Ecology and Management of Agriculture Morels Harvested From the Forests ... and Management of Morels...

170
Ecology and Management of Morels Harvested From the Forests of Western North America David Pilz, Rebecca McLain, Susan Alexander, Luis Villarreal-Ruiz, Shannon Berch, Tricia L. Wurtz, Catherine G. Parks, Erika McFarlane, Blaze Baker, Randy Molina, and Jane E. Smith United States Department of Agriculture Forest Service Research Station General Technical Report

Transcript of Ecology and Management of Agriculture Morels Harvested From the Forests ... and Management of Morels...

Ecology and Management of

Morels Harvested From the Forests

of Western North America

David Pilz, Rebecca McLain, Susan Alexander, Luis Villarreal-Ruiz,

Shannon Berch, Tricia L. Wurtz, Catherine G. Parks, Erika McFarlane,

Blaze Baker, Randy Molina, and Jane E. Smith

United States

Department of

Agriculture

Forest Service

Research Station

General Technical

Report

Authors

David Pilz

Rebecca McLain

Susan Alexander

Luis Villarreal-Ruiz

Shannon Berch

Tricia L. Wurtz

Catherine G. Parks is

Erika McFarlane

Blaze Baker

Randy Molina

Jane E. Smith

Cover Art

Abstract

Pilz, David; McLain, Rebecca; Alexander, Susan; Villarreal-Ruiz, Luis;

Berch, Shannon; Wurtz, Tricia L.; Parks, Catherine G.; McFarlane, Erika;

Baker, Blaze; Molina, Randy; Smith, Jane E. 2007.

Morchella

Summary

Morchella.

Contents 1 Introduction

5 About Morels

Morel Harvesters

75 Morel Commerce

88 Regional Summaries

108 Management and Research

121 Closing Remarks

124 Glossary

128 Acknowledgments

129 Metric Equivalents

129 Literature Cited

158

Ecology and Management of Morels Harvested From the Forests of Western North America

Introduction

Tricholoma magnivelare

Cantharellus Morchella

Morels are more

valuable than

chanterelles and

more abundant than

American matsutake;

hence in combination,

the total value of morel

Northwest is larger

than either of the

others.

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2

Ecology and Management of Morels Harvested From the Forests of Western North America

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1

“Morchella M

1

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Table 1—Names used in this publication for distinctive western morels occurring in western

North America north of Mexico

Our Morelusagea Disturbanceb cladec d Comments

M. elata

M. conica

M. angusticepse M. conica

M. angusticepse M. conica

M. angusticepsf M. atrotomentosa g

M. esculenta

M. crassipes

M. crassipes

M. esculenta

Morchella

rufobrunnea

M. deliciosa

M. deliciosa

Morchella semilibera

Mitrophora semilibera

a

b

c

M. rufobrunnea, M. guatemalensis, M. rigidoides

d

Morchella rufobrunnea Morchella semilibera

e

f

g “M. atrotomentosa

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5

About Morels

Whence Morels

Webster’s Third New

International Dictionary

2

2

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Table 2—Some non-English common names for morels

a Country Referenceb

M. conica

M. semilibera

Morchella

M. angusticeps,

M. esculenta, M. conica

Morchella

Morchella

M. crassipes

Morchella

Morchella

M. esculenta, M. conica

Morchella

M. angusticeps

Morchella

M. elata

Morchella

Morchella c

M. semiliber

M. semilibera

M. elata

Morchella

M. conica

Morchella

M. semilibera

M. elata, M. conica

M. semilibera

M. deliciosa

Morchella

M. costata

M. angusticeps, M. esculenta,

M. conica

Morchella

Morchella

Morchella

Ecology and Management of Morels Harvested From the Forests of Western North America

Table 2—Some non-English common names for morels (continued)

a Country Referenceb

Morchella

Morchella

Morchella

M. conica, M. esculenta

Morchella

M. elata

M. esculenta

M. semilibera

Morchella

Morchella

Morchella

M. conica, M. esculenta

Morchella

Morchella

M. esculenta

M. elata, intermedia, d

conica

Morchella

M. esculenta

M. esculenta

Morchella

Morchella

M. esculenta

M. semilibera

M. elata

Morchella

M. esculenta

Morchella

M. esculenta

M. semilibera

M. esculenta

M. elata

M. conica

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3

4

Distribution

3

4

Table 2—Some non-English common names for morels (continued)

a Country Referenceb

Morchella

Morchella

Morchella

Morchella

Morchella

M. conica, M. elata

M. esculenta

Morchella

a

b

c

d

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5

Eucalyptus globus Pinus radiata

6

Nothofagus

Castanea sativa Morchella conica

7

Austro-

cedrus chilensis 8

5

6

7

Castanea sativa Tuber aestivum

Morchella conica

8

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Habitats

Disturbance—

Range of habitats—Arora joked that

morels “usually grow

outdoors.”

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Trees—

9

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M. elata

Timing

Geopyxis carbonaria

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Taxonomy

Kindred organisms—

Morchella

Morchella

Morels fruit when

winter snow has

melted, the soil is

beginning to warm,

and the air is still

humid.

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Morchella Verpa Gyromitra

Morchella

Verpa

Gyromitra

Morel species—Calling morel taxonomy

“problematic” is an

understatement.

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10

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Morchella Verpa

Morchella

Ecology and Management of Morels Harvested From the Forests of Western North America

M. rufobrunnea .

M crassipes

Species Descriptions

Most morels in

North America lack

meet the standards

of the International

Code of Botanical

Nomenclature.

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The Known Morels in Mexico, a Description of a New Blush-

ing Species, and New Data on

Head:

Ribs:

Pits:

Stalk:

Spore size:

11 12

11

12

Ecology and Management of Morels Harvested From the Forests of Western North America

M. elata-M. angus-

ticeps-M. conica

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Pink morel

Head:

Ribs:

Pits

Stalk:

Spore size:

Green morel

Head:

Ribs:

Pits:

Stalk:

Spore size:

Morchella elata M conica M angusticeps

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Gray morels

Head:

Ribs:

Pits:

Stalk:

13

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Spore size:

Morchella atrotomentosa

M esculenta atrotomentosa as a “nov var ad sic int

M esculenta

Mountain blond morel

Head:

Ribs:

Pits:

Stalk:

Spore size:

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M esculenta

M esculenta

M esculenta

M esculenta

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Yellow morel

Head:

Ribs:

Pits:

Stalk:

Spore size:

Morchella

rufobrunnea

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Red-brown blushing morel—Morchella rufobrunnea

Head: Ribs:

Pits:

Stalk:

Spore size:

Clethra

M

deliciosa M. rufobrunnea M. rufo-

brunnea

M esculenta M rufobrunnea

M guatemalensis M herediana

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Half-free morel Morchella semilibera

Head:

Ribs:

Pits: Stalk:

Spore size:

M semilibera

M semilibera

Verpa bohemica

Verpa

Ecology and Management of Morels Harvested From the Forests of Western North America

Morchella semilibera

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Potentially harmful look-alikes—

Verpa Gyromitra

Verpa bohemica

Morchella Verpa Gyromitra

Helvella Disciotis

Discina

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Morchella, Gyromitra, Verpa. Gyromitra Verpa

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Biology

Overview—

14

14

To understand

how morels seem

unique among

edible forest fungi

in their adaptations,

it is necessary to

understand their

life cycle, modes

of nutrition, and

reproductive strategies.

Ecology and Management of Morels Harvested From the Forests of Western North America

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Tuber

Morchella

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Morels might be

capable of the highly

unusual feat of haploid

meiosis.

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Mycelia—

Costantinella cristata

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15

15

“individual” and its

spatial extent in the

soil is problematic.

Ecology and Management of Morels Harvested From the Forests of Western North America

Costantinella cristata

Morchella

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Mycorrhizae—

16

17 M. rotunda M esculenta M elata

16

17

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M esculenta

18

M

elata

M esculenta M elata

Morchella

Muffs—

18

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Alnus glutinosa Rubus tomentosus

Robinia pseudoacacia

Sclerotia—

Sclerotina sclerotiorum

Ecology and Management of Morels Harvested From the Forests of Western North America

Fruiting—

Factors that facilitate fruiting—

Morchella

Such fruiting may

represent a last-ditch

effort to reproduce

when the morel’s food

source disappears.

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Geopyxis carbonaria

G. carbonaria

M elata

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19

19

decomposable organic

matter could come from

forest soil organisms

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M elata

Factors that trigger fruiting—

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Morchella rotunda

20

21

Factors that support fruiting—

Fruiting body—

Morchella

20

21 a

a

a

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22

M delicious M esculenta

22

When conditions

are optimal, these

primordia can grow,

in 1 day, into morel

fruiting bodies that are

size.

Ecology and Management of Morels Harvested From the Forests of Western North America

M

conica M esculenta

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Aradus debilis Aradus

Spores—

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Population genetics—

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Inbreeding and haploid meiosis

Ecology and Management of Morels Harvested From the Forests of Western North America

Podospora anserina

Reproductive strategies—

Whether any species of

morel actually conducts

haploid meiosis in

nature is not known,

but it might be a useful

strategy for colonizing

new habitats.

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morels in western

forests remain

somewhat of an

ecological mystery.

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Cultivation

Context—

Agaricus Lentinula edodes Pleurotus

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History—

M esculenta”

Morchella rufobrunnea

Morchella

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Agrocybe aegerita Pholiota nameko

Constraints and opportunities—Morels have a

complex life cycle

that complicates the

process of scaling up

cultivation methods to

procedures.

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23

23

Morchella

Morchella

Ecology and Management of Morels Harvested From the Forests of Western North America

Prospects—

M

hortensis

Morchella

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The Handbook of Fungal Technology

Morchella

Collecting to Cooking

Harvesting—

Wild-collected morels

likely will have a value

premium in markets for

many years to come.

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There is no reason to

believe that cutting

morels does more harm

than pulling.

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Processing—

Ecology and Management of Morels Harvested From the Forests of Western North America

Marketing—

Culinary use—

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24

Toxins and Contaminants

Toxins in look-alike mushrooms—

Gyromitra Verpa

Gyromitra Verpa

24

The cardinal rule is to

always cook morels

thoroughly and start

with modest quantities

until you become

effects.

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Gyromitra gigas

Verpa bohemica Gyromitra

Verpa bohemica

Morel toxins—

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M esculenta

M conica

Another concern

is whether morels

can absorb toxic

retardant slurries.

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Public health regulations—

Gyromitra Verpa in

Gyromitra esculenta Verpa bohemica.

Gyromitra esculenta, Verpa bohemica, Helvella

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Gyromitra

esculenta Verpa bohemica

Verpa bohemica

Morel Harvesters

Personal Use

We know of no

documented instances

of consumers being

seriously poisoned by

wild mushrooms that

were commercially

collected in western

North America.

Ecology and Management of Morels Harvested From the Forests of Western North America

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Commercial

pounds

Ethnic diversity—

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Residence—

Income strategies—

Mobility—

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Patch-lines—

Harvest site portfolios—

Ecology and Management of Morels Harvested From the Forests of Western North America

tree death are scattered

and unpredictable.

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Cooperation—Information is a

particularly important

form of social currency,

and buying stations

constitute one of the

most important sites of

information exchange.

Ecology and Management of Morels Harvested From the Forests of Western North America

Motivations—

Wages—

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Opinions about forest management—

Ecology and Management of Morels Harvested From the Forests of Western North America

Morel Commerce

International

Globally, 300,000

pounds of dried morels

are traded annually.

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25

Ecology and Management of Morels Harvested From the Forests of Western North America

Economics

Value—

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The usefulness of

drying and holding

morels for sale later

in the year depends

on the cost of drying

and holding the

mushrooms and the

international supply

of dried morels in any

given year.

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26

Table 3—Estimated quantities of morels harvesteda and value of commercial mushroom permits

sold on national forests in Oregon and Washington during 2004 and 2005

January-June 2004 January-June 2005

Pounds Dollars Pounds Dollars

b

a

b

GENERAL TECHNICAL REPORT PNW-GTR-710

Markets—

Businesses—

Ecology and Management of Morels Harvested From the Forests of Western North America

Local Impacts

Social impacts—

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Environmental impacts—

Morchella

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83

Policy and Regulation

Land ownership and harvest access—

Development of regulations—

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Current regulations—

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)

Challenges—

Ecology and Management of Morels Harvested From the Forests of Western North America

Reasonable prices,

regulations that

are sensible, and

convenient times,

places, and means

for obtaining permits

all enhance harvester

acceptance.

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Regional Summaries

Mountains of Mexico

Morchella an-

gusticeps M. conica, M. costata, M. elata M. crassipes,

M. esculenta

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89

Morchella

Morchella

M. elata M. esculenta

M. guatemalensis

M. rufobrunnea

Clethra

M.

deliciosa M. rufobrunnea

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M. esculenta

Abies religiosa

Pinus patula

Abies hickelii

Morel harvesting in

Mexico is a seasonal

and traditional

activity of indigenous

and mestizo (mixed

indigenous and

European ancestry)

communities.

Ecology and Management of Morels Harvested From the Forests of Western North America

Morchella

M. angusticeps, M. conica, M. costata, M. elata,

M. esculenta, M. rufrobrunea M. umbrina

unidades de manejo ambiental

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Interior West and Rocky Mountains

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Polytrichum

Ceanothus prostratus

Ecology and Management of Morels Harvested From the Forests of Western North America

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Letters that request

collecting opportunities

mailboxes of USFS

botanists and forest

supervisors on national

forests with recent

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Coast and Cascade Ranges in Oregon and Washington

Ecology and Management of Morels Harvested From the Forests of Western North America

To date, managers

remain largely

unconcerned

about whether

morel harvesting is

sustainable.

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Boletus pinophilus

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Table 4—Number and value of mushrooma-harvesting

permits sold by the tri-forest permit collaboration of

the Malheur, Umatilla, and Wallowa-Whitman National

Forests

Year Permits sold Value

Number Dollars

a

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27

Private timberland

owners typically

concentrate on salvage

and have safety and

liability concerns

regarding mushroom

harvesters during

these operations.

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Western Canada

28

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29

Ecology and Management of Morels Harvested From the Forests of Western North America

30 31

32

33

34

30

31

32

33 a a

34

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36

37

35

36

37

Ecology and Management of Morels Harvested From the Forests of Western North America

An overlay of un-

extinguished aboriginal

rights and titles also

govern rights of access

and use of much of

British Columbia’s

public lands.

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Interior Alaska

Morchella

Ecology and Management of Morels Harvested From the Forests of Western North America

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Management and Research

Sustainability

Ecology and Management of Morels Harvested From the Forests of Western North America

Productivity

Often the only way to

reasonably monitor

sustainable production

is on the scale of

landscapes and over

periods of decades or

centuries.

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Table 5—Unbiased stand-level estimates of mean morel productivity (counts

forest stands in northeastern Oregon that were selected to represent three stand

conditionsa

1995d 1996 Stand

Site locationb conditionc Count Weight Count Weight

Morels/acre Pounds/acre Morels/acre Pounds/acre

a

b

c

d

Ecology and Management of Morels Harvested From the Forests of Western North America

British Columbia, and the Northwest Territories

Year Mean

a count weight Reference

Number/ac lbs/ac

b

c

c

c

d e

f

a

b

c

d

e

f

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Forest Management

Tree death—

A key research

question is whether

mycelial colonies or

sclerotia of morels

persist in the soil

through stand-

replacement events.

Ecology and Management of Morels Harvested From the Forests of Western North America

Timber harvesting—

Soil disturbance—

Fires—

Intentional disruption

or compaction of soil

has too many potential

negative impacts on

tree growth to consider

its use for enhancing

morel crops.

GENERAL TECHNICAL REPORT PNW-GTR-710

Other issues—Morels from western

North America could

command a premium

price if they can be

marketed as collected

from pristine areas.

Ecology and Management of Morels Harvested From the Forests of Western North America

Enhancing morel harvest opportunities—

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Harvest Management

Equitable access—

38

Ecology and Management of Morels Harvested From the Forests of Western North America

Land tenure and harvest pressure—

Liability insurance—

and regulations are

apt to be more widely

accepted if managers

involve all interested

parties in making

regulatory decisions.

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Useful Research

Ecology and Management of Morels Harvested From the Forests of Western North America

Taxonomy— Morchella

Morchella

Morchella

The jumbled taxonomy

of North American

Morchella species

and their lack of valid

species names is

perhaps the biggest

current impediment to

morel research.

GENERAL TECHNICAL REPORT PNW-GTR-710

Ecology—

Morchella

Reproduction and population genetics—

Ecology and Management of Morels Harvested From the Forests of Western North America

Morchella

Morel commodity chains

Closing Remarks

The capacity of

state, federal, and

provincial governments

in western North

America to support

the wild mushroom

industry is hampered

by a profound lack of

published information

about morel commodity

chains.

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Ecology and Management of Morels Harvested From the Forests of Western North America

GENERAL TECHNICAL REPORT PNW-GTR-710

Glossary

allele—

anastomosis—

ascomycetes—

ascus, asci (plural)—

Morchella

ascospores—

asexual

basidium, basidia (plural)

Morchella,

chromosome—

clade—

conidiophore—

conidium, conidia (plural)—

cytoplasm

Ecology and Management of Morels Harvested From the Forests of Western North America

dikaryotic—

diploid—

ectomycorrhiza—

fruiting body (American usage) or fruit body (elsewhere)—

Morchella

fungivore—

genome—

haploid—

Hartig net—

heterokaryotic—

hymenium—

hypha, hyphae (plural)—

hypogeous—

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karyon—

macroscopic—

meiosis—

multikaryotic—

multinucleate—

mycelium, mycelia (plural)—

mycology—

mycophagy—

mycophilic—

mycophobic—

mycorrhiza, mycorrhizae or mycorrhizas (plural)—

nucleus, nuclei (plural)—

polymorphic—

primordium, primordia (plural)—

rhizomorphs—

Ecology and Management of Morels Harvested From the Forests of Western North America

riparian

saprobe, saprobic—

sclerotium, sclerotia (plural)—

sensu lato—

septum, septa (plural)—

sporocarp—

Morchella

stipe—

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Acknowledgments

Ecology and Management of Morels Harvested From the Forests of Western North America

Metric Equivalents

Literature Cited39

Acker, R. 1986

Aegerter, M. 1995.

Alaska Wildland Fire Coordination Group. 1998.

f

Alexander, S.J.; Fight, R.D. 2003.

Alexander, S.J.; Weigand, J.F.; Blatner, K.A. 2002b.

39

GENERAL TECHNICAL REPORT PNW-GTR-710

Alexopoulos, C.J.; Mims, C.W.; Blackwell, M. 1996.

Amaranthus, M.; Pilz, D. 1996.

Ambrose, S.E. 1996.

Morchella esculenta

Morchella esculenta

Morchella esculenta

Morchella esculenta

Morchella esculenta

Antypas, A.; McLain, R.J.; Gilden, J.; Dyson, G. 2002.

Apfelbaum, S.I.; Haney, A.; Dole, R.E. 1984. Morchella

angusticeps

Apple, R.W., Jr. 1998.

Arkan, O. 1992. Morchella conica

Ecology and Management of Morels Harvested From the Forests of Western North America

Morchella conica

Arkan, O.; Güler, P. 1991. Morchella conica

Morchella conica

Arnolds, E. 1995

Arora, D. 1986.

Arora, D. 1999.

Arora, D. 2000.

Arora, D.K.; Bridge, P.D.; Bhatnagar, D., eds. 2004.

Barnes, S.; Wilson, A. 1998.

Benjamin, D.R. 1995.

Beug, M.W. 2004.

Birren, B.; Fink, G.; Lander, E. 2003.

f

Biswell, H.H. 1989.

GENERAL TECHNICAL REPORT PNW-GTR-710

Blanco-Dios, J.B. 2002.

Morchella conica Morchella

conica

Blatner, K.A.; Alexander, S.J. 1998.

Boa, E. 2004.

Boom, M. 1995.

.

Brock, T.D. 1951. Morchella esculenta

Bromley, D.W. 1991.

Brown, B.A.; Marin-Hernandez, A. 2000.

Brown, C. 2005.

Morchella

Morchella Verpa Disciotis

Ecology and Management of Morels Harvested From the Forests of Western North America

Burros, M. 1985.

Buscot, F. 1989. Morchella

rotunda Mitrophora semilibera

Buscot, F. 1992a.

Buscot, F. 1992b.

Buscot, F. 1992c. Morchella

esculenta Picea abies

Buscot, F. 1993. Morchella esculenta

Buscot, F. 1994.

Morchella elata Picea abies

Buscot, F.; Bernillon, J. 1991.

Morchella esculenta

Buscot, F.; Kottke, I. 1990. Morchella rotunda

Picea abies

Buscot, F.; Roux, J. 1987.

Morchella rotunda

GENERAL TECHNICAL REPORT PNW-GTR-710

California Code of Regulations. 1996.

Carpenter, S.; Trappe, J.M.; Ammirati, J., Jr. 1987.

Casey, K. 1995.

Chandra, A. 1989.

Chang, S.-T.; Miles, P. 2004.

Cobanoglu, S.; Bayram, S. 1998.

Morchella

Constantin, J. 1936.

Coombs, D. 1986.

Coombs, D. 1994.

Crisan, E.V.; Sands, A. 1978.

Crous, P.W.; Cother, E.J. 2003.

Czarnecki, J. 1986.

Ecology and Management of Morels Harvested From the Forests of Western North America

Morchella

Dähncke, R.M.; Dähncke, S.M. 1984.

Dalgleish, H.J.; Jacobson, K.M. 2005.

Morchella esculenta

de Geus, P.M.J. 1995.

DeWolf, S. 2001

s

Duchesne, L.C.; Weber, M.G. 1993. Morchella

conica Pinus banksiana

GENERAL TECHNICAL REPORT PNW-GTR-710

Fabricant, F. 1980.

Morchella

Ferndock, M. (Each, P.; Mikkelsen, A., eds.) 1986.

Fine, G.A. 1998.

Fire-Trol Holdings, L.L.C. 1999.

Fischer, D.W.; Bessette, A.E. 1992.

Fries, E.M. 1822.

Fritze, H.; Pennanen, T.; Kitunen, V. 1998.

Ecology and Management of Morels Harvested From the Forests of Western North America

Pinus ponderosa

Gadbow, D. 2001a.

Gadbow, D. 2001b

Gecan, J.S.; Cichowicz, S.M. 1993.

Gessner, R.V. 1995.

Morchella

Gessner, R.V.; Romano, M.M.; Schultz, R.W. 1987.

Morchella deliciosa M esculenta

Ghosh, S.; Majumdar, S.K. 1986.

Morchella

Gilbert, F. 1961. Morchella

Gilmore, M.R. 1919.

GENERAL TECHNICAL REPORT PNW-GTR-710

f

Giunta Regionale dell’Umbria. 1987.

Morchella conica

Gove, P.B.; Merriam-Webster Editorial Staff, eds. 1993.

Grace, B.C. 2005.

Greis, H. 1940. Morchella

Morchella

Guin, J. 1997

Güler, P. 2000. Morchella esculenta

Guzmán, G. 1977.

Guzmán, G.; Tapia, F. 1998.

Morchella rufobrunnea M guatemalensis

Guzmán-Dávalos, L.; Rodríguez-Alcantar, O. 1993. Morchella

guatemalensis Morchella guatemalensis

Ecology and Management of Morels Harvested From the Forests of Western North America

Hallen, H.; Volk, T.; Adams, G. 2001.

Hammond, C. 1999.

Hanlin, R.T.; Hahn, C.G. (illustr.). 1990.

Hansis, R. 1998.

Harbin, M.; Volk, T.J. 1999.

Härkönen, M. 1988.

Härkönen, M.; Järvinen, I. 1993.

Haugen, J. 1994.

Haugen, J. 1998.

Hemmes, D.E.; Desjardin, D.E. 2002.

Helfferich, D. 2005.

Hervey, A.; Bitis, G.; Leong, I. 1978.

Morchella esculenta

GENERAL TECHNICAL REPORT PNW-GTR-710

Hosford, D.; Pilz, D.; Molina, R.; Amaranthus, M.P. 1997.

Huffman, D.M.; Tiffany, L.H. 2001.

Morchella conica, elata y esculenta Morchella

conica, elata esculenta

Iqbal, M. 1993.

Jacobs, M.E. 1982.

Jacquetant, E. 1984.

Johnston, G. 2001

Jones, E.T.; McLain, R.J.; Weigand, J.F., eds. 2002.

Jung, S.W.; Gessner, R.V.; Keudell, K.C.; Ramano, M.A. 1993.

Morchella esculenta

Ecology and Management of Morels Harvested From the Forests of Western North America

Kasischke, E.S.; Rupp, T.S.; Verbyla, D.L. 2006.

Morchella

Keefer, M.E. 2005.

Kendrick, B. 2001.

Korhonen, M. 1986.

Kreisel, H. 2005.

Kuo, M. 2002 (December).

Kuo, M. 2005.

Kuo, M. 2006 (February).

GENERAL TECHNICAL REPORT PNW-GTR-710

Leonard, T.J.; Volk, T.J. 1992.

LeVaux, A. 2005.

Lonik, L. 2002.

Love, T.; Jones, E. 2001.

Love, T.; Jones, E.; Liegel, L. 1998.

Low, J. 1995.

Mader, R. 2005.

Malchow, D. 1991 (27 December).

Masaphy, S. 2005. Morchella

Ecology and Management of Morels Harvested From the Forests of Western North America

Matruchot, L. 1892.

Morchella

McKnight, K.H. 1987.

McLain, R.J. 2000.

McLain, R.J.; Christensen, H.H.; Shannon, M.A. 1998.

McLain, R.J.; Jones, E.T. 2001.

McLain, R.J.; McFarlane, E.; Alexander, S.J. 2005.

McRobert, G. 1985.

Methodus Consultora S.C. 2005.

Miller, H. 2003.

Miller, J. 2004a.

Miller, J. 2004b.

GENERAL TECHNICAL REPORT PNW-GTR-710

Miller, S.L.; Torres, P.; McClean, T.M. 1994.

Morchella

Minnesota Statutes. 1998. Morchella esculenta

Moerman, D.E. 1998.

Molina, R.; Massicotte, H.; Trappe, J.M. 1992.

Molliard, M.M. 1904a. Morchella esculenta

Morchella esculenta

Molliard, M.M. 1904b.

Montoya, A.; Hernández-Totomoch O.; Estrada-Torres A.; Kong A. 2003.

Montréal Process Working Group. 1999.

Ecology and Management of Morels Harvested From the Forests of Western North America

Montréal Process Working Group. 2003.

f

Moser, M. 1949. Morchella

Morchella

Mueller, G.M.; Bills, G.F.; Foster, M.S., eds. 2004.

Obst, J.; Brown, W. 2000.

Obst J.; Coedy, W.; Bromley, R.G. 2001.

GENERAL TECHNICAL REPORT PNW-GTR-710

Morchella

O’Driscoll, P. 2001.

Olynyk, J.; Bergner, K. 2002.

f

Ower, R. 1982. Morchella

esculenta

Ower, R.D.; Mills, G.L.; Malachowski, J.A. 1986 (June 17).

Morchella

Ower, R.D.; Mills, G.L.; Malachowski, J.A. 1988 (July 19).

Morchella

Ower, R.D.; Mills, G.L.; Malachowski, J.A. 1989 (September 19).

Morchella

Palm, M.E.; Chapela, I.H., eds. 1997.

Ecology and Management of Morels Harvested From the Forests of Western North America

Parks, C.G.; Schmitt, C.L. 1997.

Penticton Herald. 2004

Perry, J.P., Jr.; Graham, A.; Richardson, D.M. 1998.

Pinus

Persoon, C.H. 1797.

Peterson, R.L.; Massicotte, H.B.; Melville, L.H. 2004

Philippoussis, A.; Balis, C. 1995.

Morchella

Pietikäinen, J. 1999.

Pietikäinen, J.; Hiukka, R.; Fritze, H. 2000.

Pilz, D.; Ballard, H.L.; Jones, E.T. 2006b.

Pilz, D.; Molina, R. 1998.

GENERAL TECHNICAL REPORT PNW-GTR-710

Pilz, D.; Molina, R. 2002.

Pilz, D.; Molina, R.; Mayo, J. 2006c.

Morchella esculenta

Pugh, G.J.F.; Boddy, L. 1988.

Pyne, S.J.; Andrews, P.L.; Laven, R.D. 1996.

Ramsbottom, J. 1953.

Rédei, G.P. 2003.

Redhead, S.A. 1997.

Robbins, W.J.; Hervey, A. 1959. Morchella

Ecology and Management of Morels Harvested From the Forests of Western North America

Rolfe, R.T.; Rolfe, F.W. 1925.

:

Royse, D.J.; May, B. 1990. Morchella

Schlosser, W.E.; Blatner, K.A. 1995.

Schlosser, W.E.; Blatner, K.A. 1997.

Schmidt, E.L. 1979. Morchella

Schmidt, E.L. 1983.

Morchella esculenta

2004.

Proyecto de Conservacion y Manejo Sustentable de Recursos Forestales

Morchella elata Morchella esculenta

Morchella elata

GENERAL TECHNICAL REPORT PNW-GTR-710

Morchella esculenta

Shanley, P.; Pierce, A.R.; Laird, S.A.; Guillen, S.A. 2002.

Morchella esculenta

Morchella esculenta

Morchella esculenta

Morchella esculenta

Ecology and Management of Morels Harvested From the Forests of Western North America

Smith, S.E.; Read, D.J. 1997.

Stabenow, D. 1995.

Stamets, P. 1993.

Stamets, P. 2000.

Stamets, P. 2005.

Still, G.; MacKinnon, A.; Planden, R. 1994.

Stott, K.G.; Gill, W.; Mohammed, C.; Brown, M. 2002.

Stott, K.G.; Mohammed, C. 2004.

Sturgis, W.C. 1905. Morchella esculenta

Szuecs, J. 1958 (September 9).

Szuecs, J.; Wiles, G.M. 1950 (May 2).

GENERAL TECHNICAL REPORT PNW-GTR-710

Taylor, J.W.; Spatafora, J.; Berbee, M. 1996.

i

Tedder, S.; Mitchell, D.; Farran, R. 2000.

Tedder, S.; Mitchell, D.; Hillyear, A. 2002.

Thompson, V.V. 1994.

Tiffany, L.H.; Knaphus, G.; Huffman, D.M. 1998.

Trappe, J.M. 1996.

Trappe, J.M.; Molina, R.; Castellano, M. 1984.

Treseder, K.K.; Mack, M.C.; Cross, A. 2004.

Tumlin, H. 2005

Ulloa, M.; Hanlin, R.T. 2000.

Ecology and Management of Morels Harvested From the Forests of Western North America

1996.

2003

f

Vehling, J.D., ed. and transl. 1977.

Apicius

de re Coquinaria

GENERAL TECHNICAL REPORT PNW-GTR-710

Villarreal, L.; Gómez, A. 1997.

Villarreal, L.; Pérez-Moreno, J. 1989.

Villarreal-Ruiz, L. 1994.

Villarreal-Ruiz, L., ed. 1996.

Volk, T.J. 1990.

Volk, T.J. 1991.

Volk, T.J.; Kozak, M.E.; Krawczyk, J. 1997.

Ecology and Management of Morels Harvested From the Forests of Western North America

Volk, T.J.; Leonard, T.J. 1989a.

Morchella

Volk, T.J.; Leonard, T.J. 1989b.

Morchella crassipes

Volk, T.J.; Leonard, T.J. 1990. Morchella

Vrålstad, T.; Holst-Jensen, A.; Schumacher, T. 1998.

Geopyxis carbonaria

Picea abies

Walstad, J.D.; Radosevich, S.R.; Sandberg, D.V. eds. 1990.

Morchella

GENERAL TECHNICAL REPORT PNW-GTR-710

Wedin, M.; Döring, H.; Gilenstam, G. 2004.

Stictis Conotrema

Werner, P. 2002.

Wheeler, S. 1996.

Willetts, H.J. 1972

Wills, R.M.; Lipsey, R.G. 1999.

Winder, R. 2006. Morchella elata

Wipf, D.; Fribourg, A.; Munch, J.C. 1999.

Wipf, D.; Munch, J.C.; Botton, B.; Buscot, F. 1996b.

Morchella esculenta Morchella conica

World Bank. 1995.

Ecology and Management of Morels Harvested From the Forests of Western North America

World Bank. 1997.

Yamin, S. 2005.

Yoon, C.-S.; Gessner, R.V.; Romano, M.M. 1990.

Morchella esculenta

Yukon Energy, Mines, and Resources. 2004.

f

Zamora, M.; Torres, J.M.; Zamora, L. 2001.

Podospora anserina

GENERAL TECHNICAL REPORT PNW-GTR-710

a

True morelsb

Morchella Costantinella cristata

Morchella

Morchella angusticeps

Morchella esculenta atrotomentosa

Morchella atrotomentosa

Morchella conica

Morchella costata

Morchella crassipes

M.

esculenta

Morchella deliciosa

Morchella elata

Morchella esculenta

Morchella guatemalensis

Morchella herediana

Morchella hortensis

Morchella intermedia

M. conica

Morchella rigidoides

Morchella rotunda Morchella

esculenta

Morchella rufobrunnea

Morchella semilibera Mitrophora

semilibera

Morchella umbrina

M. esculenta

Morel look-alikes and related genera:

Discina

Disciotis venosa

Ecology and Management of Morels Harvested From the Forests of Western North America

Gyromitra gigas

Gyromitra esculenta

Gyromitra

Gyromitra Helvella

Verpa bohemica

Verpa

Other fungi:

Agrocybe aegerita

Agaricus brunnescens

Boletus

Boletus edulis

Boletus pinophilus

Gymnosporangium juniperi-virginianae

Cantharellus

Cantharellus,

Craterellus, Gomphus, Polyozellus.

Geopyxis carbonaria

Lentinula edodes

Ophiostoma ulmi

Pholiota nameko

Pleurotus

Podospora anserina

Sclerotina sclerotiorum

Tricholoma magnivelare

Tricholoma matsutake

Tricholoma Tricholoma magnivelare

Tricholoma caligatum

Tricholoma

matsutake

Tuber

Tuber

Xerophyllum tenax

Pteridium aquilinum

GENERAL TECHNICAL REPORT PNW-GTR-710

Polytrichum

Fragaria

Trillium

Shrubs:

Vaccinium

Syringa vulgaris

Holodiscus discolor

Ceanothus prostratus

Gaultheria shallon

Rubus tomentosus

Trees:

Alnus

Malus pumila

Fraxinus

Populus

Fagus

Prunus serotina

Picea mariana

Betula

Eucalyptus globus

Prunus laurocerasus

Castanea sativa

Austrocedrus chilensis

Populus

Pseudotsuga menziesii

Ulmus

Picea engelmannii

Alnus glutinosa

Abies hickelii

Carya

Pinus banksiana

Juniperus

Robinia pseudoacacia

Pinus contorta contorta

Acer

Pinus radiata

Picea abies

Quercus

Abies religiosa

Betula papyrifera

Pinus

Pinus ponderosa

Populus

Ecology and Management of Morels Harvested From the Forests of Western North America

Pinus jeffreyi Pinus ponderosa

Pinus coulteri

Nothofagus

Abies lasiocarpa

Larix lyallii

Liquidambar

Platanus

Abies

Liriodendron tulipifera

Pinus patula

Tsuga heterophylla

Larix occidentalis

Clethra

Abies concolor

Picea glauca

Pinus albicaulis

Salix

Insects:

Orgyia pseudotsugata

Aradus debilis

Lymantria dispar

Dendroctonus ponderosae

Choristoneura occidentalis

Animals:

Cervus canadensis

Ursus arctos horribilis

Canis lupus

a

b

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