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Taxonomic revision of true morels (Morchella) in Canada and the United States

April 2012
Mycologia 104(5):1159-77

DOI:10.3852/11-375

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PubMed

Authors:

Michael Kuo

K. O'Donnell

Agricultural Research Service

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Recent molecular phylogenetic studies have revealed the existence of at least 50 species of Morchella worldwide and demonstrated a high degree of continental endemism within the genus. Here we describe 19 phylogenetic species of Morchella from North America, 14 of which are new (M. diminutiva, M. virginiana, M. esculentoides, M. prava, M. cryptica, M. frustrata, M. populiphila, M. sextelata, M. septimelata, M. capitata, M. importuna, M. snyderi, M. brunnea and M. septentrionalis). Existing species names (M. rufobrunnea, M. tomentosa, M. punctipes and M. angusticeps) are applied to four phylogenetic species, and formal description of one species (M. sp. "Mel-8") is deferred pending study of additional material. Methods for assessing morphological features in Morchella are delineated, and a key to the known phylogenetic species of Morchella in North America is provided. Type studies of M. crassistipa, M. hotsonii, M. angusticeps and M. punctipes are provided. Morchella crassistipa is designated nomen dubium.

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Taxonomic revision of true morels (

Morchella

) in Canada and the United States

Michael Kuo

Department of English, Eastern Illinois University,

Charleston, Illinois 61920

Damon R. Dewsbury

Department of Ecology and Evolutionary Biology,

University of Toronto, Toronto, Ontario M5S2C6

Kerry O’Donnell

Bacterial Foodborne Pathogens and Mycology Research

Unit, National Center for Agricultural Utilization

Research, United States Department of Agriculture,

Agricultural Research Service, 1815 North University

Street, Peoria, Illinois 61604

M. Carol Carter

17070 SW Vendla Park Lane, Aloha, Oregon 97006

Stephen A. Rehner

Systematic Mycology and Microbiology Laboratory,

United States Department of Agriculture, Agricultural

Research Service, Beltsville, Maryland 20705

John David Moore

Department of English, Eastern Illinois University,

Charleston, Illinois 61920

Jean-Marc Moncalvo

Department of Natural History, Royal Ontario

Museum, and Department of Ecology and Evolutionary

Biology, University of Toronto, Toronto, Ontario

M5S 2C6

Stephen A. Canfield

Department of Foreign Languages, Eastern Illinois

University, Charleston, Illinois 61920

Steven L. Stephenson

Department of Biological Sciences, University of

Arkansas, Fayetteville, Arkansas 72701

Andrew S. Methven

Department of Biological Sciences, Eastern Illinois

University, Charleston, Illinois 61920

Thomas J. Volk

Department of Biology, University of Wisconsin, La

Crosse, Wisconsin 54601

Abstract

: Recent molecular phylogenetic studies

have revealed the existence of at least 50 species of

Morchella

worldwide and demonstrated a high degree

of continental endemism within the genus. Here we

describe 19 phylogenetic species of

Morchella

from

North America, 14 of which are new (

M. diminutiva

M. virginiana

M. esculentoides

M. prava

M. cryptica

M. frustrata

M. populiphila

M. sextelata

M. septime-

lata

M. capitata

M. importuna

M. snyderi

brunnea

and

M. septentrionalis

). Existing species

names (

M. rufobrunnea

M. tomentosa

M. punctipes

and

M. angusticeps

) are applied to four phylogenetic

species, and formal description of one species (

. sp.

‘‘

Mel

-8’’) is deferred pending study of additional

material. Methods for assessing morphological fea-

tures in

Morchella

are delineated, and a key to the

known phylogenetic species of

Morchella

in North

America is provided. Type studies of

M. crassistipa

hotsonii

M. angusticeps

and

M. punctipes

are provid-

ed.

Morchella crassistipa

is designated nomen dubium.

Key words:

Ascospore, identification key, Morchel-

laceae, morels, morphology, North America, pits,

taxonomy

NTRODUCTION

Morels (species of

Morchella

Dill. ex Pers.) are highly

valued edible mushrooms in the northern hemi-

sphere, especially in Europe and North America

(Weber 1995, Kuo 2005), but we still lack an

understanding of many aspects of their biology,

taxonomy and distribution. Molecular phylogenetic

analyses of 590

Morchella

collections (O’Donnell et al.

2011) revealed at least 41 phylogenetic species

worldwide, 19 of which appeared to be endemic to

North America. Results additionally indicated a high

degree of continental endemism for the genus, with

only two of the 41 species occurring naturally in both

Europe and Asia and none occurring naturally in

both North America and Eurasia (although a few

North American species appear to have been intro-

duced into Turkey; see Tas¸kın et al. 2010, 2012).

Because extensive sampling has indicated strict North

American endemism, existing European and Asian

species names (including among others the well

known names

M. esculenta

[L] Pers.,

M. elata

Fr.,

M. semilibera

DC and

M. conica

Pers.) could not be

applied to North American morels. We studied the

type collections of the six species originally described

from North America (

M. tomentosa

M. Kuo,

rufobrunnea

Guzma´n & F. Tapia,

M. angusticeps

Peck,

M. punctipes

Peck,

M. hotsonii

Snyder,

M. crassistipa

Snyder) to investigate the contemporary applicability

of these species names (see S

UPPLEMENTARY MATERI-

ALS

). We also studied morphological, ecological and

distributional data from 244 North American collec-

tions identified phylogenetically to assess whether

Submitted 12 Nov 2011; accepted for publication 3 Mar 2012.

Corresponding author. E-mail: tvolk @uwlax.edu

Mycologia,

104(5), 2012, pp. 1159–1177. DOI: 10.3852/11-375

2012 by The Mycological Society of America, Lawrence, KS 66044-8897

1159

these species can be defined on the basis of

morphology, ecology and distribution. Herein, five

species within the Esculenta Clade (yellow morels)

andninespecieswithintheElataClade(black

morels) from North America are formally described

and

M. angusticeps

is epitypified (FIG. 1).

ATERIALS AND METHODS

Collections were accessioned into the Morel Data Collection

Project (MDCP) database (www.mushroomexpert.com/

mdcp/mdcprecords.pl), together with collectors’ notes

and photographs. Phylogenetic identification of 244 collec-

tions was accomplished following the methods for DNA

isolation, PCR amplification and sequencing, and phyloge-

netic analysis described in O’Donnell et al. (2011).

Macromorphology of specimens was determined through

observation of fresh specimens, dried specimens, collectors’

notes and photographs of material in the fresh state. (See

UPPLEMENTARY MATERIALS for details on specific features.)

For micromorphological study horizontal scalp sections

taken from the hymenia and sterile ridges were squash-

mounted in 2% KOH. Measurements of asci, paraphyses

and elements on sterile ridges were made at 4003

magnification;ascosporemeasurementsweremadeat

10003. At least 10 mature ascospores were measured for

each specimen. Collections were deposited in the Mycology

Collection of the Field Museum of Natural History in

Chicago (F) using MDCP accession numbers or were

studied and deposited in the Fungarium at the Royal

Ontario Museum in Toronto (TRTC). Others were

returned to lending herbaria. DNA sequence data from

type specimens described in this study were deposited in

GenBank under accession numbers JQ670119–JQ670132.

RESULTS

Study of 244 North American collections identified

phylogenetically (S

UPPLEMENTARY TABLE I) revealed five

species within the

Morchella esculenta

Clade as defined

by O’Donnell et al. (2011) and nine species within the

Morchella elata

Clade (FIG. 1). Among these, a few

phylogenetic species (

Morchella rufobrunnea

M. tomen-

tosa

M. importuna

M. prava

M. frustrata

) could be

differentiated on the basis of morphological characters,

and several others (

M. punctipes

M. populiphila

brunnea

M. capitata

M. snyderi

) could be distinguished

when geographic range and/or ecological factors are

combined with morphological characters. However,

several phylogenetic species (including the pair

sextelata

and

M. septimelata

and the pair

M. esculentoides

and

M. cryptica

) remain morphologically and ecologi-

cally cryptic based on present data. Type studies (see

UPPLEMENTARY MATERIALS) supported applying four of

the six previous North American species names (

angusticeps

M. punctipes

M. rufobrunnea

M. tomen-

tosa

) to phylogenetic species. However, two previous

North American names were problematic: (i) the type

collection of

M. crassistipa

was determined to be mixed,

obliging us to designate this name as nomen dubium;

and (ii) preliminary DNA sequence data on

M. hotsonii

suggested that it might represent a novel phylogenetic

species, apparently undocumented since the 1935 type

collection. As a result of our analyses 14 new species are

described below. Description of another new species was

deferred pending additional material for study. In

addition, we identified a suite of morphological

characters for study of

Morchella

specimens (see

UPPLEMENTARY MATERIALS).

AXONOMY

Morchella rufobrunnea

Clade

The

Morchella rufobrunnea

Clade (O’Donnell et al. 2011)

is basal to the remaining

Morchella

lineage and consists of a

single species. Morphologically

M. rufobrunnea

resembles

the ‘‘yellow morels’’ as they have been defined traditionally

(Kuo 2005) and most members of the

Morchella esculenta

Clade.

Morchella rufobrunnea

Guzma´n & F. Tapia, Mycologia

90:706. 1998. F

IG.2

The original description of

Morchella rufobrunnea

ap-

peared in Guzma´n and Tapia (1998). (For an augmented

description based on material studied by the authors see

UPPLEMENTARY MATERIALS.)

Comments. Morchella rufobrunnea

is easily distin-

guished on the basis of ‘‘its abruptly conical young

cap with pale ridges and nearly black pits, and its

rufescence’’ (Kuo 2008). It appears in woodchips and

landscaping settings on the West Coast from California

to Seattle. (For a discussion of species names fre-

quently misapplied to

M. rufobrunnea

see Kuo 2008.)

Molecular phylogenetic analysis of F 03110601 and sev-

eral other commercially produced morels (O’Donnell

unpubl data) confirms

M. rufobrunnea

as the morel

cultivated commercially (US Patent numbers 4594809,

4757640).

Morchella esculenta Clade

The

Morchella esculenta

Clade (O’Donnell et al. 2011) is

sister to the

M. elata

Clade and corresponds fairly well with

the traditional morphological concept of ‘‘yellow morels’’

(Kuo 2005), amended to exclude

M. rufobrunnea

, which has

pale ridges like yellow morels but is phylogenetically

distinct, and to exclude

M. frustrata

and

M. snyderi

, which

have the colors usually associated with yellow morels but are

nested within the

M. elata

Clade. The latter two species,

however, possess pits that primarily are elongated vertically

and feature a notable sinus where the hymenophore

attaches to the stipe; most members of the

M. esculenta

Clade have less vertically arranged pits and feature

hymenophores that are typically adnate.

1160 MYCOLOGIA

Morchella diminutiva M. Kuo, Dewsbury, Moncalvo &

S.L. Stephenson, sp. nov. F

IG.3

MycoBank MB 563948

Ascomata 35–94 mm alta; capitulum conicum; costae

pallidae, colocatae recte; hymenium sufflavum colore

subcaesio; biotopium in silva frondosa in orienti America

septentrionalis, praesertim in alveis ubi

Fraxinus americana

L aut

Liriodendron tulipifera

L praesens est; sporae 20–24 3

11–16

mm. Holotypus: Biotopium in silva frondosa; USA, in

Illinoisense, ad Coles County; M. Kuo col.; specimen

typicum in herbarium F (05030404) conservatum.

Etymology.

The epithet refers to the size of the

ascomata.

Ascomata 35–94 mm high. Hymenophore 20–

41 mm high; 10–27 mm wide at the widest point;

usually conical to subconical, occasionally ovoid or

subcylindrical with a subacute apex; pitted and

ridged, with 8–16 primary vertical ridges and occa-

sional secondary vertical ridges, with scattered to

frequent, sunken, transecting horizontal ridges;

adnate at the point of attachment, or when young

attached to stipe with a slight sinus. Ridges glabrous;

yellowish to nearly whitish when young, becoming

pale brownish yellow or pale yellowish brown with

maturity; widely rounded to nearly flat when young,

but with age often becoming sharpened or eroded.

Pits primarily vertically elongated; glabrous; medium

to dark gray or grayish brown when young, becoming

pale brownish yellow or pale yellowish tan. Stipe

10–68 mm high; 3–18 mm wide; more or less equal,

or sometimes basally subclavate; finely mealy with

whitish granules or nearly glabrous; whitish. Context

whitish; about 1 mm thick in the hollow hymeno-

phore; in the stipe sometimes slightly chambered

near the base. Sterile inner surface whitish and

pubescent. Ascospores (18–)20–24(–26) 3 (10–)11–

16(–18)

mm; elliptical; smooth; contents homoge-

neous; orangish yellow in deposit. Asci 175–325 3

12.5–25

mm; eight-spored; cylindrical; hyaline. Pa-

raphyses 125–250 3 7.5–20

mm; cylindrical; apices

generally merely rounded but occasionally subfusi-

form, subcapitate, or irregular; septate; brown to

brownish in KOH (2%) on immature ascomata,

becoming hyaline with maturity. Elements on sterile

ridges 75–175 3 (7.5–)10–30

mm; hyaline to brownish

or brown in KOH (2%); septate; terminal cell

subfusiform by maturity but sometimes merely cylin-

drical, with a rounded or subcapitate apex.

Ecology.

Appearing in eastern North American

hardwood forests, especially in association with

Fraxinus americana

Land

Liriodendron tulipifera

but also found under

Carya

spp. and other hardwoods;

widely distributed east of the Rocky Mountains,

although the northern limits of its range are undeter-

mined; April and May. Specimens examined (S

UPPLE-

MENTARY TABLE

I) were collected in Illinois, Massachu-

setts, Mississippi, Missouri, Pennsylvania, Tennessee,

Virginia and West Virginia.

Comments. Morchella diminutiva

corresponds to

phylogenetic species

Mes

-2 in O’Donnell et al.

(2011). The species is characterized by its small

average size; its conical to subconical hymenophore,

which features vertically elongated pits and ridges, and

the elements on its sterile ridges, which usually are

subfusiform. It is widely distributed and common east

of the Great Plains under ash, tulip trees and other

hardwoods. Larger specimens are similar to smaller

specimens of

M. virginiana

, which has a limited

southeastern range and scattered elements on sterile

ridges that are variably shaped.

Morchella esculentoides

along with

M. cryptica

and

M. prava

, can be separated

from

M. diminutiva

on the basis of their larger size,

less conical hymenophores, and their pits and ridges,

which are not primarily vertically elongated.

Morchella

diminutiva

corresponds to ‘‘type 1’’ of the ‘‘North

American Deliciosas’’ in Kuo (2005) and, in part, to

M. deliciosa

in Weber (1995).

Morchella virginiana O’Donnell & S.A. Rehner, sp.

nov. F

IG.4

MycoBank MB 563949

Ascomata 50–125 mm alta; capitulum late conicum;

costae pallidae; hymenium sufflavum colore subcaesio;

biotopium in silva frondosa in America vulturna septen-

trionalis, praesertim in alveis ubi

Liriodendron tulipifera

praesens est; sporae 18–25 3 10–16

mmHolotypus:

Biotopium in alveo se conjungens cum

Liriodendron

tulipifera

L; USA, in Virginiaense, ad Fairfax County;

S. Rehner col.; specimen typicum in Herb. BPI (880503)

conservatum.

Etymology.

The epithet refers to the location of the

holotype collection.

Ascomata 50–125 mm high. Hymenophore 30–

70 mm high; 22–35 mm wide at the widest point;

usually ovoid with a bluntly conic to subconic apex,

but occasionally subcylindrical with a rounded apex;

pitted and ridged, with 12–16 primary vertical ridges

and scattered secondary vertical ridges, with scat-

tered, sunken, transecting horizontal ridges; adnate

at the point of attachment. Ridges finely tomentose

to glabrous; pale yellowish when young, becoming

pale brownish yellow or yellowish brown with ma-

turity; flattened when young but often becoming

sharpened or eroded in age. Pits primarily elongated

vertically; glabrous or finely tomentose; grayish

brown when young, becoming pale brownish yellow

or pale yellowish tan but often retaining grayish

hues. Stipe 25–55 mm high; 10–20 mm wide; more

or less equal, or basally subclavate; finely mealy with

whitish granules or nearly glabrous; whitish to pale

yellowish. Context whitish; about 1 mm thick in the

UO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1161

FIG. 1. One of two equally parsimonious phylograms 1674 steps long depicting phylogenetic relationships among 19 North

American and seven European species of

Morchella

inferred from a five-gene dataset containing 600 parsimony informative

characters (PIC). Sequences of

M. rufobrunnea

were used to root the phylogram based on more inclusive analyses (O’Donnell

et al. 2011). The 17 species within the Elata Clade (black morels) and eight species within the Esculenta Clade (yellow morels)

are identified respectively by

Mel

and

Mes

followed by a unique Arabic number. Highlight is used to identify nine species

1162 MYCOLOGIA

hollow hymenophore; in the stipe sometimes slightly

chambered near the base. Sterile inner surface

whitish and pubescent. Ascospores 18–25(–28) 3

10–16

mm; elliptical; smooth; contents homoge-

neous. Asci 200–325 3 15–25

mm; eight-spored;

cylindrical; hyaline. Paraphyses 125–200 3 5–15

mm;

cylindrical; apices generally merely rounded but

occasionally subclavate, subcapitate or subfusiform;

septate; hyaline to brownish in KOH (2%). Elements

on sterile ridges scattered and infrequent (often

difficult to locate or distinguish from paraphyses),

100–175 3 10–30

mm; hyaline to brownish or brown

in KOH (2%); septate; terminal cell widely cylindri-

cal, with a rounded, subcapitate, subclavate or

subfusiform apex.

Ecology.

Appearing in southeastern hardwood forests,

especially in association with

L. tulipifera

in river bottoms,

coastal plains and drainage areas, often in sandy soil;

North Carolina, South Carolina, Mississippi and Virginia;

April and May. Specimens examined (S

UPPLEMENTARY

TABLE

I) were collected in Mississippi, North Carolina,

South Carolina and Virginia.

Comments. Morchella virginiana

corresponds to

phylogenetic species

Mes

-3 in O’Donnell et al.

(2011). Based on current data it can be characterized

as intermediate in stature between

M. diminutiva

and

M. esculentoides

; its dimensions and proportions

approximate large specimens of

M. diminutiva

and

small specimens of

M. esculentoides

. It is usually

less sharply conical than

M. diminutiva

,however,

and its stipe is proportionally longer than that of

M. esculentoides

, from which it also differs in the

primarily vertical (rather than more or less random)

orientation of the pits. While

M. dimin utiva

is found

in a variety of eastern hardwood habitats in associ-

ation with

Fraxi nus

spp.,

L. tulipifera

and other

hardwoods,

M. virginiana

apparently is limited to

association with

L. tulipifera

in riparian and upland

ecosystems from Virginia to northern Mississippi

(O’Donnell et al. 2011). Elements on sterile ridges in

M. virginiana

specimens examined were scattered

and often difficult to isolate; when present they were

variably shaped, cylindrical to subcapitate, subcylind-

rical or subfusiform. Elements on sterile ridges of

diminutiva

, by contrast, were easily demonstrated

and primarily widely fusiform.

Morch ella virginian a

corresponds with ‘‘type 2’’ of the ‘‘North American

Deliciosas’’ in Kuo (2005).

Morchella esculentoides M. Kuo, Dewsbury, Moncalvo

& S.L. Stephenson, sp. nov.

IG.5,SUPPLEMENTARY FIG.1

MycoBank MB 563950

Ascomata 36–220(–415) mm alta; capitulum ovoideum cum

apice late conico aut convexo; costae pallidae; hymenium

sufflavum aetate; biotopium in silva; sporae 18–22 3 11–13

mm.

Holotypus: Biotopium in silva frondosa; USA, in Oregonense,

ad Linn County; N. S. Weber 7114 col.; specimen typicum in

Herb. OSC (138364—138367) conservatum.

Etymology.

The epithet refers to similarity with the

iconic European species

M. esculenta

Ascomata 36–220(–415) mm high. Hymenophore 23–

110(–220) mm high; 15–42(–126) mm wide at the widest

point; usually ovoid with a bluntly conical or convex apex

but occasionally subcylindrical with a convex apex or

subglobose to pyriform; pitted and ridged, with approx-

imately 12–30 vertical ridges and numerous horizontal

and oblique ridges, along with scattered, sunken,

transecting ridges; adnate at the point of attachment.

Ridges glabrous or nearly so; white to pale yellowish when

young, becoming pale brownish yellow or remaining

whitish with maturity; bluntly rounded or nearly flattened

when young, usually becoming sharpened or eroded in

age. Pits usually more or less vertically elongated but not

strictly so and not infrequently subglobose to irregular in

outline; glabrous or finely tomentose; grayish brown to

dark brown or nearly black when young, becoming pale

brownish yellow. Stipe 20–120(–240) mm high; 16–

92(–140) mm wide; usually basally clavate to subclavate;

glabrous or finely mealy with scattered whitish to

yellowish granules; whitish to pale yellowish or brownish.

Context whitish; 1–3 mm thick in the hollow hymeno-

phore; in the stipe often becoming thickened and

chambered near the base with maturity. Sterile inner

surface whitish and pubescent. Ascospores (17–)18–

22(–24) 3 11–13(–15)

mm; elliptical; smooth; contents

homogeneous. Asci 225–325 3 15–25

mm; eight-spored;

cylindrical; hyaline. Paraphyses 75–180 3 5–15

mm;

cylindrical; apices generally merely rounded but occa-

sionally subclavate to clavate or subfusiform; septate;

hyaline to ochraceous or faintly brownish in KOH (2%).

Elements on sterile ridges 75–160 3 10–27.5(–37.5)

mm;

hyaline to ochraceous in KOH (2%); septate; terminal

cell subclavate to clavate, or subfusiform to widely

cylindrical with a rounded or subcapitate apex.

Ecology.

Widely distributed and common; often

found under living and dead hardwoods (especially

living

F. americana

and dead or dying

Ulmus americana

L) but also found in apparent association with

Populus

within the Elata Clade and five species within the Esculenta Clade that are formally described herein. In addition

angusticeps

(

Mel

-15) and

M. punctipes

(

Mel

-4) are epitypified. Numbers above internodes represent bootstrap values $ 65%.

CI 5 consistency index, RI 5 retention index.

KUO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1163

FIGS. 2–10.

Morchella

species. 2.

M. rufobrunnea

F 03080501. 3.

M. diminutiva

HOLOTYPE F 05030404,

Mes

-2. 4.

virginiana

HOLOTYPE BPI 880503,

Mes

-3. 5.

M. esculentoides

F 04250405,

Mes

-4. 6.

M. prava

HOLOTYPE F 05100602,

Mes

-7.

M cryptica

HOLOTYPE F 04220401,

Mes

-11. 8.

M. tomentosa

HOLOTYPE F 06150405. 9.

M. frustrata

UC 1860809,

Mel

-2. 10.

M. punctipes

F 04240304,

Mel

-4. Bars 5 5 cm.

Mel

and

Mes

numbers refer to the phylogenetic species reported in O’Donnell

et al. (2011).

1164 MYCOLOGIA

deltoides

Bartr.,

P. balsamifera

P. grandidentata

Michx.,

L. tulipifera

Platanus occidentalis

Land

Quercus

spp. in a variety of habitats (from riparian

cottonwood-sycamore lowlands to oak-hickory forests,

mixed woods and northern hardwood forests); also

appearing in association with

Malus

spp. in old

orchards and occasionally in association with conifers

(especially

Pinus strobus

L but also with

P. resinosa

Ait.,

Abies balsamea

[L] Mill. and

Picea abies

[L] Karst.);

rarely found in areas with no trees present; widely

distributed and common east of the Rocky Mountains

from Kansas to Ontario and the northeastern United

States, south to Texas, Arkansas and South Carolina; in

western North America occurring under hardwoods in

river bottoms or in urban settings in association with

apple trees or ornamental ash plantings; appearing in

spring (March–June, depending on latitude and

altitude). Specimens examined (S

UPPLEMENTARY TABLE

I) were collected in Arkansas, California, Colorado,

Illinois, Iowa, Kansas, Massachusetts, Minnesota, Mis-

souri, Nebraska, New York, Ohio, Oklahoma, Ontario,

Oregon, Pennsylvania, South Carolina, South Dakota,

Texas, Vermont, Virginia, West Virginia and Wisconsin.

Comments. Morchella

sculentoides

is the most widely

distributed member of the genus in North America

and corresponds to phylogenetic species

Mes

-4 in

O’Donnell et al. (2011). Mature ascomata that have

developed normally can be distinguished morpholog-

ically from all other North American species of

Morchella

, except

M. cryptica

, by its medium to large

ascomata, together with the non-rufescent, pale ridges,

the generally rounded apex of the hymenophore, and

the pits, which are not strictly vertical in arrangement

but are not as asymmetrical and irregular as those of

prava

(however, occasional aberrant specimens of

esculentoides

, such as F 04150501, can appear similar to

M. prava

and must be identified with molecular

analysis). In the Great Lakes region, where the ranges

M. cryptica

and

M. esculentoides

overlap, the two

species cannot be separated reliably based on current

data without molecular analysis. (See the comments

under

M. cryptica

for details.) In western regions

esculentoides

has no close look-alikes, although

rufobrunnea

, which has rufescent ridges, and

frustrata

, which has a conical hymenophore featuring

vertically arranged pits, have similarly colored hyme-

nophores.

Morchell a esc ulentoid es

corresponds (in

part) to

M. esculenta

in Weber (1995) and to the

‘‘Classic North American Yellow Morel’’ in Kuo (2005).

Morchella prava Dewsbury, Moncalvo, J.D. Moore &

M. Kuo, sp. nov. F

IG.6

MycoBank MB 563951

Ascomata 50–100 mm alta; capitulum ovoideum cum

apice subconico; costae pallidae; hymenium pravium, canae

vel atrae, pallescens aetate; biotopium in silva; sporae 17–21

3 10–12

mm. Holotypus: Biotopium in humo harenoso

sub

Pinus strobus

Let

Acer saccharum

Marsh.; USA, in

Michiganense, ad Emmet County; M. Kuo & J.D. Moore col.;

specimen typicum in Herb. F (05100602) conservatum.

Etymology.

The epithet means crooked, irregular or

deformed and reflects the contorted aspect of the

hymenophore.

Ascomata 50–100 mm high. Hymenophore 30–

60 mm high; 20–50 mm wide at the widest point;

irregularly shaped but often more or less ovoid with a

slightly narrowed or subconic apex; pitted and ridged;

with 12–18 vertical ridges intersecting a line across the

widest portion but with numerous horizontal and

oblique ridges as well, along with scattered, sunken,

transecting ridges; adnate at the point of attachment

or occasionally attached with an inconsistent, poorly

defined sinus. Ridges glabrous or very finely tomen-

tose; thick; whitish to pale yellowish or pale tan when

young, becoming pale brownish yellow with maturity

and eventually darkening to yellowish brown or

reddish brown in places; flattened or widely rounded

when young but with age sometimes becoming

sharpened or eroded. Pits asymmetrical and irregular

in outline and size; glabrous or finely tomentose; gray

to dark gray, gray-brown or nearly black when young

and often remaining dark for a long time before

becoming pale brownish yellow or pale yellowish tan

with maturity. Stipe 25–40 mm high; 10–30 mm wide;

more or less equal above a subclavate to clavate base;

glabrous or nearly so; whitish to pale yellowish, often

discoloring reddish brown; usually developing indis-

tinct ridges and folds near the base. Context whitish;

about 1–2 mm thick in the hollow hymenophore;

usually chambered or layered near the base. Sterile

inner surface whitish and pubescent. Ascospores

(16–)17–21(–24) 3 (8–)10–12(–13)

mm; ellipsoid to

slightly subfusiform; smooth; contents homogeneous.

Asci 200–300 3 15–25

mm; eight-spored; cylindrical;

hyaline. Paraphyses 100–175 3 5–12.5

mm; cylindrical;

apices generally merely rounded or subclavate but

occasionally clavate or subfusiform; septate; hyaline to

brownish or brown in KOH (2%). Elements on sterile

ridges scattered and infrequent (often difficult to

locate or distinguish from paraphyses); 75–125 3 7.5–

25(–37.5)

mm; hyaline to ochraceous, brownish, or

brown in KOH (2%); septate; terminal cell widely

cylindrical with a rounded apex, subclavate, clavate,

subcapitate, capitate or widely subfusiform.

Ecology.

The ecology of

Morchella prava

collections

studied was not consistent. Among the seven collections

studied, few if any common ecological denominators

were apparent. Collections were made approximately

43–50uN across North America, in April, May and early

June. (See S

UPPLEMENTARY MATERIALS for further

UO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1165

ecological data from the collections studied.) Speci-

mens examined (S

UPPLEMENTARY TABLE I) were col-

lected in Michigan, Montana, Ontario, Saskatchewan

and South Dakota.

Comments. Morchella prava

corresponds to phyloge-

netic species

Mes

-7 in O’Donnell et al. (2011). The

species usually can be identified on the basis of its

esculenta

-like stature and its contorted, asymmetrical and

irregular pits and ridges. The pits are highly irregular in

outline and size, and the thick, bluntly rounded ridges

are less likely to become eroded and sharpened with

maturity than the ridges in

M. esculentoides

and

cryptica

. The hymenophore of

M. prava

often appears

like a contorted or somewhat deformed version of the

M. esculentoides

hymenophore. Significant differences

in microscopic features between

M. prava

M. esculen-

toides

and

M. cryptica

were not observed, although

elements on sterile ridges in the

M. prava

specimens

studied generally were sparsely scattered or nearly

absent, in contrast to the easily located elements of the

other two species.

Morchella prava

undoubtedly has

been misidentified as ‘‘

M. esculenta

’’ and ‘‘

M. deliciosa

’’

in many North American treatments, and it probably

corresponds to what has been labeled ‘‘

M. vulgaris

’’ in

Que´bec (Lincoff 1981). It appeared in Kuo (2005) as

the ‘‘Classic North American Morel III’’.

Morchella cryptica M. Kuo & J.D. Moore, sp. nov.

IG.7

MycoBank MB 563952

Ascomata 60–200 mm alta; capitulum ovoideum cum

apice late conico aut convexo; costae pallidae; hymenium

sufflavum aetate; biotopium in silva frondosa in meditulio

occidente Americae septentrionalis, alveis ubi

Fraxinus

americana

L praesens est; sporae 18–23 3 10–13 mm.

Holotypus: Biotopium in silva frondosa cum

Fraxinus

americana

Let

Acer

spp. praesentes; USA, in Illinoisense,

ad Coles County; M. Kuo col.; specimen typicum in Herb. F

(04220401) conservatum.

Etymology.

The epithet refers to the cryptic mor-

phology of the species.

Ascomata 60–200 mm high. Hymenophore 40–

75 mm high; 32–62 mm wide at the widest point;

usually ovoid with a convex, bluntly conical, or conical

apex; pitted and ridged; with 10–18 vertical ridges

intersecting a line across the widest portion but with

numerous horizontal and oblique ridges as well,

along with scattered, sunken, transecting ridges;

adnate at the point of attachment. Ridges glabrous

or nearly so; pale yellowish when young, usually

remaining pale yellowish with maturity but occasion-

ally darkening somewhat to brownish yellow; flattened

when young, becoming sharpened or eroded in age.

Pits often vertically elongated but not strictly so and

occasionally subglobose to irregular in outline;

glabrous; grayish yellow to gray or pale grayish brown

when young, becoming pale yellowish or pale

brownish yellow (usually concolorous with the ridges

at maturity). Stipe 50–130 mm high; 14–60 mm wide;

usually basally subclavate to clavate when mature but

occasionally more or less equal throughout develop-

ment; sometimes developing wide, shallow ridges;

finely mealy with whitish granules or nearly glabrous;

whitish. Context whitish; about 1–2 mm thick in the

hollow hymenophore; often becoming thickened and

chambered near the base. Sterile inner surface whitish

and pubescent. Ascospores 18–23 3 10–13(–15)

mm.

ellipsoid; smooth; contents homogeneous. Asci 175–

300 3 15–25

mm; eight-spored; cylindrical; hyaline.

Paraphyses 100–160 3 5–12.5(–15)

mm; cylindrical;

apices usually rounded or subclavate but occasionally

clavate or widely subfusiform; septate; hyaline to

brownish in KOH (2%). Elements on sterile ridges

scattered (occasionally difficult to locate or distinguish

from paraphyses); 75–125(–175) 3 10–30

mm; hyaline

in KOH (2%); septate; terminal cell widely cylindrical

with a rounded apex, subcapitate, capitate, subclavate,

clavate or widely subfusiform.

Ecology.

Appearing in Midwestern hardwood for-

ests, especially in apparent association with

F. ameri-

cana

but also reported under

L. tulipifera

and

Acer

spp.; fairly common in the Great Lakes region from

Ontario to central Illinois and western Pennsylvania;

April, May and June. Specimens examined (S

UPPLE-

MENTARY TABLE

I) were collected in Illinois, Michigan,

Ontario and Pennsylvania.

Comments. Morchella cryptica

corresponds to phylo-

genetic species

Mes

-11 in O’Donnell et al. (2011). Based

on current data the species cannot be reliably separated

from

M. esculentoides

on the basis of morphological

characters, although the hymenophore of

M. cryptica

frequently somewhat paler and its ridges are usually

more flattened. Microscopic features studied for the

two species are virtually identical.

M. cryptica

is not

found as frequently as

M. esculentoides

; however, both

species were collected under hardwoods, often in

apparent association with

Fraxinus

spp. Further collec-

tions of

M. cryptica

might lead to a reliable means of

separating these phylogenetic species. It undoubtedly

has been labeled ‘‘

M. esculenta

’’ in North American

treatments that cover the Great Lakes region (e.g.

Weber 1995). It corresponds with the ‘‘Classic North

American Yellow Morel II’’ in Kuo (2005).

Morchella Elata

Clade

The

Morchella elata

Clade (O’Donnell et al. 2011) is

sister to the

M. esculenta

Clade and corresponds with

the traditional morphological concept of ‘‘black

morels’’ (Kuo 2005) amended to include

M. semi-

libera

-like species and three species (

M. tomentosa

1166 MYCOLOGIA

frustrata

M. snyderi

) that can manifest pale,

esculenta

like coloration of the hymenophore. The North

American species in the

M. elata

Clade have pits that

primarily are elongated vertically at maturity and

generally feature a notable sinus where the hymeno-

phore attaches to the stipe.

Morchella tomentosa

M. Kuo, Mycotaxon 105:441.

2008. F

IG.8,SUPPLEMENTARY FIG.2

The original description of

Morchella tomentosa

appeared in Kuo 2008. (For an augmented description

based on material studied by the authors see S

UPPLE-

MENTARY MATERIALS

Comments. Morchella tomentosa

is a postfire morel

featuring densely tomentose surfaces. Because of the

dark, young ascomata it frequently has been called the

‘‘gray morel’’ by western commercial collectors. With

prolonged exposure to sunlight its colors often fade

dramatically, approximating those typical of

esculentoides

-like morels; these specimens, however,

still feature hairs that are brown in KOH (2%)and

thus can be identified microscopically. Stefani et al.

(2010) described an underground, root-like structure

beneath three Alaskan ascomata of

M. tomentosa

.Itis

unclear whether this structure, which they termed a

‘‘radiscisclerotium’’, is consistently attached to

tomentosa

ascocarps; it was not documented by any of

the collectors of the specimens we examined, and it

has not been reported by mycologists investigating the

species (McFarlane et al. 2005, Pilz et al. 2007) or by

commercial collectors for whom

M. tomentosa

is a

popular species. The three radiscisclerotia document-

ed by Stephani et al. (2010) extended to about 80 mm

into the substrate, branching several times; individual

branches of the structures were 5–15 mm thick.

Morchella tomentosa

has been treated as the ‘‘gray

morel’’ (Pilz et al. 2004, 2007; McFarlane et al. 2005)

and as the ‘‘Black Foot Morel’’ (Kuo 2005).

Morchella frustrata M. Kuo, sp. nov. F

IG.9

MycoBank MB 563953

Ascomata 60–90 mm alta; capitulum conicum; costae

minime spadice, non nigrescens; hymenium levis, minime

spadix; biotopium in silvis multigeneri; sporae 20–29 3

14–19

mm. Holotypus: Biotopium in silvis coniferibus; USA,

in Californiaense, ad Placer County; T. Bruns 3643 col.;

specimen typicum in Herb. UC (1860811) conservatum.

Etymology.

The epithet reflects the frustrating com-

bination of black and yellow morel features that

characterize the species.

Ascomata 60–90 mm high. Hymenophore 40–

60 mm high; 25–40 mm wide at the widest point;

conical; pitted and ridged; with 16–22 primary vertical

ridges and few shorter, secondary vertical ridges, with

frequent, sunken, transecting horizontal ridges; attached

to stipe with a sinus about 2–4 mm deep and 2–4 mm

wide. Ridges glabrous; pale yellowish to nearly whitish

when young, becoming pale tan with maturity; slightly

flattened when young but often becoming sharpened or

eroded with age. Pits primarily elongated vertically;

glabrous; dull grayish to pale yellowish or nearly whitish

when young, becoming pale tan to pale pinkish tan.

Stipe 20–40 mm high; 10–25 mm wide; more or less

equal or sometimes basally subclavate; glabrous or finely

mealy with whitish granules; whitish. Context whitish;

1–2 mm thick in the hollow hymenophore; in the stipe

sometimes slightly chambered near the base. Sterile

inner surface whitish and pubescent. Ascospores 20–29

3 14–19

mm; elliptical; smooth; contents homogeneous.

Asci 225–300 3 15–25

mm; eight-spored; cylindrical;

hyaline in KOH (2%). Paraphyses 100–225 3 10–25

mm;

cylindrical; apices rounded to subclavate or infrequently

subfusoid; septate; hyaline to brownish in KOH (2%).

Elements on sterile ridges 100–175 3 12.5–20

mm;

septate; terminal cell clavate or subclavate; hyaline or

with brownish contents in KOH (2%).

Ecology.

Appearing at various altitudes in mixed forests

dominated by various trees, including

Arbutus menziesii

Pursh,

Quercus

spp.,

Pseudotsuga menziesii

(Mirb.) Franco,

Pinus ponderosa

Laws.,

Pinus lambertiana

Dougl. and

Abies

concolor

(Gord. & Glend.) Lindl.; California and Oregon;

April. Specimens examined (S

UPPLEMENTARY TABLE I)

were collected in California and Oregon.

Comments. Morchella frustrata

corresponds to phy-

logenetic species

Mel

-2 in O’Donnell et al. (2011). The

species is one of a few North American members of the

M. elata

Clade with pale colors that can approximate

the ones traditionally associated with members of the

M. esculenta

Clade. However, despite its colors

frustrata

manifests the stature typical of black morels;

its hymenophore is conical, its pits are vertically

oriented and the hymenophore is attached to the

stipe with a notable sinus. Unlike

M. snyderi

,which

also can feature pale,

M. esculenta

-like colors,

frustrata

lacks a conspicuously lacunose stipe, its ridges

eventually darken in old age or upon drying and its

pits are glabrous rather than finely tomentose.

Ascospores of

M. frustrata

examined for the present

work were substantially smaller than those examined

for

M. snyderi

; analysis of further collections might

determine whether the difference in ascospore sizes is

consistent between the species.

Morchella frustrata

likely the same as the morel informally designated the

‘‘mountain blond morel’’ (Pilz et al. 2004, 2007).

Morchella punctipes

Peck, Bull. Torrey Bot. Cl. 30:99.

1903. F

IG.10

Holotypus.

USA, Michigan, Agricultural College, Iunius,

B. O. Longyear col.; in herbarium NYS (NYSF2511)

conservatum.

KUO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1167

Epitypus (hic designatus).

Biotopium sub

Liriodendron

tulipifera

Let

Fraxinus americana

L; USA, in Virginiaense, ad

Rappahannock County; T. Geho col.; specimen epitypicum

in herbarium F (05020502) conservatum.

Etymology.

Peck’s epithet refers to the surface of the

stipe, which often is punctated with mealy granules

that can darken with maturity; he used this feature to

distinguish

M. punctipes

from what we now know is the

strictly European

M. semilibera

, which according to his

concept, had a glabrous stipe.

Ascomata 25–180 mm high. Hymenophore 20–

45 mm high; 20–45 mm wide at the widest point;

broadly to narrowly conical or occasionally nearly

convex; pitted and ridged; with 14–26 primary vertical

ridges and infrequent shorter, secondary vertical

ridges and transecting horizontal ridges; attached in

a skirt-like manner to the stipe, roughly halfway from

the apex, with a sinus 8–20 mm deep. Ridges

glabrous; pale yellow to dull yellowish brown when

young; darkening to brown, dark brown, or black with

maturity; when young up to 1 mm wide, and flat with

sharp edges, but with age often rounded, sharp or

eroded. Pits vertically elongated; glabrous; whitish to

pale yellowish when immature, becoming brownish to

yellowish brown at maturity. Stipe 15–150 mm high;

8–45 mm wide; more or less equal, or tapered to apex;

often hidden by the hymenophore when young, but

lengthening dramatically by maturity; often develop-

ing shallow longitudinal furrows; fragile; in warm, wet

conditions sometimes becoming inflated, especially

near the base; white to whitish or watery brownish;

occasionally nearly glabrous but more commonly

mealy with whitish granules that sometimes darken

to brownish or brown. Context whitish to watery tan;

1–2 mm thick in the hollow hymenophore; in the

stipe sometimes chambered or layered near the base.

Sterile inner surface whitish to brownish; mealy;

sometimes grooved. Ascospores elliptical; smooth;

contents homogeneous; 20–27 3 (10–)14–18

mm;

whitish to bright yellowish orange in deposit. Asci

eight-spored; 175–350 3 15–25

mm; cylindrical;

hyaline. Paraphyses cylindrical with rounded, subca-

pitate, clavate, mucronate or irregularly inflated

apices; septate; hyaline in KOH (2%); 120–275 3

10–22

mm. Elements on sterile ridges 50–100 3

10–25

mm; septate; tightly packed in an even layer;

brownish in KOH (2%); terminal cell broadly clavate

to sub-rectangular with a flattened to broadly round-

ed or submucronate apex.

Ecology.

AppearingineasternNorthAmerican

hardwood forests, especially those containing

americana

L. tulipifera

and

U. americana

; widely

distributed east of the Rocky Mountains (although

Weber and Smith 1985 reported that it is ‘‘to be

expected primarily in the northern and montane parts

of the South’’); from late March in southern areas to

late May in northern areas. Specimens examined

UPPLEMENTARY TABLE I) were collected in Illinois,

Michigan, Missouri, Pennsylvania and Virginia.

Comments. Morchella punctipes

corresponds to phy-

logenetic species

Mel

-4 in O’Donnell et al. (2011).

Most North American treatments (e.g. Overholts 1934,

Weber 1995) regard the present species as ‘‘

Morchella

semilibera

DC,’’ distinct from other morels on the basis

of its ‘‘half-free’’ cap attachment. However, results

from O’Donnell et al. (2011) support two

semilibera

like morels in North America (species

Mel

-4 and

species

Mel

-5), morphologically similar but clearly

separated on the basis of their range and ecology.

Morchella populiphila

(phylogenetic species

Mel

-5 in

O’Donnell et al. 2011) is a western species associated

with

Populus trichocarpa

Torr. & Gray; the eastern

species (phylogenetic species

Mel

-4 in O’Donnell et al.

2011) is widely distributed in hardwood forests east of

the Rocky Mountains. Neither is the same as the

European species (phylogenetic species

Mel

-3 in

O’Donnell et al. 2011),

M. semilibera

. Although we

were unable to obtain informative phylogenetic results

from the holotype of

M. punctipes

Peck, Peck’s

description (1903) and the morphology of the

holotype specimens (see

M. punctipes

, type studies)

match the morphology of the specimens examined for

the present work, which are conspecific with phyloge-

netic species

Mel

-4 in O’Donnell et al. (2011).

Morchella populiphila M. Kuo, M.C. Carter & J.D.

Moore, sp. nov. F

IG.11

MycoBank MB 563954

Ascomata 40–150 mm alta; capitulum conicum, remisse

adherum stipiti; costae perpendicules, atrae; hymenium

fulvum; biotopium in alveis occidentis Americae septen-

trionalis sub

Populus trichocarpa

Torr. & Gray; sporae 20–25

3 12–16

mm. Holotypus: Biotopium in alveo sub

Populus

trichocarpa

Torr. & Gray; USA, in Oregonense, ad Jackson

County; N. Selbicky col.; specimen typicum in Herb. F

(03240401) conservatum.

Etymology.

The epithet refers to the association with

Populus trichocarpa

Ascomata 40–150 mm high. Hymenophore 20–

50 mm high; 20–50 mm wide at the widest point;

broadly to narrowly conical; pitted and ridged, with

12–20 primary vertical ridges and infrequent shorter,

secondary vertical ridges and transecting horizontal

ridges, attached in a skirt-like manner to the stipe,

roughly halfway from the apex, with a sinus 10–25 mm

deep. Ridges glabrous; yellowish brown to honey

brown when young, darkening to brown, dark brown

or black with maturity; when young up to 1 mm wide

and flat with sharp edges but often rounded, sharp or

eroded in age. Pits vertically elongated; glabrous;

1168 M

YCOLOGIA

FIGS. 11–19.

Morchella

species. 11.

M. populiphila

HOLOTYPE F 03240401,

Mel

-5. 12.

M. sextelata

HOLOTYPE F 07130403,

Mel

-6. 13.

M. septimelata

HOLOTYPE F 06150404,

Mel

-7. 14.

M. capitata

HOLOTYPE F 08070406,

Mel

-9. 15.

M. importuna

HOLOTYPE F 04130401,

Mel

-10. 16.

M. snyderi

F 05140401,

Mel

-12. 17.

M. angusticeps

EPITYPE F 04090601,

Mel

-15. 18.

brunnea

F 04100401,

Mel

-22. 19.

M. septentrionalis

F 05110306,

Mel

-24. Bars 5 5 cm.

Mel

numbers refer to the phylogenetic

species reported in O’Donnell et al. (2011).

KUO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1169

whitish to pale brown when immature, becoming

brownish to yellowish or grayish brown at maturity.

Stipe 25–110 mm high; 10–50 mm wide; more or less

equal, or tapered to apex; often hidden by the

hymenophore when young but lengthening dramatically

with maturity; often developing shallow longitudinal

furrows; fragile; in warm, wet conditions sometimes

becoming inflated, especially near the base; white to

whitish or watery brownish; occasionally nearly glabrous

but more commonly mealy with whitish granules that

sometimes darken to brownish or brown. Context

whitish to watery tan; 1–2 mm thick in the hollow

hymenophore; sometimes chambered or layered near

the base; fragile. Sterile inner surface whitish to

brownish; mealy. Ascospores 20–25(–29) 3 12–16(–18)

mm; elliptical; smooth; contents homogeneous; bright

yellowish orange in deposit. Asci 225–325 3 15–22.5

mm;

eight-spored; cylindrical; hyaline. Paraphyses 150–275 3

7–15

mm; cylindrical with subclavate, clavate or subcapi-

tate apices; septate; hyaline in KOH (2%). Elements on

sterile ridges 100–175 3 10–25

mm; septate; tightly

packed in an even layer; brownish to brown in KOH

(2%); terminal cell broadly clavate to sub-rectangular

with a flattened to broadly rounded apex.

Ecology.

Occurring under

P. trichocarpa

in western

North American river bottoms; distributed from

Oregon to Nevada and northern California; March

and April. Specimens examined (S

UPPLEMENTARY TABLE

I) were collected in California, Nevada and Oregon.

Comments. Morchella populiphila

corresponds to

phylogenetic species

Mel

-5 in O’Donnell et al.

(2011). Western North American field guides (e.g.

Smith 1975, Arora 1986) regard this species as

‘‘

Morchella semilibera

DC’’, distinct from other

morels on the basis of its ‘‘half-free’’ hymenophore

attachment, and identical to the half-free morel of

eastern North America, which is phylogenetically

distinct and is reported as

M. punctipes

Peck in the

present work. On the basis of the specimens

examined, comments in western field guides and

the Seattle area collecting experience of one of us

(Moore), the association with

P. trichocarpa

consistent; the range of

M. populiphila

may corre-

spond to the range of

P. trichocarpa

Morch ella

populiphila

was featured as one of two ‘‘North

American Half-Free Morels’’ in Kuo (2005).

Morchella sextelata M. Kuo, sp. nov. F

IG.12

MycoBank MB 563955

Ascomata 40–105 mm alta; capitulum conicum; costae

brunneae, interdum nigrescens; hymenium minime bru-

num usque minime puniceum; biotopium in silvis conifer-

ibus incensis in occidentali America septentrionalis; sporae

18–25 3 10–16

mm. Holotypus: Biotopium in silvis conifer-

ibus incensis; USA, in Montanaense, ad Missoula County; S.

Engstrom col.; specimen typicum in Herb. F (07130403)

conservatum.

Etymology.

The epithet reflects the fact that this is

the sixth phylogenetic species (

Mel

-6) in the

M. elata

Clade enumerated in O’Donnell et al. (2011).

Ascomata 40–105 mm high. Hymenophore 25–

75 mm high; 20–50 mm wide at the widest point;

conical to widely conical; pitted and ridged, with

12–20 primary vertical ridges and numerous shorter,

secondary vertical ridges and sunken transecting

horizontal ridges; attached to stipe with a sinus about

2–4 mm deep and 2–4 mm wide. Ridges glabrous or

finely tomentose; pallid when young; becoming dark

grayish brown to nearly black with maturity; bluntly

flattened when young, sometimes becoming sharp-

ened or eroded with age. Pits primarily vertically

elongated; glabrous; dull brownish to yellowish tan,

pinkish, or nearly buff. Stipe 20–50 mm high; 10–

22 mm wide; more or less equal or sometimes basally

subclavate; glabrous or finely mealy with whitish

granules; whitish. Context whitish; 1–2 mm thick in

the hollow hymenophore; sometimes slightly cham-

bered near the base. Sterile inner surface whitish and

pubescent. Ascospores 18–25 3 10–16(–22)

mm;

elliptical; smooth; contents homogeneous. Asci 200–

325 3 5–25

mm; eight-spored; cylindrical; hyaline.

Paraphyses 175–300 3 2–15

mm; cylindrical with

rounded, subacute, subclavate or subfusoid apices;

septate; hyaline in KOH (2%). Elements on sterile

ridges 50–180 3 5–25

mm; septate; terminal cell

cylindrical with a rounded apex, subfusoid, subcapi-

tate or block-like; with brown to brownish contents in

KOH (2%).

Ecology.

Appearing at 1000–1500 m in lightly to

moderately burned conifer forests, including forests

dominated by

Ps. menziesii

and

P. ponderosa

.Found

primarily in years immediately following forest fires but

often appearing in dwindling numbers for several

seasons thereafter; Washington, Idaho, Montana, Wyo-

ming and Yukon Territory; April–July. Specimens exam-

ined (S

UPPLEMENTARY TABLE I) were collected in Idaho,

Montana, Washington, Wyoming and Yukon Territory.

Comments. Morchella sextelata

corresponds to phy-

logenetic species

Mel

-6 in O’Donnell et al. (2011).

From a strictly morphological perspective the species is

virtually identical to several members of the

M. elata

Clade (

M. septimelata

M. brunnea

M. angusticeps

septentrionalis

), but because it apparently is limited to

conifer burn sites in western North America it can be

easily separated from all but

M. septimelata

,from

which it is morphologically and ecologically indistin-

guishable on the basis of currently available data.

Elements on sterile ridges in the latter species were

primarily subclavate to clavate in the specimens

examined, while elements in

M. sextelata

were

1170 M

YCOLOGIA

cylindrical with a rounded apex, subfusoid, subcapitate

or block-like, but this distinction is too tentative and

based on too few specimens examined for us to

express confidence that the difference is consistent.

Because several of the collections studied for the

present work (F 07130403, F 07070405) had pinkish

pits,

M. sextelata

probably was included in the concept

of the ‘‘pink morel’’ set forth in Pilz et al. (2004,

2007); however,

M. septimelata

specimens also dem-

onstrated pinkish pits.

Morchella sextelata

was treated

in Kuo (2005) as one of several ‘‘Other North

American Black Morels’’ appearing in burn sites.

Morchella septimelata M. Kuo, sp. nov. F

IG.13

MycoBank MB 563956

Ascomata 75–200 mm alta; capitulum conicum; costae

brunneae, interdum nigrescens; hymenium minime bru-

num usque leviter veridis aut minime puniceum; biotopium

in silvis coniferibus incensis in occidenti America septen-

trionalis; sporae 18–25 3 10–15

mm. Holotypus: Biotopium

in silvis coniferibus incensis; USA, in Montanaense, ad

Mineral County; S. Engstrom col.; specimen typicum in

Herb. F (06150404) conservatum.

Etymology.

The epithet reflects the fact that the spe-

cies is the seventh phylogenetic species (

Mel

-7) in the

M. elata

Clade enumerated in O’Donnell et al. (2011).

Ascomata 75–200 mm high. Hymenophore 40–

100 mm high; 30–70 mm wide at the widest point;

conical to subconical; pitted and ridged, with 14–22

primary vertical ridges and numerous shorter, sec-

ondary vertical ridges and transecting horizontal

ridges; attached to stipe with a sinus about 1–3 mm

deep and 1–3 mm wide. Ridges glabrous or finely

tomentose; brownish to brown when young, becom-

ing dark brown to black at maturity; bluntly flattened

when young but sometimes becoming sharpened or

eroded with age. Pits primarily vertically elongated;

glabrous; progressing from olive to olive-brown,

pinkish or brownish when immature to brownish or

tan at maturity. Stipe 35–100 mm high; 20–50 mm

wide; occasionally basally subclavate; flared slightly to

apex; mealy with whitish granules; white, darkening

to brownish in older specimens. Context whitish;

1–2 mm thick in the hollow hymenophore; sometimes

slightly chambered near the base. Sterile inner

surface whitish and pubescent. Ascospores (17–)18–

25(–30) 3 10–15(–20)

mm; elliptical; smooth; contents

homogeneous. Asci 175–275 3 12–25

mm; eight-

spored; cylindrical; hyaline. Paraphyses 100–200 3

5–12.5

mm; cylindrical; apices subacute or subfusiform

(occasionally merely rounded, or subclavate); septate;

hyaline in KOH (2%). Elements on sterile ridges

60–200 3 7–18

mm; septate; with brown contents in

KOH (2%); cylindrical; terminal cell clavate (rarely

subcapitate or irregular).

Ecology.

Appearing at 1000–2000 m in lightly to

moderately burned conifer forests, often near creek

beds, springs and seeps. Found primarily in years

immediately following forest fires but often appearing

in dwindling numbers for several seasons thereafter;

April–July. Specimens examined (S

UPPLEMENTARY

TABLE

I) were collected in British Columbia, Montana

and Oregon.

Comments. Morchella septimelata

corresponds to phy-

logenetic species

Mel

-7 in O’Donnell et al. (2011). On

the basis of present data the species is morphologically

and ecologically indistinguishable from

M. sextelata

(See the comments for the latter species for further

information and comparison with other

elata

-like spe-

cies.) Because several of the

M. septimelata

collections

studied had greenish pits (F 06150404, F 07140404,

F 07070401, F 07070408) or pinkish pits (F 07070408),

M. septimelata

probably has been included in commer-

cial collectors’ concept of the ‘‘pickle’’, and in the

concepts of the ‘‘pink morel’’ and the ‘‘green morel’’ set

forth in Pilz et al. (2004, 2007); however, several

capitata

specimens also had greenish pits and specimens

M. sextelata

also had pinkish pits.

Morchella septimelata

was treated in Kuo (2005) as one of several ‘‘Other

North American Black Morels’’ appearing in burn sites.

Morchella

sp.

Mel

-8

Commentary and a description of the sole ascoma

representing this phylogenetic species for the present

work are included (S

UPPLEMENTARY MATERIALS). A

formal description is deferred pending identification

of further specimens.

Morchella capitata M. Kuo & M.C. Carter, sp. nov.

IG. 14, SUPPLEMENTARY FIG.4

MycoBank MB 563957

Ascomata 60–110 mm alta; capitulum conicum vel

subconicum; costae brunneae, interdum nigrescens; hyme-

nium aerugineum vel fulvum; biotipium in silvis coniferibus

incensis; sporae 18–25 3 12–17

mm; elementa in costis

sterilibus capitata. Holotypus: Biotopium in silvis conifer-

ibus incensis; 1200 m. altitudinis; USA, in Oregonense, ad

Jefferson County; M.C. Carter col.; specimen typicum in

Herb. F (08070406) conservatum.

Etymology.

The epithet refers to the capitate ele-

ments on the sterile ridges.

Ascomata 60–110 mm high. Hymenophore 40–

80 mm high; 25–80 mm wide at the widest point;

conical to subconical (occasionally subglobose);

pitted and ridged, with 18–28 primary vertical ridges

and numerous shorter, secondary vertical ridges,

often with prominent and regular transecting hori-

zontal ridges; attached to stipe with a sinus about

1–7 mm deep and 1–7 mm wide. Ridges glabrous or

finely tomentose; dull olive, brown, brownish black or

UO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1171

brownish when young, becoming dark brown to black

at maturity; somewhat flattened when young but with

age sometimes becoming sharpened or eroded. Pits

primarily vertically elongated; glabrous; progressing

from olive to olive brown or brownish when imma-

foture to brownish or tan at maturity. Stipe 25–50 mm

high; 20–50 mm wide; occasionally basally subclavate;

often mealy with whitish granules; whitish, darkening

to brownish in older specimens. Context whitish;

1–2 mm thick in the hollow hymenophore; in the

stipe sometimes layered or chambered, especially

near the base. Sterile inner surface whitish and pu-

bescent. Ascospores 18–25 3 12–17(–19)

mm; ellip-

tical; smooth; contents homogeneous. Asci 175–300

3 15–27

mm; eight-spored; cylindrical; hyaline in

KOH (2%). Paraphyses 125–225 3 7.5–13

mm; cylin-

drical; septate; apices subclavate, subfusiform or

merely rounded (rarely subcapitate); hyaline to

faintly brownish in KOH (2%). Elements on sterile

ridges 75–200 3 10–35 +

mm; septate; brown in KOH

(2%); terminal cell usually subcapitate, capitate or

strongly swollen and subglobose but occasionally

clavate or merely cylindrical with a rounded apex.

Ecology.

Appearing at approximately 1200 m in lightly

burned spruce and fir forests. Found in the year

following summer forest fires; August. Specimens exam-

ined (S

UPPLEMENTARY TABLE I) were collected in Oregon.

Comments. Morchella capitata

corresponds to phylo-

genetic species

Mel

-9 in O’Donnell et al. (2011). Among

the confusingly similar burn-site morels in western North

America,

Morchella capitata

is distinct microscopically,

based on the specimens examined; elements on its sterile

ridges are overwhelmingly capitate, often dramatically so

UPPLEMENTARY FIG. 4). Specimens we studied of

sextelata

and

M. septimelata

, which are nearly identical to

M. capitata

in macroscopic features and which also occur

in conifer burn sites, had sterile ridge elements from

cylindrical with a rounded apex to subclavate, subfusi-

form or subcapitate. Because several of the collections

studied (08070401, 08070408) had greenish pits,

capitata

probably has been included in commercial

collectors’ concept of the ‘‘pickle’’ and in the concept of

the ‘‘green morel’’ set forth in Pilz et al. (2004, 2007);

however, several

M. sextelata

and

M. septimelata

speci-

mens also had greenish pits.

Morchella capitata

was

treated in Kuo (2005) as one of several ‘‘Other North

American Black Morels’’ appearing in burn sites.

Morchella importuna M. Kuo, O’Donnell & T.J. Volk,

sp. nov. F

IG.15

MycoBank MB 563958

Ascomata 60–200 mm alta; capitulum conicum vel

ovatum; costae aequabiles, perpendicules, atrae, similes

scalae; hymenium fulvum; biotopium in hortis et terra

perturbata; sporae 18–24 3 10–13

mm. Holotypus: Bioto-

pium in horto; USA, in Washingtonense, ad King County;

A. Thrailkill col.; specimen typicum in Herb. F (04130401)

conservatum.

Etymology.

The epithet means ‘‘assertive’’ or ‘‘in-

considerate’’ in character; the species often is the

cause of consternation and distress among gardeners

and homeowners whose territory has been invaded.

Ascomata 60–200 mm high. Hymenophore 30–

150 mm high; 20–90 mm wide at the widest point;

conical to widely conical or occasionally ovoid; pitted

and ridged, with 12–20 primary vertical ridges and

numerous transecting horizontal ridges, creating a

laddered appearance; attached to stipe with a sinus

about 2–5 mm deep and 2–5 mm wide. Ridges

glabrous or finely tomentose; pale to dark gray when

young, becoming dark grayish brown to nearly black

with maturity; bluntly rounded when young, but with

age becoming sharpened or eroded. Pits vertically

elongated in all stages of development; glabrous or

finely tomentose; opening and deepening with

development; progressing from gray to dark gray

when immature to grayish brown, grayish olive or

brownish yellow at maturity. Stipe 30–100 mm high;

20–60 mm wide; often basally clavate to subclavate;

glabrous or finely mealy with whitish granules;

developing longitudinal ridges and channels with

maturity, especially basally; whitish to pale brownish.

Context whitish to watery tan; 1–3 mm thick in the

hollow hymenophore; in the stipe sometimes cham-

bered or layered. Sterile inner surface whitish and

pubescent. Ascospores 18–24 3 10–13

mm; elliptical;

smooth; contents homogeneous. Asci 220–300 3

12–25

mm; eight-spored; cylindrical; hyaline. Paraphy-

ses 150–250 3 7–15

mm; septate; cylindrical with

rounded to subclavate, subcapitate, subacute or

subfusoid apices; hyaline or brownish in KOH (2%).

Elements on sterile ridges 125–300 3 10–30

mm;

septate; terminal cell cylindrical with a rounded apex,

subclavate, clavate, subcapitate or subfusiform; hya-

line or brownish to brown in KOH (2%).

Ecology.

Appearing in gardens, planters, woodchip

beds and urban landscaping settings in the Pacific

Northwest and northern California; March–May. Spec-

imens examined (S

UPPLEMENTARY TABLE I) were

collected in British Columbia, California, Nevada,

Oregon and Washington.

Comments. Morchella importuna

corresponds to phy-

logenetic species

Mel

-10 in O’Donnell et al. (2010).

The species is distinguished from other morels in the

M. elata

Clade on the basis of its regularly laddered,

vertically oriented pits and ridges, combined with its

urban habitat in landscaping areas, planters, woodchip

beds and gardens, primarily in western North America.

One of us (Volk) collected morels matching the

morphology of

M. importuna

in a woodchip bed on

1172 M

YCOLOGIA

the campus of the University of Wisconsin at La Crosse

in 1999.

Morchella hotsonii

Snyder (so far known only

from the 1935 type collection) is similar and also

apparently appeared in woodchip beds in the Pacific

Northwest, although it differs morphologically by

having finely tomentose surfaces (S

UPPLEMENTARY

MATERIALS

, Type studies, for further information).

Morchella importuna

was treated and illustrated erro-

neously in Kuo (2005) as conspecific with the ‘‘Classic

North American Black Morel’’; the top-middle, top-

right and bottom-middle photos on p 179 actually

represent

M. importuna

Morchella snyderi M. Kuo & Methven, sp. nov.

IG.16

MycoBank MB 563959

Ascomata 60–140 mm alta; capitulum conicum; costae

pallidae, brunnescens aetate; hymenium sufflavum vel

fulvum; stipes lacunosus aetate; biotopium in silvis conifer-

ibus in occidenti America septentrionalis; sporae 25–37 3

15–23

mm. Holotypus: Biotopium in silvis coniferibus; USA,

in Idahoense, ad Kootenai County; N. S. Weber 6554 col.;

specimen typicum in Herb. OSC (139277) conservatum.

Etymology.

The epithet honors Leon Carlton Snyder

(1908–1987), who named

M. hotsonii

and

M. crassis-

tipa

from Washington state in the 1930s.

Ascomata 60–140 mm high. Hymenophore 35–

80 mm high; 30–50 mm wide at the widest point;

conical; pitted and ridged, with 16–22 primary vertical

ridges and frequent shorter, secondary vertical ridges,

with occasional sunken, transecting horizontal ridges;

attached to stipe with a sinus about 2–4 mm deep and

2–4 mm wide. Ridges glabrous or very finely tomen-

tose; pale yellowish, becoming pale tan, then grayish

brown with maturity and darkening to nearly black

when dried; flattened when young but with age

sometimes becoming sharpened or eroded. Pits more

or less vertically elongated, at least at maturity; finely

tomentose; yellowish when young, becoming pale tan

to pale grayish brown. Stipe 35–70 mm high; 25–

40 mm wide; more or less equal, or sometimes basally

subclavate; at first finely mealy with whitish granules,

becoming prominently granulated; whitish to pale

brownish; usually becoming prominently ridged and/

or lacunose with maturity. Context whitish; 1–2 mm

thick in the hollow hymenophore, becoming layered

and chambered, especially near the base of the stipe.

Sterile inner surface whitish and pubescent. Asco-

spores 25–37 3 15–23

mm; elliptical; smooth; contents

homogeneous. Asci 225–300 3 17.5–32.5

mm; eight-

spored; cylindrical; hyaline in KOH (2%). Paraphyses

100–200 3 7.5–20

mm; cylindrical; apices rounded to

subclavate, clavate, or occasionally subcapitate or

widely fusiform; septate; hyaline to faintly brownish

in KOH (2%). Elements on sterile ridges 75–175 3

10–20

mm; septate; terminal cell subclavate, clavate,

subcapitate or widely fusiform; with hyaline to

brownish or brown contents in KOH (2%).

Ecology.

Appearing under non-burned, montane

conifers, including

Ps. menziesii

P. ponderosa

and

concolor

; April, May and early June. Specimens

examined (S

UPPLEMENTARY TABLE I) were collected in

California, Idaho, Montana, Oregon and Washington.

Comments. Morchella snyderi

corresponds to phylo-

genetic species

Mel

-12 in O’Donnell et al. (2011). A

combination of ecological and morphological features

separates the species: it appears in non-burned conifer

forests in western North America; its stipe is often

ridged and lacunose (although the stipes of many

morels can become ridged and sublacunose, especially

near the base, when mature, the stipe of

M. snyderi

often prominently lacunose and ridged throughout

most of its development) and prominently granulated;

the colors of the hymenophore approximate those

esculenta

-like morels when young, but the ridges

become smoky brown to black with development or

upon drying; its hymenophore is conical, with longitu-

dinally arranged pits and a sinus at the point of

attachment to the stipe; and its ascospores are

comparatively large. Young specimens of

M. snyderi

can appear similar to

M. frustrata

,butthelatterspecies

hasridgesthatdonotdarkenandhassmaller

ascospores.

Morchella brunnea

also is similar, but its

hymenophore is more brown when young and its stipe

is not regularly lacunose. We found one ascoma

fragment in the putative holotype of

M. crassistipa

Snyder to be phylogenetically conspecific with

snyderi

, but we were unable to recognize Snyder’s name

because the type collection was mixed as to species (see

UPPLEMENTARY MATERIALS, Type studies, for full discus-

sion).

Morchella snyderi

was not treated in Kuo (2005).

Morchella angusticeps

Peck, Ann. Rep. New York St.

Mus. 32:44. 1879. F

IG. 17, SUPPLEMENTARY FIG.5

Holotypus.

USA, New York, West Albany et Center,

Aprilis et Maius, C. H. Peck col.; in herbarium NYS

(NYSF268) conservatum.

Epitypus (hic designatus).

Biotopium sub

Fraxinus

americana

; USA, in Illinoisense, ad Coles County; M. Kuo

col.; specimen epitypicum in herbarium F (04090601)

conservatum.

Etymology.

Peck’s epithet refers to the narrow

hymenophore.

Ascomata 50–140 mm high. Hymenophore 30–

80 mm high; 25–50 mm wide at the widest point;

conical or bluntly conical; pitted and ridged, with

16–24 primary vertical ridges and occasional shorter,

secondary vertical ridges, with frequent sunken,

transecting horizontal ridges; attached to stipe with

a sinus about 2–5 mm deep and 2–5 mm wide

UO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1173

(however, at maturity the stipe sometimes becomes

swollen, obscuring the sinus). Ridges finely tomen-

tose; tan to dark brown, blackening with maturity;

flattened when young but with age sometimes

becoming sharpened or eroded. Pits primarily verti-

cally elongated; glabrous; pale tan to dull brownish

yellow (occasionally with olive shades). Stipe 20–

80 mm high; 10–30 mm wide; more or less equal or

sometimes basally subclavate to clavate; finely mealy

with whitish granules; whitish to pale brownish;

developing folds and channels, especially near the

base, at maturity; in warm, wet conditions sometimes

becoming swollen to reach, or nearly reach, the width

of the hymenophore, obscuring the sinus at the point

of attachment. Context whitish; 1–2 mm thick in the

hollow hymenophore; in the stipe becoming layered

and chambered near the base. Sterile inner surface

whitish and pubescent. Ascospores 22–27 3 11–15

mm;

elliptical; smooth; contents homogeneous. Asci 225–

400 3 17.5–30

mm; eight-spored; cylindrical; hyaline

in KOH (2%). Paraphyses 125–250 3 5–12.5

mm;

septate; cylindrical with a rounded, subclavate, cla-

vate, subcapitate, or occasionally widely fusiform

apex; hyaline in KOH (2%). Elements on sterile

ridges 100–200 3 7.5–35

mm; septate; terminal cell

widely cylindrical with a rounded apex, clavate,

subcapitate or occasionally irregular; with hyaline to

brownish or brown contents in KOH (2%).

Ecology.

Appearing under diverse hardwoods, espe-

cially

F. americana

and

L. tulipifera

; widely distributed

east of the Rocky Mountains; March, April and May.

Specimens examined (S

UPPLEMENTARY TABLE I) were

collected in Arkansas, Illinois, Massachusetts, Missis-

sippi, Pennsylvania and West Virginia.

Comments. Morchella angusticeps

corresponds to

phylogenetic species

Mel

-15 in O’Donnell et al.

(2011). Results from O’Donnell et al. (2011) support

two

elata

-like morels in North America east of the

Rocky Mountains. One of these (

Mel

-24 in O’Donnell

et al. 2011) is

Morchella septentrionalis

, characterized

by its small size (ascomata 40–75 mm high), small

ascospores ([19–]20–22[–25] 3 11–15

mm), northern

distribution (from about 44uN northward) and fre-

quent association with the deadwood of hardwoods

(especially that of

P. grandidentata

); the second (

Mel

15 in O’Donnell et al. 2011) has larger ascomata and

ascospores and is widely distributed east of the Rocky

Mountains. Although we were unable to obtain

informative phylogenetic results from the holotype of

M. angusticeps

, Peck’s description and illustration

(1879) and the macromorphology of the holotype

roughly match the specimens examined and cited

above, which are phylogenetically consistent with

Mel

15 in O’Donnell et al. (2011). However, there are

several potentially unresolved complications presented

by the holotype and by Peck’s description (see

UPPLEMENTARY MATERIALS, type studies). From a

strictly morphological perspective

M. angusticeps

virtually indistinguishable from several western North

American species, including

M. sextelata

M. septime-

lata

and

Morchella

sp.

Mel

-8 (all of which occur in

conifer burn sites), as well as

M. brunnea

, which

appears in unburned forests.

Morchella angusticeps

has

been treated as

M. angusticeps

by many authors (e.g.

Seaver 1928, Overholts 1934, Weber 1995) who

probably combined it with

M. septentrionalis

.It

appeared as the ‘‘Classic North American Black

Morel’’ in Kuo (2005), where it was combined

erroneously with

M. importuna

.Wehaveselecteda

robust, contemporary collection (F 04090601) as an

epitype representative of our concept of the phyloge-

netic species we are labeling

M. angusticeps

Morchella brunnea M. Kuo, sp. nov. F

IG.18

MycoBank MB 563960

Ascomata 45–90 mm alta; capitulum conicum; costae

fulvum, nigrescentes aetate; hymenium fulvum, subtiliter

tomenteum; biotopium in silva in occidenti America

septentrionalis; sporae 22–36 3 14–20

mm. Holotpus:

Biotopium in silvis; USA, in Oregonense, ad Jefferson

County; N.S. Weber 6199 col.; specimen typicum in Herb.

OSC (138686) conservatum.

Etymology.

The epithet refers to the brown hyme-

nophore.

Ascomata 45–90 mm high. Hymenophore 30–

50 mm high; 25–35 mm wide at the widest point;

conical; pitted and ridged, with 16–22 primary vertical

ridges and occasional shorter, secondary vertical

ridges, with frequent sunken, transecting horizontal

ridges; attached to stipe with a sinus about 2–3 mm

deep and 2–3 mm wide. Ridges glabrous or finely

tomentose; dark brown to nearly black, blackening

with maturity and when dried; flattened when young

but sometimes becoming sharpened or eroded with

age. Pits primarily vertically elongated; finely tomen-

tose; pale tan to brownish yellow. Stipe 20–35 mm

high; 8–15 mm wide; more or less equal, or

sometimes basally subclavate; finely mealy with

whitish granules; whitish; developing fine ridges, or

a few folds near the base, with maturity. Context

whitish; 1–2 mm thick in the hollow hymenophore,

becoming layered and chambered near the base of

the stipe. Sterile inner surface whitish and pubescent.

Ascospores 22–36(–40) 3 14–20(–25)

mm; elliptical;

smooth; contents homogeneous. Asci 225–300 3

17.5–22.5

mm; eight-spored; cylindrical; hyaline in

KOH (2%). Paraphyses 150–250 3 7.5–17.5

mm;

cylindrical; apices rounded to subclavate, clavate, or

widely fusiform; septate; hyaline in KOH (2%).

Elements on sterile ridges 75–160 3 12.5–27.5

mm;

1174 M

YCOLOGIA

septate; terminal cell clavate (sometimes strikingly

so), subcapitate or widely subfusiform; with hyaline to

brownish contents in KOH (2%).

Ecology.

Appearing under hardwoods, including

menziesii

and

Quercus

spp.; probably also to be

expected in non-burned conifer forests; April. Speci-

mens examined (S

UPPLEMENTARY TABLE I) were col-

lected in Oregon.

Comments. Morchella brunnea

corresponds to phy-

logenetic species

Mel

-22 in O’Donnell et al. (2011).

The species is morphologically similar to several

morels in the

M. elata

Clade (including

M. sextelata

M. septimelata

M. capitata

Morchella

sp.

Mel

-8,

angusticeps

M. septentrionalis

) but can be differenti-

ated when its ecology (in non-burned forests) and

distribution (in western North America) are consid-

ered. Among the species that are similar in appearance

M. brunnea

, only the poorly known

Morchella

sp.

Mel

-8 apparently inhabits similar western habitats.

The sole specimen we examined for

Morch ella

sp.

Mel

-8 had glabrous pits, as compared to the finely

tomentose pits of

M. brunnea

specimens examined;

further collections of both phylogenetic species may

determine whether this difference is consistent.

Morch ella brunnea

is mentioned as

M. angusticeps

in Weber (1995) and is treated as the ‘‘natural black

morel’’ in Pilz et al. (2004, 2007). It was treated as

one of several ‘‘Classic North American Black

Morels’’ in Kuo (2005).

Morchella septentrionalis M. Kuo, J.D. Moore &

Zordani, sp. nov. F

IG.19

MycoBank MB 563961

Ascomata 40–75 mm alta; capitulum conicum vel sub-

clavatum; costae fuscae, nigrescentes aetate; hymenium

fulvum usque fuscum; biotopium in silva frondosa in alta

America septentrionalis; sporae 20–22 3 11–15

mm. Holo-

typus: Biotopium sub

Populus grandidentata

Michx.; USA,

in Michiganense, ad Cheboygan County; R. Zordani col.;

specimen typicum in Herb. F (05110405) conservatum.

Etymology.

The epithet reflects the northern distri-

bution of the species.

Ascomata 40–75 mm high. Hymenophore 30–

45 mm high; 15–25 mm wide at the widest point;

conical, bluntly conical, or subclavate; pitted and

ridged, with 14–20 primary vertical ridges and

occasional shorter, secondary vertical ridges, with

occasional or frequent sunken, transecting horizontal

ridges; attached to stipe with a sinus about 2–3 mm

deep and 2–3 mm wide. Ridges finely tomentose or

nearly glabrous; tan to brown or dark brown,

blackening with maturity; flattened when young but

with age sometimes becoming sharpened or eroded.

Pits primarily vertically elongated; glabrous; pale tan

to dull brownish yellow or grayish brown. Stipe 20–

30 mm high; 8–15 mm wide; more or less equal or

sometimes basally subclavate; finely mealy with

whitish granules; whitish to pale brownish; sometimes

developing a few folds near the base with maturity.

Context whitish; up to about 1 mm thick. Sterile inner

surface whitish and pubescent. Ascospores (19–)20–

22(–25) 3 11–15

mm; elliptical; smooth; contents

homogeneous. Asci 225–350 3 15–30

mm; eight-

spored; cylindrical; hyaline in KOH (2%). Paraphyses

75–225 3 7.5–17.5

mm; septate; cylindrical with a

rounded, subacute, subclavate, clavate, subfusiform,

sublecythiform, subcapitate or occasionally irregular

apex; hyaline in KOH (2%). Elements on sterile

ridges 75–175 3 7.5–25

mm; septate; terminal cell

widely cylindrical with a rounded apex, subclavate,

clavate, widely fusiform, utriform, or occasionally

irregular; with hyaline to brownish or brown contents

in KOH (2%).

Ecology.

Appearing under hardwoods, especially

grandidentata

and

F. americana

; usually growing near

woody debris or growing directly from rotted hard-

woods; known from north of 44uN; May. Specimens

examined (S

UPPLEMENTARY TABLE I) were collected in

Michigan and New York.

Comments. Morchella septentrionalis

corresponds to

phylogenetic species

Mel

-24 in O’Donnell et al. (2011).

Based on available data the species can be character-

ized by the comparatively small size of the ascomata, its

northern distribution in eastern North America (from

approximately 44uN northward), its frequent associa-

tion with the deadwood of hardwoods and its relatively

small ascospores. It is easily confused with

angusticeps

, which is morphologically similar and

overlaps with

M. septentrionalis

in range, size of

ascomata and ascospores. (See the comments for

angusticeps

for further discussion.)

Morchella septen-

trionalis

probably has been misidentified as

angusticeps

(e.g. Seaver 1928, Overholts 1934, Weber

1995). It appears as the ‘‘Classic North American Black

Morel II’’ in Kuo (2005).

KEY TO 20 PHYLOGENETIC SPECIES OF

ORCHELLA

IN THE

UNITED STATES AND CANADA

1. Ridges dark gray to brown or black when young, or

pale when young and darkening to brown or black

with maturity . .......................... 2

19. Ridges pale yellowish, pale brownish, or rufescent

throughout development . ................ 12

2. Hymenophore only partially attached to the

stipe; sinus 8–20(–25) mm at point of attach-

ment . ............................ 3

29. Hymenophore almost completely attached to

the stipe except for a small (2–4 mm) sinus at

the point of attachment ............... 4

KUO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1175

3. Associated with

Populus trichocarpa

in western

North American river bottoms .....

M. populiphila

39. Associated with various hardwoods from the Great

Plains eastward .................

M. punctipes

4. Appearing in conifer burn sites; in western

North America ...................... 5

49. Not found in conifer burn sites; variously

distributed . . . ...................... 7

5. Surfaces densely tomentose (especially when

young), covered with projecting hairs (120–250 +

mm) that are brown in 2% KOH; hymenophore

and stipe usually dark gray to black but sometimes

becoming pale in direct sunlight . . . .

M. tomentosa

59. Surfaces glabrous to finely tomentose, not covered

with projecting hairs . . . ................... 6

6. Elements on sterile ridges primarily capitate

(often strikingly so) ...........

M. capitata

69. Elements on sterile ridges variously shaped,

only occasionally subcapitate; following two

species indistinguishable both morphological-

ly and ecologically based on current data

..............

M. sextelata

M. septimelata

7. Pits and ridges usually arranged in a ‘‘lad-

dered’’ vertical appearance (F

IG. 15); appear-

ing in landscaping areas and woodchip beds in

urban areas in western North America . . . . . 8

79. Pits and ridges not usually arranged in a

‘‘laddered’’ vertical appearance .......... 9

8. Ridges velvety when young; known only from the

1935 type collection in Redmond, Washing-

ton . . .

M. hotsonii

(see SUPPLEMENTARY MATERIALS)

89. Ridges glabrous or very finely tomentose

...........................

M. importuna

9. Found from the Rocky Mountains westward . . . 10

99. Found from the Great Plains eastward . . . . 11

10. Ridges pale yellowish when young, often darkening

somewhat later; stipe strikingly ridged and lacunose

...................................

M. snyderi

109. Ridges dark brown to black when young; stipe not

strikinglyridgedandlacunose ................

................

M. brunnea

,and

Morchella

sp.

Mel

-8

11. Distributed from roughly 44uN latitude northward

(currently known only from Michigan to New

York); ascomata 40–75 mm high; ascospores (19–)

20–22(–25)

mm long . . . .......

M. septentrionalis

119. Widely distributed from the Great Plains eastward;

ascomata 50–140 + mm high; ascospores 22–27

long ........................

M. angusticeps

12. Surfaces tomentose and usually dark gray to

black when young; covered with projecting

hairs (120–250 +

mm) that are brown in 2%

KOH; from conifer burn sites in western

North America ..............

M. tomentosa

129. Surfaces glabrous to finely tomentose; not

covered with projecting hairs ........... 13

13. Appearing in landscaping areas and woodchip beds

in urban areas, primarily in western North America

but introduced to central Michigan from California;

ridgesnotablyrufescent ..........

M. rufobrunnea

139. Ridges pale brownish to smoky brown or black, not

notably rufescent ....................... 14

14. Hymenophore usually subconic to conic,

attached to the stipe with a notable sinus;

western North America ............... 15

149. Hymenophore attached to the stipe without a

notable sinus ...................... 16

15. Ridges remaining relatively pale throughout devel-

opment, at maturity pale brownish and more or

less concolorous with pits; stipe not strikingly

ridged and lacunose ..............

M. frustrata

159. Ridges usually darkening to smoky brown or black

with maturity or when dried; stipe often strikingly

ridged and lacunose (F

IG.16) ........

M. snyderi

16. Mature ascomata small to medium (35–125 mm

high); pits primarily vertically arranged; dis-

tributedeastoftheGreatPlains .......... 17

169. Mature ascomata medium to large (50–220 +

mm high); pits more randomly arranged;

variously distributed ................. 18

17. Hymenophore usually ovoid with a bluntly conic to

subconic apex, 30–70 mm high; elements on sterile

ridges scattered and infrequent, variable in shape;

southeastern in distribution ......

M. virginiana

179. Hymenophore usually subconical to conical, 20–

40 mm high; elements on sterile ridges abundant,

usually subfusiform; widely distributed east of the

Great Plains. Northernmost distribution probably

southern Michigan. .............

M. diminutiva

18. Pits and ridges contorted, asymmetrical and

irregular (F

IG. 6); ridges often remaining

flattened or widely rounded into maturity;

elements on sterile ridges scattered and

infrequent ...................

M. prava

189. Not as above ....................... 19

19. Distributed in the Great Lakes region from Ontario

to Illinois and Virginia; morphologically indistin-

guishable from

M. esculentoides

.......

M. cryptica

199. Widely distributed and common east of the Rocky

Mountains (including the Great Lakes region); in

western North America found in river bottoms or

in urban areas . ...............

M. esculentoides

ACKNOWLEDGMENTS

We are grateful for the help of Cathie Aime, Cathy Cripps,

Darvin DeShazer, Ron Kerner and Carol Schmudde for

presubmission reviews. We thank Stacy Sink who provided

excellent technical assistance and John Haines who examined

Peck’s type collections and provided morphological data.

We wish to thank and acknowledge Nancy Weber for her

many years of invaluable contributions to the study of

Morchella

in North America and to mycological study in general.

We are grateful for the following collectors who

contributed specimens and data cited in this work to the

MDCP: Terry Allen, William Andrew, Keith Arquilla, Rex

Bartlett, Dent Benjamin, Lesley Bergemann, Glenn Brown,

Tommy Chiu, Norris Coit, Darvin DeShazer, Tanya Duchild,

Tom Emig, Chris England, Scott Engstrom, Gretchen

1176 MYCOLOGIA

Fitzgerald, Tim Geho, Jim George, Ken Greger, Gaston

Guzma´n, Bill Hartwig, Donna Herbold, Matthew Hill, John

Holmes, Julia Hoskins, John Jennemann, Sherry Kay, Dan

Kimberling, Toff Kobylarz, Dave Kowalishen, Judy Latchaw,

Curt Leitz, Jim Lessard, Rowlin Lichter, Douglas Ling, Jeff

Linkinhoker, Patrick Lyon, Ray Mason, James Mattan,

Cornelius McHugh, Ron Meyers, Keith Miller, Nathan

Mueller, Mary Munch, Kristin Musgnug, Bill Neill, Carl

Nielsen, Carolina Nurik, Miles Oleskiw, Ron Pastorino, Greg

Pecchia, Jamie Petersen, John Plischke, Judy Provo-Klimek,

Rebecca Rader, Morgan Sailors, John Schaefer, Wendy

Schaefer, Floyd Schmidt, Steve Schwartzman, Bob Sears, Neil

Selbicky, Tony Sepulveda, Robert Sharman, Mike Sheller,

Noah Siegel, Scott Smith, Dave Steortz, Greg Stevenson, Don

Stokes, Jeanne Storm, Alan Thrailkill, Dave Trummer, Nancy

Walker, Mike Wood, Joe Ziolkowski and Bob Zordani.

We thank the following individuals and herbaria for kindly

lending or processing specimens: the Stover-Ebinger Herbar-

ium at Eastern Illinois University (EIU, Gordon Tucker,

curator), the Harry D. Thiers Herbarium of San Francisco

State University (SFSU, Dennis Desjardin, curator), the

Mycology Collection of the Field Museum of Natural History

(F, Robert Lu¨cking, curator), the Oregon State University

Mycological Collection (OSC, Joey Spatafora, curator), the

University Herbarium of the University of California (UC,

Tom Bruns, curator), the University of Washington Herbar-

ium (WTU, Richard Olmstead, curator) and the U.S. National

Fungus Collections (BPI, Shannon Dominick, curator).

We also thank Joe Ammirati, Alija Bajro Mujic, Dick Bishop,

Roy Halling, Patrick Leacock, Lorinda Leonardi, Drew Minnis,

Tom Nauman, Vicky Nauman, Lorelei Norvell, Dennis Oliver,

Ron Petersen, Alice Piller, Jean Toothman and Sue Yocum for

invaluable assistance in various aspects of this study.

Studies of morels in National Capital Region Parks were

supported by Cooperative Agreement H3992050001 (Protect-

ing resources: sustaining wild mushrooms in four NCR Parks)

from the National Park Service, which provided financial

support to the University of Arkansas and (through sub-award

agreement SA0604193) to the University of Toronto. The

logistical support provided by park personnel on many different

aspects of the overall project is greatly appreciated.

The mention of firm names or trade products does not

imply that they are endorsed or recommended by the U.S.

Department of Agriculture over other firms or similar

products not mentioned. The United States Department of

Agriculture is an equal opportunity provider and employer.

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KUO ET AL.: REVISION OF

ORCHELLA

TAXONOMY 1177

Morchella prava strain M1300 RNA polymerase II subunit 2 (RPB2) gene, partial cds

Nucleotide Sequence

February 2012

Michael Kuo · D.R. Dewsbury · Kiaera Donnell · M.C. Carter · T.J. Volk

Morchella importuna strain M836 RNA polymerase II subunit 1 (RPB1) gene, partial cds

Nucleotide Sequence

February 2012

Michael Kuo · D.R. Dewsbury · Kiaera Donnell · M.C. Carter · T.J. Volk

Morchella brunnea strain M87 RNA polymerase II subunit 1 (RPB1) gene, partial cds

Nucleotide Sequence

February 2012

Michael Kuo · D.R. Dewsbury · Kiaera Donnell · M.C. Carter · T.J. Volk

Morchella populiphila strain M903 RNA polymerase II subunit 2 (RPB2) gene, partial cds

Nucleotide Sequence

February 2012

Michael Kuo · D.R. Dewsbury · Kiaera Donnell ·

Taxonomic revision of true morels (Morchella) in Canada and the United States

Abstract

Supplementary resources (14)