4 predation by soil invertebrates, or antagonistic or synergistic

interactions with other rhizosphere organisms. Measurement of the size

of the fruiting mosaics was often limited by plot boundaries.

Chapter 2 70 Table 18. The observed and expected freqencies of the number of basidiomycete species in 384 2 ¤ 2 m quadrats in hardwood forest plots. If species are mutually independent the number of different species per quadrat should approximate a binomial distribution.

quadrat frequencies ' (fobs - fexp): number of observed expected species fobs fexp fexp

0 226 206 1. 94 1 97 131 8. 81 ” 7 2 44 39 0. 64 3 14 7 4 3 18 1 8 12.53 S 1 0 6 and over 0 0

384 384 23.92

X2 = 23.92, 3 d. f. , P<0. 001

Chapter 2 71 Table 19. The observed and expected freqencies of the number of basidiomycete species in 384 2 x 2 m quadrats in spruce forest plots. If species are mutually independent the number of different species per quadrat should approximate a binomial distribution.

frequencies qllßdfßt • (fobs fexp)‘ _____i number of observed expected species fobs fexp fexp

0 54 36 9. 01 1 83 100 2.89 2 105 119 1. 65 3 ” 84 82 0. 05 4 42 35 1.45 5 ll 10 1. 33 6 4 16 2 12 . 7 1 0 ‘ 8 and over 0 0 EZ E 16. 38 X2 = 16.38, 5 d. f. , 0.01>P>0. 001

Chapter 2 72 The spatial patterns of the fungi were different between the two

forests and were highly dependent of the distribution of the canopy trees.

The mosaics formed by major species in the spruce forests often approached

the size of a single plot. Extrapolating to the scale of an entire spruce

forest, fruiting mosaics, and presumably the corresponding mycelia, of

common species can occur over extensive continuous areas. These patterns

indicate that a large percentage of the spruce trees in these homogeneous

forests have the same fungal species associated with their root zones.

At the finer scale of the quadrat in the spruce forest, extensive overlap

or intermingling of mosaics occurs, probably because single trees can

support several species of fungi. In contrast, fruiting patterns in the

hardwood forests indicate that only a relatively small proportion of the

rhizosphere was occupied by ectomycorrhizal fungi. The fruiting mosaics

· were relatively small and restricted to the vicinity of ectomycorrhizal

trees or were·excluded from the rhizospheres of endomycorrhizal trees and

ferns.

Indices of spatial aggregation based on heterogeneity of mean

densities not only demonstrated that sporocarps occurred non-randomly but

appeared to be useful for describing and comparing the degree of sporocarp

aggregation and the dispersion of the sporocarps aggregates. The indices

confirmed mathematically the investigator's intuitive perception of

sporocarps patterns and the differences in patterns among species. The

patchiness index was especially useful for illustrating the differences

in sporocarp dispersion between the more structurally homogeneous spruce ‘ forest and the more structurally heterogeneous hardwood forest.

Chapter 2 73 Spatial autocorrelation was of limited value for describing the spatial patterns of sporocarps. Large areas where sporocarps occurred in contiguous quadrats were often not autocorrelated probably because of the small plot size relative to the size of the fruiting mosaics (Tables

12, 13, 14, 15). A more satisfactory impression of spatial pattern was provided by inspection of the frequency maps (Appendix B). For example, inspection of the frequency of S. gitgingm in plot H4 indicates an occupied quadrats are closely associated with the root zones of Qgggggg ggbgg. However, the join count statistics were not significantly greater than expected under the null hypothesis of random pattern.

Statistically positive or negative associations between two species could occur for several reasons. The two species could be mutually beneficial on inhibitory or one could exert a synergistic or antagonistic influence on the other. When environment is variable, the two species may have similar or exclusive tolerance ranges for environmental conditions (Pielou, 1974). When a large number of 2 X 2 comparisons are made, the results of a few (5 Z if the significance level is 0.05) will correspond to a low probability simply by chance. In the 45 comparisons of major spruce species, two or three or the associations could have been chance events. In the 20 comparisons of major hardwood species, one association could have been a chance event. Rejection of the null hypotheses of independence of species might mean that the quadrats are dependent. This kind of positive association might be observed when a coarse·grained mosaic of two species is sampled with closely spaced quadrats (Pielou, 1974). If the mosaics of both are large relative to

Chapter 2 74 the study area, then extensive overlap of the large mosaics of each

species will result in greater joint occurrence than expected. When viewed at a larger scale over the entire forest, the overlap of could be a chance event without assuming mutual benefit or heterogeneous habitat.

The three positive associations in the spruce forest could have been caused by overlapping mosaics because all the species that were positively associated had high freqencies. The involvement of one species, Amnjgg flgyggggjg, in all the negative associations seems more than coincidental. Fruiting of Amanitg flgygggnig was noticeably constant in certain quadrats over the three years. Further studies designed to test the hypothesis that root zones colonized by A. flgyggggig cannot be infected by other mycorrhizal fungi might be profitable.

Chapter 2 75 QHAEIERL. S.INQES.I§QERH§§LlI«AlN1lIEEQBE§.'L&QE1llE n S9Ll'lliEM

Most North Amercian Russulas remain inadequately described or are

without descriptions readily available in the North American literature.

Most North American taxa were described in the late 1800°s or early 1900°s

by C. H. Peck, C. H. Kauffman, H. C. Beardslee, G. S. Burlingham, W. A.

Murrill, and R. Singer. These mycologists were largely unaware of the

complexity of the genus, the need for detailed macroscopic descriptions

~accompanied by accurate illustrations, and the usefulness of comparative

micromorphology for consistent delimititation of taxa. Kgsgglg, perhaps

_ more than any other genus of agarics in North America, has suffered from V wholesale misappllcation and overextension of names of European taxa by

both early and modern mycologists.

The first attempt of any comprehensive treatment of North America

Russulas employing modern morphological descriptions was by Singer

(1957). He provided descriptions for about 80 Russulas of North America.

No keys to taxa were included, but the taxa were arranged in a systematic

framework as outlined in other works (Singer 1949, 1962). The

investigator willing to collect the necessary macroscopic data and to

examine the micromorphology of the pileus cuticle, hymenial cystidia,

spore ornamentation etc. could determine many North American taxa with

Chapter 3 76 some degree of certainty. However, Singer°s descriptions were

- unillustrated and often based on examination of limited numbers of fresh

specimens. Futhermore many descriptions were based on populations from

widely separated geographic without indication of which specimens the

descriptions were based on. As a result, Singer°s treatment has been

difficult to utilize and his conclusions on specific taxa must be

interpreted cautieusly.

The first truely usable and methodologically sound treatments of North

American Russulas were those of R. L. Shaffer (1962, 1964, 1970a, 10970b,

1972, 1975). Shaffer provided meticuously detailed descriptions, keys,

and illustrations in monographic form for several subsections of

Kgggglg. Most Russulas encountered in northeastern North Amercia

belonging to subsections treated by Shaffer can now be determined with a

high degree of confidence.

Studies of Russulas in the Southern Appalachians are limited.

Burlingham (1915) mentioned the occurrence of a few taxa in the mountains

of southwestern North Carolina. Beardslee (1918) provided brief

macroscopic descriptions for a few taxa in the vicinity of Asheville,

North Carolina. L. R. Hesler°s series of papers on fleshy fungi of the

Southern Appalachians and his book, Mgggggggg gf ggg Qgggg ßmggggg

(Hesler, 1960), contain brief descripticns and photographs of a few taxa.

Hesler's greatest contribution to the knowledge of the Southern

Appalachian Russulas was his accumulation of collections, notes, and

photographs of specimens from eastern Tennessee and western North

Chapter 3 77 Carolina. Hesler°s attention to recording fresh characters combined with his type concepts of Rggsglg (Hesler, 1960; Hesler, 1961) made his specimens an invaluable resource. He was able to apply names to many of his specimens, often accurately, and these specimens are the basis for the checklist of the Russulas of the Great Smoky Mountains National Park

(GSMP) (Petersen, 1979). Hesler went so far as to compile a key to

Kggsglg based on his field observations anu notes from his type studies.

The inclusion of large numbers of taxa known to Hesler only from dried materlal (especially taxa described by W. A. Murrill) limited the · usefulness of this key.

Singer (1938, 1939) described several Russulas, including some new species, from the GSMNP based on specimens and notes of A. H. Smith. The descriptions were very brief, published in French, and therefore largely ignored by North American mycologists. During the 1940°s, Singer spent two summers at Mt. Lake, Virginia. Descriptions of some of the Russulas collected during these visits were included in his treatment of North

American Russulas (1957). One new species, R. gpgglgghjgggig Singer,

(1948, 1957) was described from Mt. Lake, Virginia.

Finally, in his monographs (1962, 1964, 1970a, 1970b, 1972, 1975),

Shaffer cited several specimens of Russulas as occurring in the Southern

Appalachians based on his visits to the GSMP and material from Hesler°s herbarium.

Chapter 3 78 I present here a sumary of the taxa that I have determined to date

from the higher elevations of the Southern Appalachians. The taxa are

arrranged by the sections of Romagnesi (1967) except for the subsections Aruhaainaa, Amaeninaa, Mudastinaa, and which are either

absent in Europe or which I choose to recognize at a lower taxonomic level

than Romagnesi.

ß QE BSÄÄSLZLA IN IHLE EQRESIS

RUSSULA (Pers.) S. F. Gray

Subsection ARCHAEINAE Heim

RUSSULA EARLEI Peck, Neg Xggk ßggtg Mg;. Bull. No. 67: 24, pl. N.,

figs. 5-10. 1903.

This unusual species occurs in high elevations of the Allegheny

Plateau, the Valley and Ridge province, and the Blue Ridge Mountains.

See Bills and Miller (1984) for a description and discussion.

Section PLORANTINAE Bataille

RUSSULA BREVIPES Peck, Annual Ban- Hau Xszzls §.t.a1;.e Muaeum

43: 20. pl. 2, figs. 5-8. 1890.

Chapter 3 79 Rggggla hggyjpgg and its varieties are common throughout the region.

See Shaffer (1964) for descriptions and discussions.

Section NIGRICANTINAE Bataille

RUSSULA DENSIFOLIA (Secretan) Gillet, Hymgn. p. 231. 1874.

. Romagnesi (1967) indicated the taxon referred to as R. gggsijglig by

American authors may not be the same as Agagiggg ggnsjjglig Secretan and

perhaps should referred to R. aggifglia Romagnesi. Rggsglg Qgggjfglig

and its variants are commonly encountered in the Appalachians. See Shaffer

(1962) for an account of North American populations.

Section INDOLENTINAE Melzer and Zvära

RUSSULA VARIATA Benning, ßg;. Qgz. 6: 166. 1881.

I have never seen this species in spruce—fir forests, but it is

extremely common in various hardwood forest types. See Singer (1957) and

Shaffer (1970a) for descriptions and discussion. Neither Singer (1957),

Hesler (1960), nor Shaffer (1970a) mentioned examining a type specimen

for the name K. yggigtg determined by Mary Benning. During a visit to

NYS in the spring of 1984, I was unable to locate a type specimen. The

original painting of R. ygrigtg by Mary Benning, however, was deposited

at NYS.

· Section VIRESCENTINAE Singer

Chapter 3 80 RUSSULA CRUSTOSTA Peck, Annual Reg. New Xggk ßtgtg Mg;. ·

39: 41. 1886.

To my knowledge, R. grggtggg does not occur in the spruce-fir forest

type, but it is common in various hardwood forests. See Singer (1957)

and Shaffer (1970b) for descriptions and comparison with the closely

related K. yjrgsgggs.

RUSSULA VIRESCENS (Schaffer : Secretan) Fries. Epjggjgjs

Sys;. Myggl p. 355. 1838.

Bgsgglg yjggsgeng appears to be restricted to forests dominated by the

Fagaceae (Romagnesi, 1967). My observations in the Appalachians agree.

See Romagnesi (1967) and Shaffer (1970b) for descriptions and

discussions.

RUSSULA POLYCYSTIS Singer, ßyll. §gg._Mygg1. Zgngg 55: 238. 1939.

7 Prior to this study, Rgggglg gglygygtis had never been reported

occurring beyond the type locality under spruce on Mt. LeConte, Tennessee.

Presently, it is known only from high mountain regions of West Virginia,

Virginia, and Tennessee, but probably occurring throughout Southern

Appalachian Piggg rghggs comunities. It may represent a Buggy}; species

Chapter 3 81 truely endemic to the high•elevatiou forests of the Southern

Appalachians. See Bills (1984) for description and discussion.

Subsection AMOENINAE Singer

RUSSULA MARIAE Peck, Annggl Rgn. Egg Xgrk Sgggg Eng. 24: 74. 1872. Rxmsnla alashuana Murrill. M1¢.o.1szg;La 3¤= 362- 1938- Russia M¤rri11„ Llsmua 6= 217- 1943- Russyla Murrill. Llmia 6= 218- 1943-

ßgggnlg mggigg is the most common and widespread member of the subsection Amggnjngg Singer in eastern North America. A complete description and taxonomic discussion can be found in Bills and Miller

(1984).

It occurs singlely to gregarious on soil or humus in deciduous forests or mixed deciduous-coniferous forests. The author has observed

itfruitingin association with a variety of ectomycorrhizal trees including 2e.¤1.lal.au1;.a„ Eazuazrandiiella, mazmumimu, Issagaaanadmxais, and ornamental Riggg ghigg. Habitat notes with herbarium specimens indicate

frequent associations with Qngggng species. RUSSULA76: ACICULOCYSTIS990. 1984. Kauffman ex Bills & Miller, Eggglggig - Keuffman ex Singer um am-

ßyl], Sg;. Egggj,. Zggngg S5: 243. 1939.

Chapter 3 82 Rgggglg mgrigg Peck, ;gg;g Singer, Sygggig

11: 185. 1957.

This taxon is only known from the Appalachian Mountains of West

Virginia, Virginia, North Carolina and Tennessee and nearby Piedmont regions where it is very common. It fruits single to gregarious on soil or bryophytes, especially in disturbed habitats, e.g. eroded areas, road or stream banks, trails, or surface mines. It has been noted fruiting under Quazmla minus,. Q., s.o.<;.c.in;La and athar app- . Batnln laura. lisnza sanauansls. Blnua uizxiniaua. Biss: ahiaa. and athar Binuu spp- . and Qggyg spp. from June to September. See Bills and Miller (1984) for description and taxonomic discussion.

RUSSULA FLAVIDA Frost an Peck, Annual Kan um Karls Slaßn Mus- V 32: 32. 1880.

Rgggglg mggigg var. flggjgg (Frost in Peck) Singer,

ßgll. Sg;. Myggl. [ggggg 55: 244. 1943.

ßgggglg flggigg fruits singly to gregarious on soil or humus, in forests dominated by Qgggggg spp. such as Q. p;jgg;, Q. ggggiggg, [ggg; xranslif.o.l.La.mu.1.a. l1aunaaanadansla¤nd£lmlaspp~

Known from New York southward to Florida and westward to eastern Texas. lt has been reported from eastern Asia (Chiu, 1945; Hongo, 1960).

Fruiting July to November. See Bills and Miller (1984) for description and discussion. ·

Chapter 3 83 RUSSULA OCHROLEUCOIDES Kauffman, Mggglggig 9: 165. 1917.

— Russula maria: var- snbilmzida Singer. Bull- Sm- Mysel-

{gggg 55: 244. 1939.

Bg;;g1g dggg Burlingham, Mggglggig 16: 19. 1924.

Busäula suhgghrglggga Murrill, Mxgglggig 30: 363. 1938.

Bg;;g1g 1;g1;pg;g Murrill, Llgggjg 6: 212. 1943.

Kg;;g1g 1ggj;g;;ifg;mj; Murrill, Llgygjg 8: 266. 1945.

. Buääulß lßlääääßiifgliß Murrill, Llgggig 8: 267. 1945.

Rg;;g1g gghgglgggggdgg fruits solitary to gregarious on soil or humus

in forests dominated by Qgggggg spp., Qgggg spp., {ggg; ggggdifglig Ehrh., B.a1u:.La, B- laura. Isszgananadsnsia. Assxsasshamm. and

ßhgggggggggg spp. It has been reported from Newfoundland to Florida,

westward to Michigan from June to October. Dr. Richard Homola informs

me that it is fairly common under American beech in Maine. In Virginia

and West Virginia, it is also common under American beech but can also

be found in thin, dry oak-hickory forests. Although this species never

fruited in the quantitative study areas it was found under beech within

20•50 m of plots H5 and H6. See Bills (1984) for description and

discussion.

Subsection MODESTINAE Singer

RUSSULA MODESTA Peck, Mgg Xggk $3;;; Mg;. ßgll. 116: 78. 1907.

Chapter 3 84 Rgggglg mggggtg is one of the most common Russulas in the mixed northern hardwood-red spruce forests of West Virginia and Virginia. It fruits singlely to gregarious on humus under Eggg; gggngifgljg, ßgtglg

gggghgggm and Rhgggggggggn mgxjggm from July to September. Known from

New England southward to mountainous regions of West Virginia, Virginia and Tennessee. This taxon was found in close proximity to all the quantitative hardwood plots but never fruited within their boundaries.

See Bills (1984) for description and discussion.

Section HETEROPHYLLINAE Maire

RUSSULA VESCA Fries, Epiggigig $3;;. Myggl. p. 352, 1838.

Romagnesi (1967) indicated this was perhaps the most common Rgggylg in central Europe. Singer (1957) and Shaffer (1970b) indicated it is common in northeastern North America. QI have observed it fruiting abundantly in yellow birch, beech, and eastern hemlock forests on the

Allegheny Plateau and in the Blue Ridge Mountains.

RUSSULA BRUNNEOLA Burlingham, N. Age;. E1. 9: 233. 1915.

This species has been described in detail by Singer (1957) and Shaffer

(1970). Shaffer also provided excellent illustrations of its important microscopic features. Bgssglg hrgnngglg has been reported from

Washington State and throughout the northeastern United States and

Chapter 3 85 southeastern Canada. This species has now been observed in the higher

elevations of the Southern Appalachian Mountains. I have collected it

in forests of Biggg ggbgns and mixed forests of R. ggbggs, Ahigg .f.r.es.e:.:L, Bssnla and Bamm xzandifalia- It prvbably ¤<=<=¤r¤

throughout these forests types from West Virginia to southern North

Carolina. The southernmost specimen I have seen was collected under Igggg

gggggggsis at Highlands, North Carolina (L. R. Hesler 24698). See Bills

(1984) for a photograph and discussion. -

RUSSULA HETEROPHYLLLA (Fries) Fries, Epigrisis $3;;. Myggl. · p. 352. 1838.

This is a tenative determination of a partially decayed specimen that

fruited one time in plot H4 (see Chap. 1, Table 7). I referred this

specimen to R. hgggggphgllg because of its pale lamellae (a spore print

was unobtainable), green to pale greenish yellow pileus, radially

streaked pileus surface, dermato- and hymenial cystidia with contents

inert in SV, ciliate epicuticular hyphae, and low isolated spore

ornamentation.

Section GRISEINAE J. Schaeffer

RUSSULA REDOLENS Burlingham, Mygglggig 13: 133. pl. 7, fig. 6

. & fig. 6. 1921.

Chapter 3 86 ßyggylg gggglgn; can be found fruiting solitary to gregarious on humus in mesic northern hardwood forests or northern hardwood·red spruce transition zones dominated by Iggy; ggngjfglig, ßgyylg g11gghgyjg¤;j;, Qalemuambra, Iiliaamsriaana. Asarsasshaum. A- humus ggggyjng, and Iggginyg mgglggna from late July until late August.

Presently it is known only from Vermont, eastern West Virginia and southwestern Virginia, but probably more widely distributed within the above forest types in eastern North America. See Bills (1984) for description and discussion.

Section FOETENTINAE Melzer and Zvéra

RUSSULA GRANULATA (Peck) Peck, Annual Kan- Hau Xazk Staia Muaaum

53: 843. pl. C, figs. 1-5. 1901.

Ry;;y1g gggnylgyg is widely distributed in southeastern Canada and northeastern United States and extends southward through the higher - elevations of the Southern Appalachian Mountains. It has been reported from the higher elevations of the GSMNP (Singer, 1957; Shaffer, 1972) and from near the summit of White Top Mt. which is covered by a red spruce forest in southwestern Virginia (Singer, 1957). Ry;;ylg gggyylgyg is the most abundantly fruiting Kugaulg in high·elevation red spruce and northern hardwood forests in southeastern West Virginia occuring with nearly the same frequency in both forests types. In this area, its fruiting season starts in late June and extends into early September with the peak sporocarp production in late July and early August. I have

Chapter 3 87 observed it fruiting abundantly on surface mines reforested with £;ggg

gbjgg. Rnggnlg ggnnnlggg also fruits in cove eastern hemlock forests at

lower elevations (about 850 m). Singer (1957) noted that this species was non-selective in its habitat and fruited in both coniferous and hardwood forests. Personal observations and notes with herbarium

specimens indicate K. gngnglggg may be associated with Eigen gnhgng, B. maziaua, i¤tr¤d¤<=ed 2- ahiaa. Ahias Ialsasaa. A- fxasari, Issxa aanadaxaia. Eamazrandifalia, B- ¤.¤.¤1z1£ar.a, and Osama rslzra-

Shaffer (1972) provided a thorough and detailed description of B. ggnnnlgtg. See Bills (1984) for photograph and discussion.

RUSSULA SUBFOETENS W. G. Smith, lgngn. ßgg., Lgnggn

11: 337. 1873.

Rnggnlg gnbfggtgng is one of the most conspicuous Russulas of the high mountain areas. It is often misdetermined as R. lgngggggggi Melzer or

R. fggtgng (Pers.) Fr. See Romagnesi (1967) and Shaffer (1972) for descriptions and discussion.

Section FELLEINAE Romagnesi

RUSSULA SIMILLIMA Peck, Annngl Ren. New Xg;k Stagg Mg;. 24: 75.

1872.

Chapter 3 88 Kussula .f.e.1.1.e.a subsp• simillima (Peck) Singer, L.1.119.a

22: 707. 1951.

Ry;;y1g ;imi1ljng Peck, fruits in association with Iggy; gggngifglig at high elevations in August and September. It was reported previously in the Southern Appalachians by Singer (1939), Hesler (1949), and and

Shaffer (1970a). Ry;;y1g gimjllimg is almost identical morphologically with.ßy;;y1g_fg11gg (Fries) Fries which is associated with Iggy;_;glgg;jgg in Europe. A critical study is needed to determine the relations of these two Russulas to each other and to the genus Iggy; in the Northern

Hemisphere. E

RUSSULA COMPACTA Frost in Peck, Annygl Rgn. Egg Xggk ßgggg Ey;.

32: 32. 1879.

Ry;;y1g gggpgggg is one of the best known Russulas in eastern North

America. It was well described by Shaffer (1970). It occurs commonly in various hardwood forests types in the Appalachians. On two occasions, on Canaan Mt., Tucker Co., West Virginia, and on Briery Knob, Pocahontas

Co., West Virginia, I observed it fruiting in red spruce or red spruce-yellow birch stands.

I Section ROSEINAE Singer

RUSSULA PECKII Singer, Eygglggig 35: 147. 1943.

Chapter 3 89 This species is widespread in the northeastern United States and

southeastern Canada. It reaches its most southern distribution in the

red spruce-Fraser fir and red spruce-northern hardwood forests of the

Southern Appalachian Mountains. Rgssglg pggkii has been collected in the · higher elevations of the GSMNP near the southern limits of the red

spruce-Fraser fir forests (Hesler, 1945, in part, as K. pggpggjgg Quel.

& Schulz.; Shaffer, 1970). In addition to herbarium material, I have seen

fresh material from the red spruce·Fraser fir zone of Roan Mt., North n Carolina-Tennessee. I agree with Singer°s observations (1943) that it

is constantly associated with conifers, specifically Ahjgg and Rjggg.

Shaffer (1970) provided a detailed description of this species. As he

pointed out, this fungus can be easily recognized by the dry, velvety,

dark red to pink pileus, crenulate lamellar edges, long, clavate, red to

pink, white-based stipe, mild taste and strong red reaction of the

incrusted cuticular hyphae with SV. See Bills (1984) for a discussion

and photograph.

Section EMETICINAE Melzer and Zvara

RUSSULA EMETICA (Schaeffer) Persoon, Qhs. Myggl. 1: 100. 1796.

Buggy]; gmgtjgg is perhaps the most misused epithet in agaricology.

The circumscription of this taxon has been concisely limited by Romagnesi

(1967) and Shaffer (1975). It appears to be rare in the Southern

Appalachians. After four seasons of active searching, it was found once

Chapter 3 90 on a hummock in a bog under red spruce, Dolly Sods, Tucker Co. West

Virginia (GB 837).

RUSSULA SILVICOLA Shaffer, ß91h. Ngyg H9Qg;9g;9 51: 229. 1975.

figs. 13-18.

Shaffer (1975) stated that this is the most common species of the

Eßgßigißßß in eastern North America. Numerous collections of R. 911919919 were made during this study. Its fruits in a variety of habitats ranging from high-elevation spruce-fir forests and reforested surface mines to ”My low-elevation oak-hickory forests. collections agreed well with

Shaffer°s description (1975) and specimens collected and determined by him (MICH).

RUSSULA456.1960.BETULARUM Hora, Igggg. ßgig. My;91. $9;. 43:

Rgggglg 9999199 var. 999919rgm (Hora) Romagnesi, L9;

BQ§§§l§§, p. 401. 1967.

Bills (1984) was the first report of this species in North America where it was found in eastern West Virginia and southwestern Virginia. It is known from birch and birch-spruce forests of western Europe. Most of the collections described were solitary to scattered on humus in forests of

spp. or in stands of ß9;g19 ßllßghßßigßßiß and BhQ§Qd§ßQIQß spp. One of the collections (GB 906) fruited on a reforested surface mine under Zi;99

Chapter 3 91 gpjg;August.and young ßgtnlg gllgghgnjgnsis. Fruiting from late° July to late

RUSSULA KROMBHOLZII Shaffer, Llgydig 33: 82. 1970. .

A Azazism Abbild- Bbssbmlb-

ßghnmmg 9: 6. pl. 64, fig. 5-6. 1845.

Rnsgnlg ggggnngnnggg (Kromb.) Britz., Rg;. Zhl. 54: 99.

1893. non. Peck. Annngl Ren. Neg Xggk §;g;g Mg;. 41: 75. 1888.

Rnggnlg yingggg Burlingham, N. Ang;. E1. 9: 217. 1915.

In my experience, B. krgmbhglzii is one of the most common and most

confusing Russulas in eastern North America. I have recognized at least

three "types" based on fruiting pattern, phenology, and macroscopic

characters. Microscopically, these forms were indistinguishable. But

they all exhibited the essential morphological features described by

Romagnesi (1967) and Shaffer (1975). These forms may represent

"ecotypes", genetically distinct populations adapted to specific

habitats. Further biosystematic studies might be warranted to determine

how distinct genetically these populations are. Below are my general

impressions of these three forms.

Type 1. This form is comon in the vicinity of Blacksburg and in the Blue 1 Ridge Mountains._ Sporocarps characteristically occur in large gregarious

fruitings with as many as 20-40 sporocarps. Fruiting is associated with

Qngggns species in the late spring or early summer (late May to mid-June

Chapter 3 92 in Virginia). Sporocarps range from yellowish green (rarely) to blackish ‘ purple to blackish red. Sporocarps arre often massive with pilei up to

18 cm broad and stipes up to 3-5 cm wide. The stipes range from

non-cinerescent to strongly cinerescent. Hesler (1945; as R. giggggg)

probably described this form.

Type 2. A form that fruits singly or in small gregarious clusters (2-10

sporocarps) occurs at higher elevations. It is ususally associated with

beech or northern red oak during late June to early August. The pileus

is medium to large, dark purple, but often partly or entirely yellowish

green or grayish green. The stipe is strongly cinerescent. Rolf Singer

collected this form at Mt. Lake, Virginia (FH) and determined it as "K.

yjnagga, half-green form". This is the form that occurred in the

quantitative study areas.

Type 3. My impressions of this form are based on three or four

observations in West Virginia and northwestern New Jersey. It appears

to be associated with eastern hemlock, but black birch and northern red

oak were present at some of the collection sites. Further observations

are needed to confirm the habitat. The pileus was consistantly purple,

blackish purple to to dark red, often with yellowish discolorations and

medium to large in size. The stipes were not cinerescent. Most of

Burlingham's specimens of R. yingggg from Vermont and the type locality

on Long Island, New York (NY) appear to be this form.

Chapter 3 93 RUSSULA AQUOSA Leclair, ßgll. Sgg. Myggl. Eggggg

48: 303. pl. 34. 1932.

Rggsglg gggtigg ssp. §g§Q§§ (Leclair) Singer,

Rey. Myggl. 1: 292. 1936.

Buggy}; gggggg was treated in detail by Romagnesi (1967) and Shaffer

(1975). It was encountered sporadically in the margins of bogs fruiting under red spruce, yellow birch, and eastern hemlock and in dense red spruce forests fruiting on byrophytes and/or rotten wood.

_ Section INTEGROIDINAE Romagnesi

RUSSULA CLAROFLAVA Grove, Miglagg Ngtgggljst 11: 265. 1888.

Buggy}; glgggflgyg is commonly found fruiting singly to gregarious on soil, humus or bryophytes in forests dominated by Riggg spp., Ahigg spp.,

ßggglg spp., Rgpglgs spp., and.A1ngg spp. This species has a circumboreal distribution. Populations extend into the higher elevations of the southern Appalachians where they are associated with Rjggg gyhggg, Abjgg

Fruiting from July to October.

This relatively wel1·known species has been previously described in the American literature as R. flgyg Romell. Although this species has been treated in a number of popular works, a complete, modern description, illustrating microscopic features, has not been available in the North

Chapter 3 94 American literature until Bills and Miller (1984) described Southern

Appalachian material.

Section DECOLORANTINAE Maire, sg.

RUSSULA DECOLORANS (Fries) Fries, Eggrjgjg $3;;. Myggl.

p. 361. 1838.

Buggy}; gggglggggg is one of the best known and most easily recognized

Russulas. It occurs commonly in boreal conifer forests of Europe, although its presence in North America is poorly documented. I have collected it at one location under red spruce and yellow birch on Briery

Knob, Pocahontas Co., West Virginia (GB 476). The specimens agreed well with Romagnesi°s description (1967) and specimens collected in Sweden and

Finland. Beardslee°s (1918) report of it fruiting on Mt. Mitchell, North

Carolina is probably accurete. His macroscopic description was consistent with the European concept and his cerrespondence with G. S.

Burlingham indicated that he had learned the species first-hand from L.

Romell during a visit to Sweden.

Section RHODELLINAE Romagnesi

RUSSULA OPERTA Burlingham, Mygglggig 16: 18. 1924. pl.4, fig. 2.

ßgggglg gpggtg is a little known taxon described from Windham Co.,

areas compared ChapterVermont. 3 Specimens from the quantitative study were 95 directly with Bur1ingham°s type specimen (NY) and were found to be identical. Singer (1957) indicated that R. gpggtg is synonymous with R. pggillg Peck and that several morphological forms exist throughout North

America. A systematic revision of this taxon is in preparation and will be published at a later date.

Chapter 3 96 £HAP.'IEB&.. NQIEESMIHIHEEQB£§ISQEI1E

S9LZIHERN

Three taxa of Lggtgrigg of the high-elevation forests of the southern

Appalachian Mountains of North America are redescribed and illustrated based upon new morphological and habitat information. Taxonomic discussions are provided to refine further their circumscriptions and clarify their infrageneric relationships. Among the taxa redescribed are

L. ljggygtgllyg Smith & Hesler and L. ggglgtgs (Peck) Burlingham, two of the most common agarics in red spruce forests of West Virginia and

Virginia. They were determined only with great difficulty because of the abundance of similar taxa within their respective infrageneric groups and lack of comparative, illustrative materials. Lgggggjgg fgggilig

(Burlingham) Hesler & Smith, also redescribed, is a poorly known species with a restricted distribution and may be endemic to the high elevations of the Southern Appalachians. In addition, the taxonomic segregation of

L. gjnggggg Peck var. giggrggs and L. ginggggg Peck var. fgggtgggm Hesler & Smith is reconsidered.

Basidiospore size and shape are from optical sections in side view and exclude the ornamentation. Capitalized color names are from Ridgway

(1912) and those numerically designated, e.g., 6C7-5, are from Kornerup Chapterand Wanscher4 (1978). Spore-print and some lamellar colors are from 97 Romagnesi (1967). Macro- and microchemical tests were made according to

the methods of Romagnesi (1967) and Singer (1975). The letters SV stand

for sulfovanillin, and GSMNP for the Great Smoky Mountains National Park.

All specimens are deposited at VPI unless stated otherwise.

QEIAXA

LACTARIUS LIGNYOTELLUS Smith & Hesler, Brittonia 14: 410.

pl. 22, lower fig. 1962.

FIGS. 14: 3, 15: 5-8

Riley; 2-4.5 cm broad, convex to plane with margin inrolled when young,

becoming plane to umbillicate with margin uplifted, or wavy in age; margin

crenate or not, sometimes eroded or slightly rimose in age; surface dry,

dull, velvety, rugulose or radially rugulose, azonate, black, dark

blackish brown to brown, Clove Brown, Seal Brown, 8F8-7, 7F8-5, Vandyke

Brown, 6F8-6, Mummy Brown, 6E8-7; trama thin, brittle, thickest at disc,

1-2 mm thick at midradius, white to slighty yellow, unchanging or

occasionally staining pale vinaceous pink or Vinaceous-Cinnamon

overnight, sometimes brown around larval channels; odor and taste not

distinctive. Lage; white, milk-like, usually abundant, unchanging.

Lgggllgg adnate to subdecurrent, medium to subdistant, with 2-3 tiers of

lamellulae, sometimes forked, up to 7 mm broad at midradius, acute in _

front, white when very young, soon pale yellow, Cream Color, Cream-Buff,

' pale orange yellow, SA3-2; edges even Chapterfinally Apricot4 Buff or to minutely98

pruinose, pale brown to dark brown, rarely concolorous with lamellar

faces. Sting 3-7 cm tall, 0.4-1 cm wide at midpoint, equal or slightly

flared or fluted at the apex, terete, often curved; surface dry, dull,

velvety, concolorous with pileus or lighter brown, 5D4, white over lower

one-fourth, with white mycelium at base; trama thin.with interior stuffed

or hollow; soft, white to pale yellow, unchanging when cut, or rarely

staining pale vinaceous pink or Pinkish Vinaceous after several hours.

ßggidigspgggg pale yellowish orange (Romagnesi II d-III c) in mass.

7.5-9.5 X 7.5-10.5 um, globose to subglobose; ornamentation amyloid, up

to 2 um high, consisting of large spines, irregular ridges or crests, and verrucae, connected by low irregular ridges and fine lines, with verrucae W often forming short catenulate ridges; suprahilar area usually with a thick, broad, irregular amyloid patch. ßggjgig 55-68 X 10-13.5 um,

clavate, 4-sterigmate. Rlggrgggstidia 5-40 um X 4-6.5 um, filamentous,

flexuous or contorted, occasionally branched or septate, with blunt or

irregularly lobed ends, projecting up to 10 um beyond the basidia, arising

from laticiferous hyphae at various levels of the trama, with or without refractive contents, hyaline to pale yellow in KOH, inert in SV.

Qhgjlggyggjgjg 20-75 X 4-9 um, abundant, filamentous to narrowly clavate,

0-2 septate, arising as branches from inflated basal cells, pale brown to hyaline in KOH. Lamgllgr trama pseudoparenchymatous, with abundant

laticifers; laticifers 6-9.5 um in diam, with yellow refractive or granular contents in KOH. Rjlggg ggtiglg 120-180 um thick, without gelatinous or incrusting materials, a trichoderm, forming a

"virescens-structure", consisting of filamentous to narrowly clavate

Chapter 4 100 5x •„ewaxefn~\„_[ v 6 •¢•»

7 8gg

Figure 15. L. lignyotellus microscopic features°‘ (GB 161).: 5. P5°“’°°"“"“°‘ dermatocystidia, arising from short chains of inflated calls;

dermatocystidia 20-65 X 6-10 um, pala brown in KOH; inflated calls 9-24

X 8-24, globose to cylindrical or irregularly inflated, often pigmented

as dermatocystidia. Zglggg ttggg consisting of sphaarocysts, connactiva

hyphaa and abundant laticifers; hyphaa 2-4.5 um in diam, laticifers 4.5-10

um in diam, with yellow refractive contents in KOH. §tigg_tgtit1g 70-130

um thick, similar to pileus cuticle, a trichoderm of filamentous to

narrowly clavate caulocystidia arising from short chains of inflated

calls; caulocystidia 35-92 X 7-10 um, pala brown in KOH; inflated calls

8-25 X 7-22 um, globose to cylindrical, sometimes pigmentad as

caulocystidia. ßtjgg tggmg composad of nests of sphaarocysts surroundad

by densaly interwoven connactiva hyphaa, connactiva hyphaa 2-5.5 um in

diam, hyaline in KOH, laticifers abundant, same as in pileus trama. Qlggg

gggggtjggg absent from all tissues.

Hgtjt, hghjtgt, ggg gjgtgghgtjgg. Solitary to subgragarious, often A fruiting over extensive areas on naedle litter, humus, deep moss and leafy

liverworts [ßggzggig ttilghgtg (L.) S. F. Gray], or rarely on well-decayed, moss-covered wood. Associated with Rjtgg ggtggg Sarg.,

Apjg; fggggti Poir., or Igggg ggggggggjg (L.) Carr. throughout the

high-elavation forests of North Carolina, Tennessee, Virginia, and West

Virginia. July to September.

Mgtgtjgl ggggjggg. USA: North Carolina: Swain Co.: Nawfound Gap,

GSMNP, L. R. Hesler 11346 - A. H. Smith 7372 (TENN). Tennessee: Sevier

Co.: C1ingman's Dome, GSMNP, L. R. Hesler 20961 (holotype, TENN), L. R.

Chapter 4 102 Hesler 39014 (TENN); Roaring Fork, near Mt. LeConte, GSMP, R. H. Petersen

35126, (TENN); Unicoi Co.: Roan Mt., L. R. Hesler, 27 Sep 1936, L. R.

Hesler, 22 Aug 1937, (both FH). Virginia: Grayson Co.: Mt. Rogers, GB

665; Madison Co.: Limberlost Trail, Shenandoah National Park, V. Cotter

771. West Virginia: Pocahontas Co.: Black Mt., GB 117; Briery Knob, GB

490; Kennison Mt., GB 161; Rocky Knob, GB 387, GB 222; Tucker Co.:

Headwaters of Otter Creek, GB 825.

iv similar tv L· liznxvms Fr•

and L. fgllg; Smith & Hesler with which it could be easily confused.

Macroscopically, L. llgnygggllus is distinguished from L. llggygtgg Fr.

by its smaller stature, absence of lilac or pink staining or only faint

or delayed staining, brown lamellae edges, and paler spore deposit.

Lggggglgg llggygggllgg, L. llgnygggg, L. fgllgg, and the varieties of L.

llgnygggg are so similar that they cannot be distinguished by microscopic

features. All these taxa are usually associated with Rlggg and Ahlgg.

Hesler and Smith (1979) did not consider L. llgnygtgllgg as one of

the varieties of L. llgnygggg because of the apparent absence of lilac or pink staining of damaged tissues. Based upon examination of dozens

of fresh sporocarps over four seasons, damaged tissues of L. llggygggllgg

rarely stain. Occasionally weak pink or vinaceous stains develop in cut

sporocarps after several hours, especially in the stipe bases. In light of the variation in staining reported for the other members of the L. llggygtgg species-complex, the absence of staining or presence of feeble

Chapter 4 103 staining of L. ljgggggellge appears to be insufficient for segregating it from this species-complex.

Another distinguishing feature of L. ljggyggellge is its brown·pigmented cheilocystidia which are manifested macroscopically as brown—margined lamellae. However, brown•margined lamellae have also been described in L. f;11;3, L. ligngggge var. g;nede¤;ie Smith & Hesler, and

L. ljggygggg var. me;gig;;g; (Smith & Hesler) Hesler & Smith. In addition, I have seen brown-pigmented cheilocystidia in some European specimens of L. ljgngggge. The cheilocystidia of these Leggegige species are morphologically and probably developmentally similar to their pileo— and caulocystidia. The development of brown pigments in the cheilocystidia may be under similar regulation as the pigments of the pileus and stipe. In sumary, L. lignyggellge cannot be clearly delimited from the L. ljgngggge species—complex based on morphological features.,

Within their geographical range, however, populations of L. ljggyggellge are relatively uniform, do not exhibit the range of variation reported for the northern populations, and can be recognized as a distinct morphological species.

Most of the taxonomic decisions on the L. lignyggge species·complex in North America appear to have been based upon specimens from outside the range of {ige; gehen; and within the range of {. m;;i;g; Mill., {. glegg; (Moench) Voss, and western {ige; species. Leggegjge ljggyggellge and the L. ljggggggg variants seem to be allopatric populations, A11 specimens determined as L. liggyggge in the Southern Appalachians have

Chapter A 10A either been L. llgyyegellye or other members of the subgenus

Rllyyhegely;. The Leeyegly; featured as L. llgyyeyy; by Hesler (1960,

p. 98) was a taxon within another species group of the subgenus

Rllyghggely;. Systematic sampling of populations of these Lactarii in .

R. gyheye forests in Pennsylvania, the Catskills, and Adirondacks may provide information relating the Southern Appalachian variant, L. llggyeyellye, to the northern populations. Consideration of the present-day distributions of these variants in light of the history of distribution of Rleee and Ahle; in the Northern Hemisphere could lead to the development of hypotheses of how the L. llgyyeyy; species-complex co-evolved with Rleey and Ayle; species.

LACTARIUS OCULATUS (Peck) Burlingham, Bull. Torrey Bot.

Club 34: 89. 1907.

FIGS. 14: 1-2, 16: 9-12

ELeeye;ly; eybgylel; eeyleyye Peck, New York State Mus.

Bull. 67: 37. pl. 83. 1903.

Rlley; 1-4.5(-5.5) cm broad, convex with sharp papilla and incurved margin when young, soon plano-convex or plane with papilla, finally umbillicate with uplifted or wavy margin; margin striate or not, obscurely striate or translucent·striate, sometimes sulcate, crenulate or undulatingg striations extending up to one-third way to disc; cuticle not separable; surface subviscid to lubricous when wet, usually moist to dry, shiny to silky, dull when faded, smooth to minutely rugulose, glabrous,

Chapter 4 105 l hygrophanous, azonate or obscurely zonate when fading, dark reddish brown

to reddish brown, Chestnut, 9F7-5, 8F7-5, 8E8-5, light reddish brown,

8D6·5, brown, with papilla retaining dark colors upon drying, becoming

pinkish brown or light orange brown, Hazel, 7C7-4, from margin inward upon

fading, finally fading to pinkish orange, pale grayish orange, 6B5-4,

Pinkish Buff, Pinkish Vinaceous, 6A3-2, or pink, 7A3; trama pliant to

brittle, concolorous with faded pileus, often water-soaked, thin, up to

1.5 mm thick at midradius, slowly staining pale yellow or not when cut;

odor not distinct; taste slightly bitter, soapy, or not distinctive.

Late; white, milk·like, unchanging or drying pale yellow on lamellae,

staining white paper yellow or not, sparse. Lgmellag adnate, subdecurrent

to decurrent in age, close to medium, with many lamellulae of various

lengths, faintly intervenose or not, acute to subacute in front, 1-5 mm

broad at midradius, pinkish cream when young, soon concolorous with paler — colors of pileus, i 6A5-4; edges even.

ßtigg 1.5-5(-5.5) cm tall, 0.4-0.8 cm wide at midpoint, terete or

occasionally compressed, equal or tapered toward the apex; surface moist,

waxy or dry, longitudinally rugulose, sometimes with a faint canescent

bloom, concolorous with paler colors of the pileus, with pinkish buff to

white hairs over the base; trama thin with hollow interior, brittle, often

water-soaked, slowly pale yellow or not when cut. Qhgmiggl ggggtjggg

(pileus trama): FeS04,-no reaction; gum guaiac—slow1y bluish green.

ßggjgjggpgggg white or pale yellowish white (Romagnesi I a-b) in mass,

8-10(-10.5) X 5.5-7(-7.5) um, obovate to broadly ellipticalg

ornamentation amyloid, up to 1.2 um high, mostly 0.5-1 um high, highly

variable, consisting of spines, irregular, conical to obtuse verrucae,

Chapter 4 106 1 9

‘ 1 · éäagäe äy O

4 z

·— mm „

10pm @ *2 Il

42°¤**· ä Ä

Figure 16. L. oculatus microscopic features GB 510).: 9. Pileus . cuticle, tangential section. 10. Cheilocystidia. 11. Basidiospores. 12. Pleurocystidia.

Chapter 4 107 with verrucae often aligned into short ridges, with few to many isolated

particles, with verrucae and particles partially intercounected by fine

lines or short, low ridges, forming a partial reticulum; suprahilar area

depressed or not, usually unornamented or with fine particles or lines

originating from it. ßggjgjg 33-48 X 9-12 um, clavate to broadly clavate,

4-sterigmate. Rlgggggygtjgig of two types: filamentous type 30-45 X 3-4

um, flexuous, embedded among basidia, arising from subhymenium, without

refractive contents; second type 60-110 X 8.5-14 um, projecting up to 65

um beyond the basidia, aciculate, subulate, obclavate to fusoid, with

acuminate, constricted or appendiculate apices, arising from the

subhymenium, completely or partially filled with hyaline to pale yellow

refractive contents in KOH, purple to purplish black in SV.

Qhgjlggyggjgjg similar to larger pleurocystidia, 30-40 X 6.5-9 um,

subulate, obclavate to fusoid, with tapered or acuminate apices, with or

without contents similar to those of larger pleurocystidia. ßghhymggigm

5-20 um thick, pseudoparenchymatous. Lamgllgg tggmg consisting of

tightly interwoven hyphae, pseudoparenchyma, and laticifers; laticifers

3-10 um in diam, with yellow granular or refractive contents in KOH,

strongly SV+, purple to purplish black in SV. Riley; ggtiglg one-layered,

140-180 um thlck, usually with a thin gelatinous zone at the surface

(observe in Melzer°s), consisting of tightly interwoven hyphae, pseudoparenchyma, and laticifers protruding at various levels, with “ ascending to horizontal free hyphal ends at the surface, with lower levels poorly differentiated from pileus trama; hyphae cylindrical to

irregularly lobed or inflated, septate, up to 20 um in diam, with yellowish brown walls and hyaline contents in KOH. Rilggg tggmg

Chapter 4 108 consisting of nests of sphaerocysts, connective hyphae, and scattered to

abundant laticifers; laticifers 4-15 um in diem, similar to those in

lamellar trama. Stiga gatjtla 20-60 um thick, not gelatinous, consisting

of tightly interwoven hyphae and pseudoparenchyma, with ascending or horizontal free hyphal ends at the surface; hyphae branched or not, septate, with blunt apices, mostly 3-5 um in diam, but with some inflated

cells up to 15 um diam. §§i§§;§{§m§ consisting of nests of sphaerocysts, connective hyphae, and laticifers; laticifers similar to those of

lamellar trama but often larger, 4-21 um in diem. Qlamg tbggattjgga absent from all tissues.

Habit, Habjtat, ang ajatgjbatjbg. Solitary to gregarious on needle

litter, bryophytes, including Sbbaggam spp. and ßaazagaa ttjlgbata, and sometimes on well-decayed wood. Fruiting in association with Ritaa spp.,

Abjaa spp., Riga; attgbaa L., and 2. gaajngaa Ait. Known from Wisconsin across southeastern Canada to New England and southward to Georgia in the

July September. mountains. to J

Hatatial agamigad. CANADA: Ontario: Woods west of South March, J. W.

Groves, 15 Aug 1963, DAOM. 93008 (BPI). Quebec: St. Aubert, J. W. Groves,

1 Sep 1958, DAOM. 59905 (BPI). USA: Georgia: Rabun Co.: L. R. Hesler

22079 (TENN). Michigan: Stinchfield Woods, A. H. Smith, 25 Sep 1961

(TENN). New York: Hamilton Co.: Lake Pleasant, D. B. Young, Aug 1909

(NYS); C. H. Peck, 14 Aug (NYS); North Elba, C. H. Peck Sep, (holotype,

NYS). Tennessee: Sevier Co.: Clingman°s Dome, GSMNP, L. R. Hesler 30326

(TENN). Virginia: Grayson Co.: Cabin Ridge, GB 340; Highland Co.:

Chapter 4 109 Headwaters of Buck Run, GB 516; Smyth Co.: Clinch Mt. Wildlife Management

Area, GB 688. West Virginia: Pocahontas Co.: Black Mt., GB 510, GB 924;

Kennison Mt., GB 497; Rocky Knob, G. Bills, 2 Sep 1982, GB 919; Randolph

Co.: Kumbrabow State Forest, GB 356.

Qhgggygtjggs. This is one of the most common agarics fruiting in ßjggg

gghggs or R. ;g§g3g~Ap1g; fgggggi forests and R. gghggg bogs in West

Virginia and Virginia. The distinguishing features of LL ggglgggg are

its (1) reddish brown pileus that soon fades to vinaceous buff or pale

pinkish buff but retains the dark colors on the papilla; (2) fruiting in

association with Rjggg, Ahigs, and northern Rings species; (3) a slightly

gelatinous pileus cuticle consisting of both interwoven, filamentous and

inflated hyphae, and (4) relatively large, subulate to lanceolate, SV+

pleurocystidia. Microscopic examinations is usually needed to identify

L. ggglgtgg because of the difficulty in detecting the gelatinous cuticle

and its similarity to other taxa in sections (Hesler &

Smith) Hesler & Smith (1979) and Ihgjgggli Hesler & Smith (1979).

Microscopic features are illustrated here to facilitate identification

and for comparison with other taxa in subgenus Rgssglggjg.

LACTARIUS FRAGILIS (Burlingham) Hesler & Smith var. FRAGILIS,

North American Species of Lactarius. p. 503. 1979.

FIGS. 1: 4, 17: 13-15

ELgg;g;jg gggphgggtg subsp. fgggjlig Burlingham, Mem.

Bot. Club 14: 99. 1908.

Chapter 4 110 Riley; 2-6.5 cm broad, convex with papilla and incurved margin when

young, soon plane with papilla, finally umbillicate or depressed with

uplifted or wavy margin; margin not striate, sometimes crenulate or

sulcate; surface dry, rugulose, radially rugulose, or concentrically

rugulose towards the margin, sometimes faintly scurfy or concentrically

areolate, subhygrophanous, dark reddish brown, Chestnut, Tawny, 8D7-5,

8C7-5, orange brown or reddish orange brown, Ochraceous Orange, Tawny

Ochraceous, Cinnamon Rufous, 7F7, 7E7, 6D8-6, 608-6, fading to

Ochraceous-Buff, Orange, SA6-5, or grayish orange; trama brittle, 0.5-2

mm thick at midradius, pale yellowish cream to pinkish buff, unchanging ” when cut, or rarely slightly pinkish violet after several hours; odor

fragrant, of fenugreek (Iriggggllg fggngm;g;agggm L.) or Lgggggigs

ggmphgggggg (Bull. : Fr.) Fr.; taste mild or slightly bitter. Lage;

watery or like dilute milk, unchanging, sparse to abundant. Lgmgllgg

adnate, subdecurrent decurrent, medium to subdistant, with many

lamellulae of various lengths, intervenose, often forked near stipe,

sometimes anastomosing, subacute in front, up to 8 mm broad at midradius,

pale orange, 5A4-3, when young, ochraceous, Orange—Rufous, 5B7, in age,

often with faces dusted white from maturing basidiospores; edges thick,

even. ßtigg 2-7 cm tell, 0.4-1 cm wide at midpoint, terete, or sometimes

irregularly compressed, equal or tapered toward the apex, straight or

curved; surface dry, minutely rugulose, sometimes minutely pruinose,

concolorous with pileus, or with pinkish buff base, with a few pink hairs

at the base; trama hard when young, brittle in age, solid to hollow, with

cortex concolorous with surface and inner trama pale pinkish cream,

Chapter 4 111 unchanging when cut. Qhgmiggl gggggjgng (pileus trama): FeSO4 -pale gray; gum guaiac-slowly dull bluish green. -

ßgsidigspgggg pale yellow (Romagnesi II b-c) in mass, 6.5-8 X 6.5-7.5 um, globose to subglobose; ornamentation amyloid, up to 1.8 um high, consisting of thick ridges and crests, medium to fine lines, sharp to truncate spines, and conical verrucae, with few if any isolated verrucae, forming a nearly complete to often dense reticulum; suprahilar area unornamented or ornamented as the rest of the spore. ßggjgig 38-48 X

9.5-11 um, clavate to broadly clavate, 4•sterigmate. Hymggjgl gyggjgjg

35-50 X 4-5 um, filamentous to narrowly clavate, flexuous, embedded among basidia, arising from hymenium or subhymenium, hyaline to pale yellow in

KOH, inert in SV, without refractive contents. Sgbhymggjgm up to 10 um thick, pseudoparenchymatous, often indistinguishable from lamellar trama.

Lgmgllgg tggmg composed of tightly interwoven hyphae, pseudoparenchyma, and laticifers; cells up to 15 um in diam; laticifers 5-15 um in diam, flexuous, with yellow refractive contents in KOH, greenish yellow in SV.

Rjlggg gggiglg two-layered, 95-145 um thick; epicutis 75-100 um thick, without gelatinous materials, cellular to pseudoparenchymatous, composed of globose, subglobose or irregularly inflated or lobed cells, with cells up to 25 um in diam; subcutis 20-50 um thick, consisting of tightly interwoven, horizontally oriented hyphae; hyphae septate, highly branched, 4-10 um in diam, with greenish brown walls in KOH. Rllggg ggggg composed of nests sphaerocysts, connective hyphae, and abundant laticifers; laticifers S-22 um in diam, similar to laticifers of lamellar trama. §;jgg ggtjglg 25-145 um thlck, not gelatinous, similar to pileus epicutis, consisting of pseudoparenchyma and tightly interwoven hyphae;

Chapter 4 112 13 ¢r . \ Qlllyqgigß' 4r

/' xéähsrgl

10um 15 — -~‘ 14 _ ‘ @ ä

17. L. fragilis (GB @Figure;1£;:|;<1

Chapter 4 113 hyphae filamentous to irregularly lobed or inflated, mostly 3-6 um in

diam, but with inflated cells up to 15 um in diam. Stiga ttama composed

of nests of sphaerocysts, connective hyphae, and scattered to abundant

laticifers; laticifers 8-20 um in diam, similar to those in lamellar

trama. Qlamp gggnagtigga absent in all tissues.

Habit, hapjtat, gg aiattihatjgn. Solitary to gregarious or

subcaespitose on humus or soil. Noted fruiting under [agga gtagajjglja Ehrh- , Bamala Britt mmans. Ass.: aasahamm Marsh- .

and Hagamalia yjtgtajaga L. at elevations above 1035 m (3400 ft). Known

from the Blue Ridge Mountains of western North Carolina, eastern

Tennessee, and southwestern Virginia. August.

Matatjal agagjgag. USA: North Carolina: Transylvania Co.: Pink Beds,

G. S. Burlingham 33-1907 (holotype, NY), D. Guravich 493 (MICH).

Tennessee: Sevier Co.: Ramsey Cascades, GSMNP, L. R. Hesler 35242 (coll.

D. Jenkins) (TENN). Virginia: Grayson Co.: Pine Mt., GB 877; Stone Mt.,

° GB 438.

Qtaatyatjgaa. This is one of the most distinctive taxa of the subgenus

Raaaalagaa. It might be confused with L. gamphgtatta which may fruit in

the same forest at the same time. Lagtarita fragilia can be easily U distinguished from L. aamphgtatga by its ochraceous to light orange

colors, often larger sporocarps, more distant lamellae, and globose to

subglobose basidiospores ornamented with a heavy, complete to nearly I complete reticulum. These distinctive macroscopic or microscopic

Chapter 4 114 features have not been illustrated previously. The two-sterigmate

basidia mentioned by Hesler and Smith (1979) were not observed, The

specimen cited by Hesler and Smith (1979) from Knoxville, Tennessee (L.

R. Hesler 35705) was L. gggphggatgs.

Lgggggjgg fgggjlig var, zghiggs Hesler & Smith, a taxon similar to L.

fgggilig var. fgggjljg, is apparently restricted to the Pacific Northwest

(Hesler and Smith, 1979; A. Methven, pers. comm.), It differs from the

Southern Appalachian populations in sporocarp colors, mycorrhizal hosts

(presumably Qygggyg spp.), and spore ornamentation with finer, more

delicate ridges. These differences coupled with the extreme geographic

isolation of the two taxa suggests L. fgggilig var, gyhiggg should be

recognized at the specific level,

LACTARIUS CINEREUS Peck var. CINEREUS, Annual Rep. New York State

Mus. 24: 73. 1872.

=Lgg;g;jg§ ginggggs Peck var. fgggtgggg Hesler & Smith,

North American Species of Lactarius. p. 396, 1979.

The recognition of L, ginggggg var. faggtgrgg (section Igistgg Hesler

& Smith) is an unnecessary taxonomic distinction, Hesler and Smith (1979) differentiated L. gingrggg var. fgggtggm from the type variety based upon comparison of their specimens with Peck°s (1870, 1884) brief accounts of the taxon rather than on living populations that agreed morphologically with Peck's descriptions. Populations identical with Peck°s descriptions

Chapter 4 115 were never cited by Hesler and Smith (1979). According to Hesler and

Smith, the features delimiting var. faggtgrum from var. gigggggs were

the pale yellow spore deposit rather than.white, olive-gray pileus colors

rather than gray, and longer hymenial cystidia in var. fggeggggm.

Comparison of the holotype of var. faggtgrgm shows it is identical

morphologically with Peck°s type and with specimens of L. gigggggs var. fgggggggm from a wide geographical area. Hymenial cystidia of Peck's type

were up to 72 um long, certainly in the range described for var. fgggtgggm. ‘Whi1e comparing var. gjggrggs to L. yietgs (Fr. : Fr.) Fr.,

Peck noted that in var. gjnggegs the "pellicle is separable". Hesler and

Smith (1979) used this character in the key to the stirps Yigtgs to distinguish var. gingrggs with a "cleanly separable" pileus cuticle from var. IQZSLQIHE which has a pileus cuticle "not cleanly separable". But within the description of var. faggtggym, Hesler and Smith described the pileus as having the "pellicle separable". My observations indicate the pileus cuticle is separable one·third to two·thirds way to the disc. Dark gray, light gray, and olive·gray sporocarps occur in close proximity.

Two distinctive characters often present in L. gjngrggs, not mentioned by Hesler and Smith (1979), are the pale pink cast to the lamellae and the pale pink to pinkish gray zone at the stipe apex.

The distributions of L. giggrgus and the closely related L. hlgggjgg

(Fr. : Fr.) Fr. coincide with the distribution of Zaggs (Neuhoff, 1956;

Hesler and Smith, 1979; Korhonen, 1984). Lggtgrjgs hlgggigg is associated with Egggg sylyggjgg L. in Europe. Lggtggigs gingrggs is known to occur with all ecotypes and varieties of E. gggggjjglig in North America, except

Chapter 4 116 [. gggggjfglig var. mgxigggg (Martinez) Little in Mexico. Chiu (1945)

reported L. gjggggg; under Qggrgg; in southwestern China, although [ggg; species are in the same area. Chiu's observations need to be reconfirmed.

The genus [ggg; probably evolved in and dispersed from the Indo-Malasean region during the upper Cretaceous with subsequent migrations eastward to Europe across Asia and westward to North America via Beringia during the Tertiary (Takhtajan, 1969; Van Steenis, 1972). The most primative

[ggg; species occur in eastern Asia (Takhtajan, 1969). These two

Lggggrig; may have evolved from a common ancestor associated with [ggg; in Asia. Studies of Lgggggig; species associated with the [ggg; species of Eurasia could elucidate a co·evo1utionary pathway of the L. gigg;gg;·L. hlgggigg group with the genus [ggg;.

Mggggjgl ggggjggg USA: Maryland: Montgomery Co.: Cabin John Woods,

V. K. Charles & E. E. Dicks, 23 Sep 1937 (BPI). Michigan: Washtenaw Co.:

Cedar Lake, A. H. Smith 80716 (holotype of L. giggggg; var. fgggggggg,

MICH). New Jersey: Warren Co.: Delaware Water Gap, GB 476. New York:

Rensselaer Co.: Sandlake, C. H. Peck Jul (holotype of L. gigggggg var. gigggggg, NYS). Vermont: Windham Co.: Newfane Hill, V. K. Charles & G.

S. Burlingham, 31 Aug 1939 (BPI). Virginia: Grayson Co.: Stone Mt., GB

436; Pittsylvania Co.: near Danville, OKM 6127, OKM 6128 (coll. J.

Lindsey). West Virginia: Pocahontas Co.: Rocky Knob, GB 418, GB 491.

Chapter 4 117 QHAKIEBL. QEIN1liEILQKE§I§QE IHE '

The distributional patterns of vascular plants of the high elevations

of the Southern Appalachians and their relationship to boreal floras are well known. The floristic and structural similarities of the Southern

Appalachian spruce-fir·birch forests to those in New England and New York have been compared by several investigators (Oosting and Billings, 1951;

Stephenson and Clovis, 1983). Likewise, the affinity of the bryophyte and lichen flora of the high elevations of the Southern Appalachians to boreal regions has been recognized (Anderson, 1971; Dey, 1976, 1978).

However, the fungal flora of these high-elevation communities and boreal communities has never been compared.

Biogeographic studies of many groups of higher fungi are impeded by differences in taxonomic concepts and from the low numbers of taxonomic specialists. However, enough information is now available on the ectomycorrhizal species of the agaric genus Lggtgrigg to compare their distribution in boreal forests with those in the high-elevation forests of the Southern Appalachians. Four seasons of sampling Lgggggjgg species in the high-elevation spruce—fir-birch_forests of Virginia and West

Virginia by the author has provided new information relating the distributions of northern Lggtarigg populations to those of the Southern

Chapter 5 118 Appalachians (Bills, 1985). In addition, monographs of Lggtggggg

(Burlingham,-1908; Neuhoff, 1956; Hesler and Smith; 1979) and regional

treatments of Lggtgrigs species of northern latitudes (Kühner, 1975;

Homola and Czapowskji, 1981; Knudson and Borgen, 1982; Laursen and

Ammirati, 1982; Korhonen, 1984) impart taxonomic consistency and enable

the geographical distributions of many Lggtggigs species to be compared

with the geographical distributions of their potential mycorrhizal hosts.

The Lggtggigg flora of the Southern Appalachians is relatively well

known and is rich in taxa. Approximately one-third of the North American

taxa occur within the Great Smoky Mountains National Park (Petersen,

1979). Hesler's and Smith°s (1979) monograph of North American Lggtggjyg was based largely on material collected by Hesler in the mountains of

Tennessee, North Carolina, and Georgia, and to a lesser degree on

specimens collected and described by Gertrude S. Burlingham from Brevard

and the Pink Beds in southwestern North Carolina. However, Hesler and

Smith (1979) stressed the preliminary nature of their monograph and pointed out that geographic distributions of many species were

incompletely known and that many problematic groups of species still existed.

The main objectives of this study were: (1) to summarize both old and new information on the geographical distributions of Lggtggjgg species in the subalpine spruce-fir-birch forests of the Southern Appalachians and their relationships with higher plant communities; (2) to demonstrate that the Lggtggigg species exhibit geographical distributions similar to

Chapter 5 119 those of the higher plants of the region, and (3) to present hypotheses

that might explain the observed distributions of these Lgggggiug species.

In this study, the Southern Appalachians were defined as the

unglaciated Allegheny and Blue Ridge Mountain Systems south of 40° N

latitude. In eastern West Virginia and western central Virginia, red

spruce (Zjggg ruhen; Sarg.) and occasionally balsam fir (Abjgg hglggmgg

(L.) Mill.) occur as scattered trees to well developed, extensive stands

at elevations >97S m. Red spruce and Fraser fir (Ahjg; fgggggi Poir.)

are usually restricted to elevations >l375 m in southeastern Virginia,

western North Carolina, and eastern Tennessee. The best development of

spruce-fir forests occurs at elevations >1800 m in Virginia, North

- Carolina, and Tennessee. When spruce and fir co-occur, the canopy is

usually dominated by spruce at lower elevations with the percentage of

fir increasing as altitude increases (Oosting and Billings, 1951; l

Whittaker, 1956). ßgtglg gllgghgnignsig Britt. is the most common

co-dominant of spruce and fir in these subalpine forests. Geographically,

these forests appear to be a southern extension of the boreal and

subalpine spruce-fir forests of the Northeast, but because they differ

in floristic composition and climate, they have been regarded as related

but distinct forest types (Whittaker, 1956).

The literature cited above and the collections and notes of Hesler

(TENN) and of the author (VPI) were used to determine geographical

distributions and potential mycorrhizal host relationships. The two most

intensively sampled areas were the main ridge of the Great Smoky Mountains

Chapter 5 120 from Mt. LeConte to C1ingman's Dome, by Hesler (periodically for over 30 years) and the Cranberry Glades area of southeast West Virginia by the author. W

HABJIAIS

ANDHabitatsof Lggtagjgs species depend on the composition of the vascular plant vegetation because they form ectomycorrhizae with members of the

Pinaceae, Fagaceae, Betulaceae, and some Salicaceae (Trappe, 1962;

Giltrap, 1982; Miller, 1983). Native trees of the high elevations available for mycorrhizal formation with Lggtgrigg species include Rjggg nmans, Abisa b.a.1s.ams.a (small p¤p¤1¤ti¤¤¤), A- frasau, Tanga aanadensia (L-) Carr-, Rimlasxxabiu L-, B- laute L-, H- ggpygjjggg Marsh. (small populations), ß. pgpgljjglig Marsh. (small populations), Alggg gygggg (Du Roi) Spreng., Qgggggg alb; L., Q. ggbgg

L., Rggglgg gggggjgggtgtg Michx., and 2. tggmulgiggg Michx. The introduction of Riggg ghigg (L.) Karst., Ring; ggsjgggg Ait., B. gylyggtgig L., and Lg;15 dggiggg Mill. for lumber or reclamation may have introduced exotic Lggtgriug species and provided new habitats for native species. Some Lggtggjgg species are characteristic of Sphagggm bogs

(Hesler and Smith, 1979; Korhonen, 1984). Both Sphggggm glades and bog forests are common in eastern West Virginia but are less common in the high mountain areas to the south (Core, 1966). Other Lggtggigg species exhibit fidelity to arctic or subalpine vegetation (Kühner, 1975; Knudson and Borgen, 1982; Korhonen, 1984). True alpine areas are absent in the

Chapter 5 121 Southern Appalachians, but treeless heath and grass balds resemble

physiognomically tundra or tree lines (Core, 1966), and could permit

arctic species to persist at southern latitudes (Ramseur, 1960; White eb ag., 1984).

The Laebaggea flora (19 species) of the high~mountain areas is closely allied with that of northeastern North America (28 species; Table 20).

The geographic distributions follow the same general patterns as those ‘ of vascular plants, bryophytes, and lichens. Laebaggea distribution patterns were classified as follows: circumboreal (or nearly), extending partially or throughout the high-mountain areas; northeastern North

America (NE), extending partially or throughout the high-mountain areas; widespread in the eastern deciduous forest (ENA); widespread in northern hemisphere, and endemic,

The extension of some boreal Laebaggea species into the spruce·fir-birch zones of West Virginia, and Virginia, but not of North

Carolina or Tennessee, is consistent with the transitional composition of the vascular plant flora of the region (Core, 1966; Stephenson and

Clovis, 1983). Laeeargba gbgmgngaa; occurs in ßeeaga aggegbaggegaga-Alge; ;agg;a bogs in West Virginia and Virginia but rarely under ßeegga in North Carolina (A. Methven, pers. comm,). Laeeaggg; hyeggga; has been found on surface mines reforested with Rgeea abge; in West.£;:.asez.1.·Bet11.laforestsVirginia and in Eisen man-ms·Ab.i.e.s

in West Virginia and Virginia. Large populations of Laeeaggg; hegyg; were found as far south as the spruce-covered mountains surrounding

Chapter S 122 Table 20. 0iaQribuQiun uf Lgiggiyg in burual fur•aQ• and hida-•1•vaQiun fur•aQa uf Qha SuuQh•rn Auuulachiana IS. A. ).

l•) Qr•• Suaeiaa Pruaanua ur 0iaQ1-ibutiun Aaauciahd abaunua l-) ganara in S. A.

ggg l$¢uu. : Fr.) Fr. Ciranburaal Picua. Abiua. Pinua - B•Qu1• mgigyg lFr. a Fr. Ciranburual L. Fr.! · • L. gguggjggg BriQz. Ciranburual B•Qu1a gggqg Fr. Circuvburaal. latula L. l$hcr•d.l - (Fr. Fr. Ciranburaal B•Qu1a L. z Fr.! - • L. lP•ra. : Fr.! Fr. Ciranhurual Puuulua L. Ligggjgg Fr. Circnnburual Picaa. Abiua · ‘! ;p_], • L. Haalar I $niQh NE' cunifara ggg; NE Thuja ? L. saw. - ggg}; R Alnua L. Saiih - NE Alnua L. gjgjgg Pk. - • L, bjpggqgg Pk, ME. Qu HV Picua. Tang • „ L. bgagjgg (Fr. : Fr.) Fr. Ciranburual. Qu NV.VA Picaa. Abiaa. B•Qula • B•Qu1• L. ggiggn (Fr. : Fr.) Fr. Ciraanburual. Qu R • L. llull. : Fr.! Fr. Ciranburaal. Qu R. TN B•Qu1a • L. g_igg (Fr.) Fr. Ciranburual. Qu HV Piuaa. Pinnn ¤=L- amithaa Pk-! • L. Ischuaff. : Fr.) Fr. Ciranburaal. Qu R l•Qu1a • L. (Fr. : Fr.) Fr. Circnnburaal. Qu R Pica:. Pinua • L. gg_gg_i_i Pk. NE. Qu R. TN Picua. Tanaga. Pinua. Faqua. Quran • L. gqgigg Pk. NE. Qu R. TN Picua. Abiaa. Pinua Tang. B•Qula.

L. NE. Qu R. TN Picaa. Abiua. Pinua • PinuaL.L. IPk.lBur1. NE. Qu R. GA Picuu. Abiaa. • Burl. NE. Qu R. TN Tang • L. ggiggg Pk. NE. Qu R. TN Pica. Abiua. Tang.

B•Qu1a. Alnua • L. gggggigg Pk. Hiduaun-und ENAz Picua. Abiaa. Pinua Quran. Butula. Tang • L. ggggü Burl. Miduauruad EMA Pinua. Tauga. Faun. Quran • ' L. ginurg Pk. Hiduauruad EMA Fugua • Q•«••raQ• L. g_=,•_•~;|·;g_gjy; Ißull. : Fr.) Niduauraad Piuaa. Qura.n.

S. F. Gray Fagnn • L. $•niQh I N•a1•r Endunic Qu NV. VA. Picaa. Abiua. Tang R. TN • H•a1•r Fagua. L. jggilig lBn.•r1.l I S•iQh Endunic Qu VA. R. TN Picua. Tang

‘NE · nurQheaaQ•rn MurQh Aulricl *ENA •aaQ•rn NurQh Anurica -

Chapter 5 123 the Canaan Valley in north central West Virginia. Lggtgglgg hlhhggggg

was only found once on a surface mine reforested with £lggg,gblgg in West

Virginia and may have been introduced.

The high•mountain Lggtgglgg flora (19 species) appears to be depleted — compared to the potential Lggtgrigg flora of a boreal spruce•fir-birch

forest (28 species). Several Lggtgrlgg species common in boreal

communities are absent despite the presence of favorable habitats in the

high mountains (Table 20). For this comparison, all the Lggtggjgg species

of both regions were assumed to have been sampled. The depletion in

Lggtgglgg species could be explained by some interrelated hypotheses

originating in island biogeographic theory (MacArthur and Wilson, 1967;

Simberloff, 1974). Mountain peaks, including those of the Southern

Appalachians, have been considered insular islands (Whittaker, 1960;

Vuilleumier, 1970; Johnson, 1975; White gt gl., 1984). The small size

and isolation of the mountain·top spruce·fir-birch communities may have

led to an insular depauperization of their flora. In the Southern

Appalachians, eight of the ten areas higher than 1680 m exhibited a steep

slope for the log vascular plant species·log area curve (White gt gl.,

1984) which is characteristic of sets of insular islands. In addition,

many rare vascular plants of the region exhibited a patchy distribution

among the high peaks. These two lines of evidence indicated the insular

nature of the high peaks. Whittaker (1956) hypothesized that a

post—P1eistocene warming period caused an upward migration of the

Chapter 5 124 spruce-fir·birch zone resulting in extirpation of many boreal species.

Subsequent to a return to modern climate and downward migration of the · e subalpine forests, reimmigration of boreal species between these peaks or from the north has not been possible. Insular depauperization may cause island communities to become overabundant in some species and impoverished in others compared to mainland communities (Simberloff,

1974). Some Lggtgrjgg species may have been replaced by other species of the genus or by other mycorrhizal fungi.

Environmental, climatic, or edaphic factors different from those in the boreal or subalpine forests of the Northeast may have selected for different Lggtgrigg species or in favor of other ectomycorrhizal competitors and may have eliminated some Lagtarigs species in the Southern

Appalachians. Mean and minimal temperatures and annual precipitation tend to be higher in the Southern Appalachians than in the Northeast. A greater percentage of the precipitation occurs as rainfall in the Southern

Appalachian spruce-fir forests. Greater snowfall and longer snowpack may insulate the rhizosphere for a longer time in the Northeast. A significant amount of moisture may accumulate from fog moisture in the montane and coastal spruce-fir forests of eastern North America. Also, soil parent materials and soil forming processes vary among the different regions of the eastern spruce-fir forests.

Some Lggggrigs populations may never have migrated into the Southern

Appalachians or may have continued northward migration beyond the

Appalachians during the Holocene. As the Wisconsin ice sheet retreated,

Chapter 5 125 boreal tree species migrated northward from different population centers

in southern North America (Davis, 1983). Migration rates differed among

trees depending on their individualistic responses to climates, dispersal

rates, and establishment requirements. According to Davis (1983), Lggjg

lggjgjgg (Du Roi) Koch migrated from the Great Plains moving across

Pennsylvania and central New York missing the Southern Appalachians.

Populations of Ahigs hglsgmgg rapidly spread northward from the southeast

along the east side of the Appalachians and then expanded northward and

westward. Rings hggksigng Lamb. and R. ggsingsg moved northward in a

similar pathway but at a much faster rate than other boreal species.

Presumably, Lgggggigs populations associated with these boreal trees

would also migrate northward from different population centers with each

species following different routes at different rates.

Another factor possibly contributing to the depletion of the Lggtggjgg A flora is the reduction in area of the Appalachian spruce•fir·birch forests

during the last century. These forests have been reduced from an

estimated one million acres to about 100,000 acres by logging and fires

.(Korstian, 1937; Allard and Leonard, 1952; Core, 1966). Habitats for

Lgggggigg species have also been eliminated by the destruction of Abjg;

jgggggi by balsam wooly aphids (Agglggs piggg Ratz.) (Fedde, 1973).

Do other ectomycorrhizal fungi of the high-mountains areas follow

these distribution patterns? Few other genera are as taxonomically well

known as Lgggggigs making it difficult to judge. However, most Rgggglg

species known from these spruce-fir-birch forests are also northern in

Chapter 5 126 distribution (Bills and Miller, 1984; Bills, 1984). At lower elevations,

the floristic fidelity of higher fungi with boreal communities is lost

rapidly, often with subtropical, tropical or eastern Asia disjunctions

occurring (Petersen, 1976; Hongo and Yokoyama, 1978). Also the number

of endemic species of Lggtggigg and other ectomycorrhizal fungi increases

at lower elevations (Bas, 1969; Hesler and Smith, 1979; Petersen, 1979).

Other components of the fungal communities within these high-elevation

forests are in need of documentation and comparison with fungal communities of true boreal

forests.Chapter5 127 AERENDJLX A,. LLKZAIIQN QB SBBLEZE M12 PLQ'I§.. $10.,.,. HESI

- 3LI.R§iIN.IA,.

Location of plots 128 JN 'I . . E ' w Ä I /{ ~‘ 4 ,' -· \ \ // // .' E, B ag ,« 1// (N"/ ·/ Y (M 1 x “· p _— ”« ‘— r' «‘ /“ 2/

.‘ _l ¤„ \‘ ,1)r ß 1 aw\·~« sw w F ,_ ‘° 2;+ .· y (— L L X; -— äÜ\‘ „ ~ XQ xx?2 · g'

‘ ‘— _/‘\ ,‘ ~_ ir / / v Z \1 j {5 _· 1 ~' LX \— vz ma we ,~ j ”° {3 '_\‘/,3/ J Il „· J. \\ ·> {I UI ,_ I ·. »— x ·‘ Rg Y ‘_ <' ¤7 1 ·— "« ·Ö » » ,- ~y __,J, /4, J, . ,.4,„ „. / ·' ,1 ‘, °‘ wr L5_./ 4 2;- ,3 ,· '» _/ ,„ XR ig ‘T ’“ K‘—"; Yi ·ä ,' _¥ ‘— il _! rrIIII ni ,‘ r , 1 'f· 1, g . ·' 2;*_‘_ , ' 1 ;_ 'Y ‘~ 5* .‘ -_ Y27 ;¤ I, w „‘ r\¥ Yu &" g? ,' ‘~ // ‘· J, jf L _· ,> {ZA; <; L Y «‘«··t ‘· . -· ·„ .» _‘ r, x

—·Figure18. Locations of plots S1 and S2, Woodrow quadrangle, West Virginia.

Location of plots 129 /“ w -};/9 LB__ 1 ézéigv __ N E S L; __ .· __ _ _' 3

1 __;, -7 *33,

‘~ I ’ 4/ 3 vv { l‘ *_ E y :‘ 1; 3 j :_ ,994 ·/‘- ~ 61 \\3 ‘* L*9

.3 *8 \ ·' /' / / / [ L" *'

1 K L

L[ ‘. 1[2 ‘_ g vi N;

,~Figure19. Locations of plots H3 and H4, Hillsboro quadrangle, West Virginia.

Location of plots 130 A I. Ä ’ (Q ”°’ v ; ·‘.Q .. ‘ 6- \ ~ I 9)* \\ . Ä AÄ I d}I , '~ Iäw ‘ Q \~\ 3/ . IÄJBJX Ä Ä M Gate

‘° ‘ I I Ö dI I { „ ” ‘ " g t d Q { „ . · -1 Q, _° \"‘~ I I Q C -f@<>sI ‘ Ä . — S8 S7

IIf/X

Q I a ‘ I . Ä " " QI Ä};. . A:% — \> . \ L„ jagd 34.E / _ <_ 5,, H . BM /_/‘ Id _ kd d BM d 3479 3794‘/

BM L IF ·, Q J E L E E L - S IQ I: I Q . / I B ” , ",._ Ö f\ Figure 20. Locations of plots S7, S8, H5, and H6, Lobelia quadrangle, West Virginia.

Location of plots 131 I /. II I I \‘ I s\ 60·1I \. 4 mi ~, _/ I- · xx 1 I ·· __;

II

/ lf 1

I ". ‘L \F I Qß /1/1 I /’ 1 'fäaaxao H12 I

\I L\ .1 LF ?“ L' ~ „_ F 1 /

„_ "' /1 ‘_ 1 ,r’ ‘\I r I L ( -¤ \^ I __ FQ I · fl) angry L L \ -1_’„~ Q-xyücp I

°I

IFigure21. Locations of plots S9, S10, H11, and H12, Lobelia quadrangle, West Virginia.

Location of plots 132 HABJZELQQDFrequencyABPLNILIX L. MBS QB MALQK SLRLEZE AND

Maps 133 12345678 4 1¤IllU¤UI UUUCIUED ZIIUUIIII UIIUDIUI 4¤UEIIIID6l¤UUIUUI DIUUIUEUIUIUCCHU 6I¤ IIDIH IIIIUIDD GIUIIIIUU IEIUUIUU YIIIUIIUI IUUIUIDU 8¤I¤UUDU- IUIUUUUU S1 w

UCIIDIDU IIIIUCHI UUEUUUDC IIUCIUII DUUUIDUI IIUDUIUE DUIUIUUI UUUUUIII IIIIDUUU UIDDUIII IIIHDDDD UUUUUUUE IUIIDIUI UUIDUUUU 6 IIIIIIII IUUUIUII S 7 5 3

IIIDIIDU UIIIIIIU IIIIIIDU HUIIIIII HIDIUIII IUIIUIII UIHIUIII UIIIUIUI IIIIIIII IIIIIIII IIIIIUII IIDIUIII IIIUIIIU DUIIIUUI IIIIIDII IIIIIDIU S 9 5 10 Figure 22. Frequency of Lactarius oculatus in spruce plots.

Frequency Maps 134 12345678 1U¤II¤U¤I UIIHIUUE ZDDIUDDUU IIIIUIUU 6IUI¤UIII UIIUIIUC 4I¤I¤IIII IIIICEII 6IIUIIIII IIIIIIII SIIIUUDDD IIDDDIII VIUIIUUUD IIIDUUII SIUUUIUUH IIIUIIUU M M

DDDDDIII JIDDUUUU UIUHUDDI IIIHCIDU DUUUHUII HDIUUCUD UIIIIIID UIHUUDCU IIIIIIII UIDHUUUU IIIIIIII IIDUUUII IIIIIIII IUDDHIII IIIIIIID HDUUCUOD S7 S8

IIIUIIII IIIUIIII IIIIIIII IIIIIIII IIIDUIUH IIIIDIUI UIUIIIUU IUIIIIDI UIIIIIII IDIIIIID UIIIIIII IIIIIIII IIIIIIUI IDIIIIII IHIIIIII IIIIIDII S 9 5 10

Figure 23. Frequency of Clavulina cristata in spruce plots.

FIQQUBIICY MGPS 135 1 2 3 4 5 6 7 8 1IIlIIIII IIIIIIII ZIIIIIIII IIUIIIII slIIIIIII IIIIIIUI 4IIIIIIII IIIIIIII sIIIIIIII IIIIIIII SUIIIIIII IIIIIIII YIIIIIIII IIHIIIII lSIIIIIIII IIIIIIII S1 w

6 IIEIUIII UUIIIIII · IIHUIIII IDIDIIII IUIIUIDI UIUIIIII IIIIIUUI IIIIIIII IIIIUUUI IIIIIUII IIIIDUUI IIIUIIII IIIIIIII IIIDUUEI IIIIIIII IIIIHDIU S 7 5 3

IIIIDDII IIIIIDII IIIDUIDI IIIIIIIE IUIIIUUD UIIIIDUI IIIIUIUI DIIIIDUI IIIIIIIU IUIIIIHI IIIIIIID IIIIUUII IIIIDUUU IIIIIIII IIIIDDUU IIIIIIII 5 9 5 10

Figure 24. Frequency of Lactarius vinaceorufescens in spruce plots.

Frequency Maps 136 12345678 1IDIIIII¤ IIIUIIUI ZIIIIIIII IIIIDIUD 3IIIIIIII IIIIIUID 4IIIIIIII IIIIIIII 6IIIIIIII IIIIIIIU GIIIIIIII IIIIUIII YIIIIIIII IIIIIIII SIIIIIIII IIIIIIII _ M M

HIIIIDUI DDIIIIUD IIIIIIHD UIUIIIUU IDUDIUID UIHIIIUU IIIUIIII UDIUDUUI IIIIIIII ‘UI¤UHIII IIIIIIII UDIIUUIU IIIIIIII UIIIIIIH · IIIIIIII DIIIIIUD S7 1 S8

IIIIIUII IIIIIIII IIUIIIIU IUIIIIII IIIUIIID IIIIHIUI UIIUIIII UIIIIUII DIIIIIID UIIUIIID HIIIIIIU IIIIIIII IUDUIIII IIIIIIIU IIDIIIIU IIIIIIUU S 9 5 1 0 Figure 25. Frequency of Boletus badius in spruce plots.

Frequency Maps 137 12345678 1IIIIIIlI HIUIIIDU ZIIIIIIIU UICIIIII aI¤IIlII¤ IUIUIIII 4IlIIUIII OUUDIIII 6IIIIUIII DDDIIIII GIIIIIIII DUUIIIIU YIIIIIIII UIIIIIIU BUUIIIHII DIIDIIIU S1 w

IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIEI IIIIIIII IIIIIIII IIIIIIII IIIIIIIU IIHIIIII IIIIIIUI IIIIIIII . IIIIIIII IIUUIIII S 7 5 3

IIIIUIDU IIIIIIII IIIDIIDD IIIIIIII IIIIDIII IIIIIIII IIIIIIII IIIIIIII III¤¤¤¤I IIIIIIII 5 IIIUIIII IDDIUIII IIIHDIGI IIIIUIII ¤¤¤II¤¤¤ IIIIIIUI S9 510 g Figure 26. Frequency of Amanita flavaconia in spruce plots.

Frequency Maps 138 12345678 1IIIIIIII IIIIIIII ZUUIUIIII UUUIIIII 6I¤IIIIII IUHUIUII 4I¤EIUIII IHDUDUUI 6UIEI¤III HUIIUIUI 6DIIII¤II IHIIUUII VIIIIIIII IDIIIIII ~ BIIIIIIII ‘ IIIIIIII M M

IIIIUIII IIIIIIID IIIIIIUI IUUIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII . IIIIIIII IIIIIIII IIIIIIII IIIIIIII S7 S8 y

IIIIIIII -IIIIIII 9 UIIIIIUU IIIIIIII UUIIIIII IIIIIIII UIIIIIII IIIIIIII ·IIIII¤I¤ IUIIIIII IIIIIIII IIIIUIII IIUUCUDI IIIIIIIU IIUIIUII IIIIIIHI $9 510

Figure 27. Frequency of Lactarius lignyotellus in spruce plots.

Frequency Maps 139 12345678 2 1¤IIIIIII IIIIIIII ZIDDIIIII IIIIEIII 6IIIIIIII IIIIUIII 4IIUIIIII IIIIUHII 6IIIIIIII IIIIHIIU 6IIIIIIII IIIIIIUI . YIIIIIIII IIIIIIII 8IIII¤III IIIIIIII S1 M

IIUIIHII IIIIIIIU IIIIIIDI IIIIIIII 5 UIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII. IIUIIIII IIIIHIII IIUIIIII .IIIIIIII IIIIIIII IIIIIIII II-IIIII S7 _ sa

IIIHIIII IIIIIIII IIIIIIII IUDIIIII IIIHHUII IIIIIIII IIIIIUII IIIIIIII IIIIUUUH IIIIIIII IIDUIUII IIIUIIII IIUIIIII IIIIIIII IIIIUIII IIIUIIII S 9 5 10

Figure 28. Frequency of Inocybe umbrina in spruce plots.

Frequency Maps 140 1 2 3 4 5 6 7 3 1¤IIIIIIl IIIUUIII ZUUIIIIII IIIUIIII sIIIIIIII IIIIIIII 4IIIEIIII IIIIIIII sIIIIIIII IIIIIIII GIICDUIII IIIIIIII VIIUIIIII IIIIIIII 8IIUIIIII IIIIIIII S1 w

IIIIIIII IIIIIIID IIUIIIII IIIIIIIU IIIIIIII IIIIIIID IIIIIIII IIIIIIII IIIIIIII IIIIIIID IIIIIIII IIIIIIDU IIIIIIUC IIIIIIII UUIIIIIU IIIIDUHI · S7 ss

IIUIIIUI IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII UIIIIIII IIIIIIII IIIIIIII ss 510 Figure 29. Frequency of Russula granulata in spruce plots.

FIGQIJBIICY MBPS 14], ’ 12345678 · 1II¤¤l¤ID IIIIIIII ZIIUUIIII IIIIIIII 6IIIIIIII IIIIIIII 4IIIIIIII IIIIIIII 6IIIIUIUI IIIIIIII GIIIIIIII IIIIIIII YIIIIIIUU IIIIIIII BIIIDIDUU IIIIIIII S1 w

IIIIIIII IIIIIIDI UIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII S7 S8

7 IIIIIIII IIIIIIII _ IIIUIIII IIIIIIIU IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIHIIIII IIIIIDUI IIIIIIII IIIIIIII IIIIIIII S 9 5 1 0 Figure 30. Frequency of Amanita inaurata in spruce plots.

Frequency Maps 1}*2 3 12345678 2 1lIIIlIII IIIIIIII ZIIIIIIII IIIIIIII 6IIIIIIII IIIIIIII 4IIIIIIII IIIIIIII 6IIIIIIII IIIIIIII GIIIIIIII IIIIIIII YIIIIIIII IIIIIIII . 8IIIIIIII IIIIIIII S1 w

IIIIIIDD IIIDIIII IIIIIIIU IIIIIIII IUIIIIII IIIIIIII IHIIDIUD IDIIUIII UIIIIIDI IIIIIIII IIIIDIII IIIIIIII IIIIIIII IIIIIIHI IIIIIIII IIIIIIII ‘ S 7 5 3

IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII UIIIIIII IIIIIIII ¤¤¤IIIII IIIIIIII S 9 5 10_

Figure 31. Frequency of Lactarus camphoratus in spruce plots.

Frequency Maps 143 123456784 ‘IIII¤III UDIIIIII g ZIIDIIDII UUIIIIII a¤IDIIIII IIIIIIII 4II¤UIIII IIIIIIII sIIIIIIII IIIIIIII SIIIIIIII IIIIIIII VIIIIIIII IUIIIIII SIIIIIIII IIIIIIII g H3 H4

IIIIUUII IIDIIIII IIDIIIII IIIIUIII IIIDIIII IDIIIIII ‘ IIIIIIII IIIIIIII IIUIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII 6 H5 H6

UUIIIIII IIIIIIII ·IIIIIIII IIIIIIUI° IIIIIIII IIIIIIII IIIIIIII IIIIIIII g IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII H11 H12 Figure 32. Frequency of Lactarus camphoratus in hardwood plots.

V Frequency Maps 144 12345678 ‘III¤¤III IDIIIIII ZIIIUIIII IUIIIIII 3IIIIIIII IDIIEIII 4IIIIIIII IIUIDIII 6IIIIIIII IIIIIIII 6IIIIIIII IIIIIIII VIIIIIIII IIIIIIII 8IIIIIIII F IIIIIIII H3 H4

DIIIIUII IIIIIIII IUIIIUII IIIIIIII IIIIIIII DUIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIUI IIIIIIII IIIIIIII IIIIIIII H5 H6

IHIIHCII IUIIIIII IIDIIIII IIUIIIII IIIIIIII IIIIIIII IIIIIIII IUUIIIII IIIIIIII IIIIIIII IIIIIIII IIIIUIII IIIIIIII IIIIUIII IIIIIIII IIIIIIII H11 H12

Figure 33. Frequency of Russula granulata in hardweod plots.

Frequency Maps 145 1 2 3 4 5 6 7 8 ‘IIIIIIII IIIIIIII ZIIIIIIII IIIIIIII 3lIIIIIII IIIIIIII 4IIIIIIII IIIIIIII 6IIIIIIII IIIIIIII GIIIIIIII IIIIIIII 7IIIIIIIl IIIIIIII SIIIIIIII IIIIIIII H5 H4

IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIDIIIIU IIIIIIII DUDUIIUU IIIIIIII HUUDIIDD IIIIIIII UUUDIIII IIIIIIII UUUUDIII H5 H6

IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII H11 H12

Figure 34. Frequency of Boletinellus merulioides in hardwood plots.

Frequency Maps 146 — 1 2 3 4 5 6 7 8 ‘IIIIIIII IIIDUIID ZIIIIIIII IIIUIIIU SIIIIIIII IIIIUUII 4IIIIIIII IIIDIIUD sIIIIIIII UIUDIIIU SIIIIIIII IIIDUUUU 7IIIIIIII IIIHIIII 8IIIIIIII IIIIIIII H3 7 H4

IIIIIIII UIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII 1 H5 H6 _

IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII H11 H12

Figure 35. Frequency of Scleroderma citrinum in hardwocd plots.

Frequency Maps 147 4 1 2 3 4 5 6 7 8 ‘IIIIIIII IIIIIIII ZIIIIIIII IIIIIIID SIIIIIIII IIIIIIII 4IIIlIIII DUIIIIII 6II¤IIIII IIIIIIII 6IIIIIIII IIIIIDII 7IIIIlIII IIIIIIII 8IIIIIIII IIIIIIII H3 H4

IIIDIIII DUIIIIII IIIIIIII UUIIIIII IIIIIIII IIIIIIII IIIIIIII UIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII H5 H6

IIIIIIII IIIIIIUI . IIUIIIII IIIIIIII _ IIIIIIII IIIIIIII .IIIIIIII IIIIDDII IIIIIIII IDUIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII IIIIIIII H11 H12 Figure 36. Frequency of Laccaria laccata in hardwood plots.

Frequency MBPS 148 AREEND.IXQ..IBEELz2£MDBHlLQQAIIQN§INlL§RDW£lQDANDSRRLLQERLQIS.„

Abbrev1ac1o¤s for mess.

P=Bis;s.ams.bens S=Ass1s.a.mh¤mm l F=£¤xusxxuxsii£9.11¤ C=P.¤ms;s.s.e:9.tina Q=Q1ez.c1mr1;1zz.¤ A=Eu:sinuaameri:„ana

Tree locations 149 1 2 3 A 6 6 1 g ‘IIIIIIIl =lIIIIIII =lIIIIIII ·IIlIIlII =IIIIIIIIP

P P PIIIIIIII ·IlIIIlIl S1

P Figure 37. Tree locations in plot S1.

Tree locations 150 • 1 2 2 4 6 6 1 ·lIIIIIll =IIIIIIlI =IllIlIIl

·IlIllIIIP ·IIlIIlI* ·IlIIIHlI 7IlIlIIIl ·IlIlIIll S 2

Figure 38. Tree locations in plot S2.

Tree locations 151 1 2 3 4 S 5 7 I GIIIIIIIIG IHIIIIII G IIIIIIII G IIIIIIII IIIIIIIIGG IIHIIIII G G IIIIIII G G IIIIIII H3

Figure 39. Tree locations in plot H3.

Tree locations 152 1 2 3 I 5 O 7 Ü _ ·IlIIlIIl ·IllIIIII ·IllIIllI ·IllIIIIl SII-II--I ·IlIIIlII «IIllIlII ·IIIllIlI H 4

Figure 40. Tree locations in plot H4.

TIQG 1OC8tiOIlS 153 • • 1 2 s e s 7 IIIIIIIH · IEIIIIII 7 IIIIIIII · IIIIIIIÜ 7 IIIIIIII 7 IIIIIIII 7 IIIIIIII C 8 H5

Figure 41. Tree locations in plot HS.

Tree locations 154 ’

1 2 3 4 5 s 7 | *IlIIlIII =lIIIIIII =IIlIIIII ·IIIIIIII 5 I°IIIllI ~llIIIlII »IIIIIlIl •I-I-IIII H 6 '

Figure 42. Tree locations in plot H6.

Tree locations 155 • A 1 2 3 6 6 7 6 ·lIllIlIl ·I.-II-.- ·IlIIIIII ·IllIIlII SHIIIIIII «lIIIIIIl 7IIIIIIII ·IIIIIIll · S 7

Figure A3. Tree locations in plot S7.

TISB 10C&tiOI1S 156 1 2 3 4 s 6 7 ' 5IIIIIIII5 ÜIIIIIII 1 5 IIIIIIII · IIIIIIII 5 IIIIIIII 5 Ill-IIII 5 IIIIIIII 5 IIIIIIII S 8

4 Figure 44. Tree locations in plot S8.

l Tree locations 157 1 2 3 4 5 ( 7 g G IIIIIIII G IIIIIIII G IIIIIIII · IIIIIIII G IIIIIIII G IIIIIIII G IIIIIIII G IIIIIIII S 9

Figure 45. Tree locations in plot S9.

Tree locations 158 1 2 I 4 I I 7 I ‘lIllIIII ·IlIIIlII ·lIIIIIII ·IlIIIIII ·IlIIIIlI ·IlIIlIIl 7IIIIlIIl ·IIIIIIIl S10

Figure 46. Tree locations in plot S10.

Tree locations 159 ‘ 2 s 4 6 6 1 g *IIlIlIlI ·IlIIIIIl~IlIIIIII ·IllIIIII 5

F ·IIIIlIlIC 7 F ·IIlIllII H 1 1

Figure /+7. Tree locations in plot H11.

Tree locations 160 1 2 3 4 5 6 7 3

VIIIIII C I VIIIIIIII VIIIIIIII B 4 F VIIIIIIII VIIIIIIII VIII C IIII VIIIIIIIIH12

Figure 48. Tree locations in plot H12.

TIQQ 1OC8tiOIlS 161 AREENDIX IL. BAK DAIA l2§.1i.1„.

Copies of the raw data sets for fungal species and tree species are obtainable from the author, Dr. Orson K. Miller, Jr. , and Dr. Golde I.

Holtzman. An additional set was deposited in the New York Botanical

Garden Archives.

Appendix D. Raw data 1981-83. 162 B ibliography163

I

__l____l_. 1939. Two new species of Rugsggle together with the spore ornamentation of some of our American Russulas. Myeglegie 31: 490-498.

Cain, S. A. 1938. The species-area curve. Age;. Mid. Net. 19: 573-581.

Campbell, C. L. and S. P. Pennypacker. 1980. Distribution of hypocotyl rct caused in snapbean by Bhizettenie selmi- 70: 521-525.

Christensen, M. 1981. Species diversity and dominance in fungal communities. ln: Ihe fgngel eegggnjty. Eds D. T. Wicklow and G. C. Carroll. Marcell Dekker, Inc., New York. pp. 201-232.

Chui, W. F. 1945. The Russulaceae of Yunnan. Lleygie 8: 31-59.

Cliff, A. D. and J. K. Ord. 1973. Sgetjgl egtgee;;elet1g¤. Pion, London. · 175 p.

Cole, L. C. 1949. The measurement of interspecific association. Eeglegy 30: 411-424.

Core, E. L. 1966. Yegetetjen gf lee; Y1;g1gie. McClain Printing Co., Parsons. 217 p.

Cotter, H. V. T. and G. F. Bills. in press. Comparison of the spatial patterns of the sexual and vegetative states of ßeletjgellge

Davis, M. B. 1983. Holocene vegetational history of the eastern United States- Pp- 166-181- In:. ef the united Staus,. hl,. 2... Ihe Heletene- Ed- . H- C- Wright- University of Minnesota, Minneapolis.

Deacon, J. W., Donaldson, S. J., and F. T. Last. 1983. Sequences and interactions of mycorrhizal fungi on birch. Ble; gg ßeil. 71:257-262.

Dey, J. P. 1976. Phytogeographic relationships of the fructicose and foliose lichens of the southern Appalachian Mountains. Pp. 398-416. In: hiuerx ef the hieta ef the sedthern hrt lu- Algae end thai- Eds-. B- 0- Parker and M- K. Roane. University of Virginia, Charlottesville.

. 1978. Fructicose and foliose lichens of the high-mountain areas of the southern Appalachians. ß;gglegjet 81: 1-93.

Dominik, T. 1961. Studium nad mikotrofizmem swierka pospolitego - hp1.Pice excelsa (Lam.) Lk. w Polsce. lnete Bee, Leege £;„ 1961, 209: 1-24.

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Edens, D. L. 1973. The ecology and succession of Cranberry Glades, West Virginia. Ph. D. Dissertation, North Carolina State University, Raleigh.

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Fedde, G. F. 1973. Impact of the balsam wooly aphid (Homoptera: Phylloxeridae) on cones and seed produced by infested Fraser fir. Qgygg. Eyggmg}. 105: 673-680.

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. 1981. Quantification of sporocarps produced by hypogeous fungi. }y Ihg fgygg} gggmgyigy. Ed., D and G. C. Carroll. Marcell - Dekker, Inc., New York. pp. 553-568.

Gauch, H. G. 1982. Multivariate analysis in community ecology. Cambridge University Press, New York. —

Giltrap, N. J. 1982. Production of polyphenol oxidase by ectomycorrhizal fungi with special reference to Lgggggig; spp. Iggy;. Egig. Mygg}. Egg. 78: 75-81.

Greig-Smith, P. 1983. Quantitative plant ecology. 3rd. Ed. University of California Press. Los Angeles. 359p. ~

Hering, T. F. 1966. The terricolous higher fungi of four Lake District woodlands. Iggy;. Eg. Mygg}. Egg. 49: 369-383.

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. 1945b. Notes on Southern Appalachian fungi. VII. J. Igyyg;;gg. Aggg. Eg}. 20: 363-372.

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______. 1960. A study of Eg;;g}g types. Mgm. Iggggy Egg. Q}gg 21: 1-59.

. 1967. The genus Eygg}ggg in southeastern North America. B.e.ih- Nam Haduidaia 23= 1·196- and A- H- Smith- 1979- Nnnth Anuisau sp.¢.¢.i¤ 9.£ Lgggggig;. University of Michigan Press, Ann Arbor. 650 p.

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_ Hora, F. B. 1959. Quantitative experiments on toadstool production in woods. Igggg. E;. Myggl. ßgg. 42: 1-15. ______. 1960. New checklist of British agarics and boleti part IV. Validations, new species and critical notes.I;ggg. ßgjg. Myggl. §gg. 43: 440-459.

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Kauffman, C. H. 1917. Tennessee and Kentucky fungi. Mggglggjg 9: 159-165. _....l- 1918- Ih; Aza:.is.¤:&.¤s Qi Mishizan- V¤1s- I. II- Mishizan Gm.1- Biel- $1:;:- Pxbl- 26. hiszl- Sg: 5. 924 p-

Knudson, H., and T. Borgen. 1982. Russulaceae of Greenland. Pp. 216-243. lg: Agggjg ggg gjgjgg gygglggy. Eds., G. A. Laursen and J. A. Ammirati. University of Washington, Seattle.

Korhonen, M. 1984. Sgggggg gggkgg. Otava, Helsinki. 223 p.

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