JANUARY-MARCH 2019... V

MYCOTAXON

VOLUME ONE HUNDRED THIRTY-FOUR (1) — TABLE OF CONTENTS

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HISTORY & NOMENCLATURE

North American mycology comes of age:

the 19th century participants and their roles RONALD H. PETERSEN 1

TAXONOMY

Coronosporidium ecuadorianum gen. & sp. nov.

from submerged decaying leaf from Ecuador DayneT Sosa,

ADELA QUEVEDO, FERNANDO ESPINOZA, LIZETTE SERRANO,

FREDDY MAGDAMA, Marcos VERA, SIMON PEREZ-MARTINEZ,

ELAINE MA.osso, RAFAEL E. CASTANEDA-RuI1z 111

Trichoglossum tetrasporum, newly recorded from India

SANJIT DEBNATH, KRIPAMOY CHAKRABORTY,

BADAL KuMAR DatTTA, PANNA Das, AJAY KRISHNA SAHA 119

Perenniporia mopanshanensis sp. nov. from China

CHANG-LIN ZHAO & XIANG Ma 125

New records of Bilimbia and Toninia from China MEI-JIE SUN,

SHU-KUN YAN, RONG TANG, CHUN-XIAO WANG, Lu-LU ZHANG 139

Anamylopsora altaica sp. nov from Northwestern China

PaRIDA AHAT, ANWAR TUMOUR, SHOU-Yu Guo 147

Four Pyrenula species new to China JrE-MENG Fu, ANDRE APTROOT,

ZHONG-LIANG WANG, Lu-Lu ZHANG 155

Spadicoides matsushimae sp. nov., and

Anisospadicoides gen. nov. for two atypical Spadicoides species Min Qtao,

DeE-WE!I Lt, ZE-FEN Yu, Kat ZHANG, RAFAEL F, CASTANEDA-Ruiz 161

New records of Hymenoscyphus, Parascutellinia, and

Scutellinia for Turkey Aut KeLes 169

Lemonniera yulongensis sp. nov. from Yunnan, China

ZE-FEN YU, YI-FAN LV, Bo FENG, MIN Qiao 177

VI ... MYCOTAXON 134(1)

Contributions to species of Xylariales in China—2.

Rosellinia pervariabilis and R. tetrastigmae spp. nov.,

and a new record of R. caudata

XIN XIE, Liti Liu, XU ZHANG, QINGDE LONG, XIANGCHUN SHEN,

SARANYAPHAT BOONMEE, JICHUAN KANG, QiRuI Li 183

Catalog of Penicillium spp. causing blue mold on

bulbs, roots, and tubers FRANK M. DUGAN & CarL A. STRAUSBAUGH 197

Sulzbacheromyces yunnanensis, a new record for Thailand

NAKARIN SUWANNARACH, JATURONG KUMLA, KANITTA SATIENPERAKUL,

WITCHAPHART SUNGPALEE, SUTHEERA HERMHUK, PIYAWAN SUTTIPRAPAN,

KRIANGSAK SRI-NGERNYUANG & SAISAMORN LUMYONG 215

MycoBioTa (FUNGA) NEW TO THE MYCOTAXON WEBSITE

A checklist of the non-gilled fleshy fungi (Basidiomycota)

of Kerala State, India (SUMMARY)

T.K. ARUN KUMAR, ANJITHA THOMAS,

KRISHNAPRIYA KUNIYIL, SALNA NANU, VINJUSHA NELLIPUNATH 221

JANUARY-MARCH 2019...

NOMENCLATURAL NOVELTIES AND TYPIFICATIONS

PROPOSED IN MYCOTAXON 134(1)

Anamylopsora altaica Ahat, A. Abbas, S.Y. Guo & Tumur

[FN 570571], p. 150

Anisospadicoides R.F. Castafieda, Qiao & Z.F. Yu

[MB 827953], p. 162

Anisospadicoides macrocontinua (Matsush.) R.F. Castaneda, Qiao & Z.F. Yu

[MB 827954], p. 165

Anisospadicoides macroobovata (Matsush.) Qiao, Z.F. Yu & R.F. Castaneda

[MB 827955], p. 165

Coronosporidium R.F. Castaneda, Quevedo & D. Sosa

[IF 554557], p. 113

Coronosporidium ecuadorianum R.F. Castaneda, Quevedo & D. Sosa

[IF 554558], p. 113

Lemonniera yulongensis Z.F. Yu

[MB 827810], p. 178

Perenniporia mopanshanensis C.L. Zhao

[MB 827580], p. 132

Rosellinia pervariabilis Q.R. Li & J.C. Kang

[MB 827512], p. 188

Rosellinia tetrastigmae Q.R. Li & J.C. Kang

[MB 828157], p. 190

Spadicoides matsushimae R.F. Castaneda & D.W. Li

[MB 809530], p. 165

VII

Vil ... MYCOTAXON 134(1)

ERRATA FROM PREVIOUS VOLUMES

VOLUME 133(2)

Front cover, bottom line FoR: 133(#) READ: 133(2)

VOLUME 133(4)

p- 729, line 3 (masthead) FoR: pp. 729 READ: p. 729

Back cover (online edition only):

REPLACE single bottom line by

(Kaur & al.—Figs 2,3; p. 678)

Maninder Kaur, ARTIST

Back cover (printed hard copy only): after bottom line

ADD: Maninder Kaur, ARTIST

JANUARY-MARCH 2019...

REVIEWERS — VOLUME ONE HUNDRED THIRTY-FOUR (1)

The Editors express their appreciation to the following individuals who

have, prior to acceptance for publication, reviewed one or more of the

papers prepared for this issue.

Meredith Blackwell

Rafael F. Castafieda-Ruiz

Dinushani Anupama Daranagama

Edit Farkas

Tatiana Baptista Gibertoni

Shouyu Guo

Luis EP. Gusmao

Liu-Fu Han

Wojciech Janisiewicz

Samantha C. Karunarathna

Steven Koike

De-Wei Li

Michael Loizides

Patinjareveettil Manimohan

Eric H.C. McKenzie

Troy McMullin

Josiane Santana Monteiro

Karen K. Nakasone

Lorelei L. Norvell

Shaun R. Pennycook

Liliane Petrini

David H.S. Richardson

Ertugrul Sesli

Harrie J.M. Sipman

Jie Song

Steven L. Stephenson

Larissa Trierveiler Pereira

Ibrahim Tirkekul

Jan Vondrak

Xiu-Guo Zhang

x ... MYCOTAXON 134(1)

2019 MyYCOTAXON SUBMISSION PROCEDURE

Prospective MycoTaxon authors should download the MycoTaxon 2019 guide,

review & submission forms, and MycoTaxon sample manuscript by clicking the ‘file

download page’ link on our INSTRUCTIONS TO AUTHORS page before preparing their

manuscript. This page briefly summarizes our “4-step’ submission process.

1—PEER REVIEW: Authors first contact peer reviewers (two for journal papers;

three for mycobiota/fungae) before sending them formatted text & illustration

files and the appropriate 2019 MycoTaxon journal or weblist reviewer comment

form. Experts return revisions & comments to BoTH the Editor-in-Chief

<editor@mycotaxon.com> and authors. ALL co-authors MUST correct and proof-

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Place first author surname + genus + ‘MycotTaxon’ on the subject line, and

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3—FINAL SUBMISSION: All coauthors thoroughly revise and proof-read files

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4—FINAL EDITORIAL REVIEW & PUBLICATION: The Editor-in-Chief conducts a

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The PDF proof and bibliographic & nomenclatural index entries are sent to all

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MyYCOTAXON ONLINE— www.ingentaconnect.com/content/mtax/mt

The Mycotaxon journal publishes four quarterly issues per year. Both open access

and subscription articles are offered.

JANUARY-MARCH 2019... XI

FROM THE EDITOR-IN-CHIEF

ON TIME! (FOR A CHANGE)—Loyal Mycotaxon authors and readers are

undoubtedly weary of the constant stream of editorial apologies following yet

another issue released 1-2 months AFTER the end of a March, June, September,

or December quarter. We are equally tired of apologizing to you. Thus it is

with immense pleasure that we announce the release of MycoTaxon 134(1) in

Marcz! To ensure future timely journal delivery, we [i] urge authors to send

their manuscripts to the EprIror-IN-CHIEF immediately after receiving their

final review from the NOMENCLATURE EDITOR and [ii] set a June 15 closure

deadline for 134(2), the 2019 April-June issue. Submissions received after that

date will be scheduled for the following (July-September) issue.

MyYCOTAXON 134(1) presents 14 papers by 64 authors representing 10 countries

and peer reviewed by 30 expert reviewers. The issue opens with Ronald H.

Petersen’s 110-page lively and welcome historical treatise on 19" century

mycologists in North America. This provides us the opportunity to introduce

two new journal divisions—HisTORY & NOMENCLATURE and TAaxONOMY—

to supplant our former ‘RESEARCH PAPERS’ designation. [“Biodiversity &

ecology” still find their home under the Mycosrota/FUNGAE summaries of

papers posted on the MycoTaxon website, here announcing a checklist of

fleshy non-gilled fungi from Kerala uploaded in February. |

The 2019 January-March MycoTaxon also proposes Two new genera

(Anisospadicoides with two species from PERU and Coronosporidium from

ECUADOR) and SEVEN species new to science representing Anamylopsora,

Lemonniera, Perenniporia, and Rosellinia from CHINA; Coronosporidium

from Ecuapor; and Spadicoides from PERU.

New species range extensions represent ascomycetes Hymenoscyphus,

Parascutellinia, and Scutellinia for TurKEy and Trichoglossum for INDIA,

ascolichens Bilimbia, Pyrenula, Toninia for mainland Curna, and the

basidiolichen Sulzbacheromyces for THAILAND.

Plant pathologists especially will welcome the paper covering Penicillium

causing blue mold on bulbs, roots, and tubers that tabulates where to find

references to descriptions, culture methodologies, phylogenies, and hosts for

each species.

Warm regards,

Lorelei Norvell (Editor-in-Chief)

22 March 2019

MY COTAXON

January-March 2019—Volume 134, pp. 1-110

https://doi.org/10.5248/134.1

North American mycology comes of age:

the 19" century participants and their roles

RONALD H. PETERSEN

Ecology & Evolutionary Biology, University of Tennessee, Knoxville TN 37996-1100

CORRESPONDENCE TO: repete@utk.edu

ABSTRACT—Mycology in North America in the early 19th century hung by a single thread.

By century’s end there were numerous workers in diverse mycological fields. The players,

their lives, and their roles sketch the development and growth of the science to set mycology

on a steady course by century’s end.

Europe's head start

Although botany had a thread of presence in colonial and post-colonial

North America, Europe's society developed somewhat earlier than American,

with selected national capitals achieving cosmopolitan status before 1800.

London, Paris, Madrid, Rome, Vienna—all vied for sophistication in the midst

of poverty, disease, and superstition. But as the emergence of the binomial

system of plant naming (Linnaeus 1753—SprEcIES PLANTARUM) took root

post-mid-century, local plants were collected, herbaria initiated, and floras

written and often illustrated (Petersen 2001). Even examples of fungi were

being publicized: in England, the 1787-1813 CoLoRED FIGURES OF BRITISH

FUNGI OR MUSHROOMS by James Sowerby (1757-1822), William Curtis's

BOTANICAL MAGAZINE (first issued in 1787), and the 1788-1791 AN HisTory

OF FUNGUSES ABOUT HALIFAX by James Bolton offered a sampling of local

mushrooms. In Paris, the 1791-1812 HisTOIRE DES CHAMPIGNONS DE FRANCE

by Bulliard (1752-1793) offered hundreds of fungi—all illustrated—while the

1790 TRAITE DES CHAMPIGNONS by Paulet (1740-1826) also contributed. In

the future Germany, J.C. Schaeffer (1718-1790) published four volumes of

2 ... Petersen

FUNGORUM QUI IN Bavaria... (Schaeffer 1762-1767), and Batsch (1761-1802)

summarized fungi from farther north in his ELENcHus FUNGoRum (Batsch

1783-1789). From Scandinavia came the 1790 Beata RuRIS by Holmskjold

(1731-1793) and, of course, the SPECIES PLANTARUM by Linnaeus (1707-

1778). This productive era in European mycology has been best summarized

by Ainsworth (1976).

The narrow strip of British colonies along the North American Atlantic

coast saw some botanical explorers imported from England in the 17" century.

During the 18" century, Catesby, the second Clayton, the Bartrams (father

and son), Linnaeus’s student Pehr Kalm, and others explored botanically, but

their plunder made its way back to their homes in Europe (Petersen 2001).

As the 18" century rolled into the 19", new explorers arrived. André Michaux

(1746-1802) and his son, John Fraser (1750-1811) and his son, John Lyon

(1765-1814), Thomas Nuttall (1786-1859), and others prowled the forests, often

into the wilderness west of the colonized coast. Their quest, however, was prompted

not by pure botanical curiosity but rather by a desire to introduce American

plants into European gardens. Whether from patriotic or financial motives, many

thousands of plants were shipped to England and France accompanied by timber

and furs from the more northern climes of Canada. The botanical exploitation of

North America was gaining momentum (Petersen 2001).

During this formative period, the colonies fought a revolutionary war

against British rule (1776-1779) and although emaciated, the colonies

prevailed, resulting in independence. But the principle of independence

did not make the colonies independent in anything but a political sense.

The colonies owed a debt to France, Germany, and Poland for aid during

the Revolution. Trade with Britain had included not only imports from the

homeland but exports of American goods, both directions now strangled

in the wake of war. Moreover, Tories (those in the colonies who supported

British rule) remained in place. It would be another decade before the colonies

would coalesce and adopt a constitution, with the colonies metamorphosing

into states of a Union. Historically, the following century would see dizzying

events, inventions, mercantile intercourse, immigration, urbanization—all

punctuated by “The War between the States.” From mycologists who could

be counted on less than one hand, there grew a population of both fingers

and toes. These roughly eight decades and the development of mycology ona

“new continent are the subject of this sketch.

In the first quarter of the 19" century, significant mycological compilations

were Offered, chief of which attempted to arrange the fungi into some

Mycology in 19th century North America... 3

taxonomic system, sometimes patterned on quasi-religious schemes. In three

ground-breaking treatises—the 1801 Synopsis METHODICA FUNGORUM by

Christiaan Hendrik Persoon (1761-1826), the 1816 Das SysTEM DER PILZE

UND SCHWAMME by Nees von Esenbeck (1776-1856), and the first volume

(1821) of Systema Myco.ocicvum by Elias Magnus Fries (1794-1876)—each

author praised his predecessor(s) yet reorganized the fungi to suit his personal

philosophical framework. Fries’s volumes especially became a primary source

for mid-century mycologists (Petersen & Knudsen 2015), including most

Americans.

The Atlantic Coastal Plain of North America drains the eastern Continental

Divide Watershed. Just to its west, the Piedmont Plateau is elevated by some

feet or yards. This slight discrepancy in altitude, it was found, could be

used for waterpower as the rivers and streams dropped. Waterwheels were

installed, manufacturing increased, and population centers grew apace. Once

steam power was developed, the manufacturing hubs did not evaporate; only

the means of power changed. As the 19"" century wore on, one might have

expected that mycology, an almost undiscovered corner of natural history,

would have followed this string of population centers and major coastal port

cities. This, as will be shown, would be a mistake.

In the newly minted United States, Lewis David de Schweinitz (1780-1834;

Fic. 1) was born and raised in America (unlike his illustrious mycological

forebears) as part of a Moravian colony in Pennsylvania. In 1798, his family

rejoined the original colony in Germany, where Lewis obtained his divinity

credentials. While there he botanized with his mentor Johannes Baptista

Albertini, with whom he published a compilation of fungi in their region

(Albertini & Schweinitz 1803). When Schweinitz returned to America, he

remained involved with botany, including fungi. His first major mycological

paper (Schweinitz 1822) summarized his observations of the fungi in North

Carolina. His second offering (Schweinitz 1832), which covered the middle

areas of eastern North America, was the first large mycological paper

compiled, written, and published in the United States (Hewett & al. 2016,

Arthur & Bisby 1918, Fries 1946, Johnson 1835, Rogers 1977) except for one

earlier paper by Bosc (1811). Unfortunately, with Schweinitz’s death in 1834,

any possible first-hand American mentoring also died. American mycological

research came to a pause except for a very weak dribble of specimens from

America to Europe. As the only source of named vouchers for North American

fungi, the Schweinitz fungus herbarium was to become the target of repeated

visits and fragmentation (Petersen 1979).

4 ... Petersen

Fic. 1. LEw1is DAVID DE SCHWEINITZ

[Mycological Notes no. 44, 1916]

When Schweinitz sailed to and from North America and Europe, his

mode of transportation would have been a wooden, 2-3-masted ship,

probably rigged for cargo (and ballast) with little room for passengers.

His berth would have been spartan and strolls on deck invited

seasickness. Such a ship was at the mercy of vicissitudes of winds and

weather. Weather prediction was primitive, with “Red skies in morning,

sailors take warning; red skies at night, sailors delight” bearing some

truth.

Mycology in 19th century North America ... 5

Antebellum mycology

Whether it is called the “American Civil War” or the “The War Between

the States,” the events and the period were and remain indisputable. The

causes were several and remain to this day blurred but filtered by time, so

that a few sentiments still reverberate in American culture.

In the wake of the Revolution, with the northern states developing

densely commercial manufacturing centers while the south continued

its rural production of cotton, rice, indigo, etc., northern congressional

representatives had legislated high tariffs to protect its industries, but

such tariffs favored the north and punished the south. In the 1830s

South Carolina voted to declare the tariff bill null and void (“sovereign

nullification”), and this forced a legal discussion on “states’ rights.” Bound

up with states’ rights was the issue of slavery. Many in the north viewed

slavery as inhumane and cruel, while in the south, slaves were property,

just as were houses, horses, crops, and land—to be owned outright as

chattel. After all, slaves were bought, sold, exchanged, and otherwise

treated as property. The south saw its sovereign right to own slaves as

being threatened, and as new states were admitted to the Union, there was

a potential legislative imbalance of slave versus non-slave in the federal

government. And if individuals had rights (to own slaves), then states had

rights, one of which was to nullify federal laws in individual states, and

if that failed, to secede. And thus the issue was joined. This era brought

disruption to all society in the United States, and thence to other countries

ccc

of the Americas and to Europe. “Much good, Henry W. Ravenel asserted,

had blessed Blacks from enslavement. They had increased in number since

coming to the United States, a sure sign of health and happiness, and were

enabled here to be useful while daily making progress in attaining a higher

level of civilization” (Haygood 1983: 228).

If a significant route to mycology as a discrete pursuit in the 19"

century could be identified, it would proceed through a personal interest

in “natural history,’ thence through botany. In botany there already existed

individual American workers (rarely “professional”) participating in

collecting, identifying, and preserving green-plant specimens. For the two

leading American mycologists of the first half of the century, this was their

path of avocational migration. Both came to this activity in the 1840s, and

(although one was a northerner) both were located in the southern states.

Their backgrounds, however, were remarkably different.

6 ... Petersen

Fic. 2. MosEs ASHLEY CURTIS

[Berkeley & Berkeley 1986]

Moses Ashley Curtis (1808-1872; Fic. 2) was already a frequent correspondent

of Asa Gray (1810-1888; Fic. 3) and John Torrey (1796-1873). He was

considered THE authority on southern mountain plants, which Torrey and Gray

coveted for their work toward a NORTH AMERICAN FLORA (Petersen 1980a).

Curtis, a Massachusetts product, first moved to Wilmington, North Carolina,

on the Coastal Plain, employed as a tutor, but after obtaining a Doctor of

Divinity he became an Episcopal minister as a life’s work. He had collected

flowering plants around Wilmington and published lists (Curtis 1835, 1843),

but diminishing returns of new taxa turned his mind to other plants. He

Mycology in 19th century North America ... 7

Fic. 3. ASA GRAY

[Neilans, 1963]

corresponded with and sent specimens to Edward Tuckerman (1817-1886),

the Harvard lichenologist, but soon became intrigued with fungi. Curtis came

to the fungi following laudatory words from Torrey and Gray and through

them, from William Jackson Hooker (1785-1865), the venerable Kew

botanist. Curtis recognized his fledgling status, but over time he assumed the

perceived role of overseer. But on beginning to take this new interest seriously

and having nowhere to turn for help in recognizing and classifying various

fungi, Curtis wrote to his old botanical friend, Asa Gray. Gray recognized

the dearth of mycologically inclined Americans and advised Curtis to

contact Miles Joseph Berkeley (1803-1889; Fic. 4) in England, who was

already sharing his Anglican clerical duties with his mycological avocation.

8 ... Petersen

‘

Yt sin

eee!

wwe:

Fic. 4. MILES JOSEPH BERKELEY

[Grevillea, vol. 1. Frontispiece]

Curtis wrote a beseeching letter in 1846 (Petersen 1980a) and, considering

the times, Berkeley “quickly” replied with encouraging words and willingness

to receive and identify specimens from Curtis. (The arrangement, prevalent

at that time, was for the supplicant to send specimens for identification

while retaining a duplicate to which the name could be applied. The modus

operandi has changed little to the present.) Thus began a 24-year relationship

bracketing the Civil War, between the two clergymen cum mycologists: one

master, the other acolyte. The post-Schweinitz mycological hiatus was closing.

Mycology in 19th century North America ... 9

For Curtis, one irksome item was the proposal of names for American

plants by European workers, caused by specimens exchanged or exported

from North America to foreign collaborators. Nevertheless, over his years

with the fungi, Curtis would send a couple thousand specimens to Berkeley,

who compassionately tacked Curtiss name on them (e.g., Boletus spraguei

Berk. & M.A. Curtis).

Very soon thereafter (14 Apr 1847) Curtis wrote to Berkeley: “On the

Santee, still further south, I have a mycological associate who is collecting

pretty diligently, & the only person in the country, besides myself, who collects

Fungi but I hope for assistance from Alabama also” [presumably from Peters

in Moulton, AL].

Once involved with fungi, the only American predecessor of any note

immediately reappeared in Curtis’s interest. The pioneer work of collecting,

classifying arranging higher plants in eastern North America had already

brought Schweinitz into Curtis’s orbit, but his early contribution on fungi

covered a region in German city-states during the beginning of the 19%

century (Albertini & Schweinitz 1803, Hewett & al. 2017). However, upon

returning to the United States, Schweinitz studied the fungi he found on his

routes between North Carolina and eastern Pennsylvania where he ministered

over his Moravian congregations. Moreover, Schweinitz not only published

on fungi but amassed a sizable herbarium that was preserved at the Academy

of Science in Philadelphia some 21 years after his death.

In 1848, Curtis set his sights on examining the Schweinitz specimens. The

curator of the Philadelphia herbarium, Elias Durand (born Elie Magliore

Durand, 1794-1873) initially refused Curtis's request. Undaunted, during

1850 Curtis took advantage of a trip to the northeast to “drop in” on the

herbarium; upon receiving permission to inspect the specimens and to take

a fragment when enough material existed, proceeded to take large enough

pieces to share with Berkeley (as well as a few other parties).

Henry William Ravenel (1814-1887; Fic. 5) represented the latest generation

of southern family plantation owners. His home territory was the Coastal

Plain of South Carolina, his crop chiefly Sea Island cotton and rice, and his

work force slaves; his father and grandfather had been plantation and slave

owners before him. After the usual academy education, Ravenel entered South

Carolina College in 1829 at the age of 15 (the minimum age for enrollment).

The student body numbered about 100 and the faculty six (Haygood 1983:

44). The curriculum included a bit more science than usual, but Ravenel,

10 ... Petersen

Fic. 5. HENRY WILLIAM RAVENEL

[Neilans, 1963]

graduating in 1832, would have had a grounding in the classics. Although at

this time there was a concerted effort to enlarge the natural history library, the

collection was no rival to those of older (northern) institutions, and natural

history was not to be generally represented for another 20-30 years. (Asa Gray

would not come to Harvard as a botanist until 1842 nor would laboratory-

based teaching be offered for another decade or more.) Whatever respect and

love of natural things Ravenel would gain had to come from within.

Mycology in 19th century North America... 11

It should be noted that during Ravenel’s college days (1829-1832), one

campus society debated whether South Carolina should secede from the

Union. Although decided in the negative, the topic presaged actual secession

by over 30 years (Haygood 1983: 55).

Sometime near his graduation, Ravenel was persuaded to begin studies

for a medical degree, but his father countered by offering his son a plantation

of his own, where slave labor would relieve Henry of the rigors of a rural

medical practice. In 1835 (age 21) Henry became engaged to Elizabeth

Gallard Snowdon, daughter of another wealthy planter. Her father’s wedding

gift included “18 Negro slaves” and several shares of bank stock. From Henry's

father came Northampton Plantation, 804 acres in the neighborhood of

present-day Lake Moultrie, SC. “By the end of 1839 Ravenel, at the age of

twenty-five, had a wife and family, land, slaves, the equipment and knowledge

necessary to raise cotton, a winter house at the plantation, and a summer home

in Pinopolis” (Haygood 1883: 76). A year later Ravenel bought an additional

eight parcels for $6800: money seemed of little concern, and Ravenel bought

a small microscope and books whenever he pleased.

In September 1846, Ravenel wrote to Curtis, thereby contacting another

southern green-plant botanist even as Curtis himself was writing Gray, “can

I get the like help [as Tuckerman in lichens] on the fungi? Can you introduce

me to any botanist, American or foreign, who understands these things &

will exchange with me, or will name my specimens for the gift of them?”

(Haygood 1983: 102). It was not long before Ravenel, too, began to migrate

toward mycology, although never giving up other botanical pursuits.

Dr. Ezra Michener’s (1794-1887) voluminous correspondence, which

contained letters from some of the most eminent botanists of his time,

including Curtis, Ravenel, Tuckerman and Gray (Harshberger 1893, 1899),

appears to have been destroyed (Shear & Stevens 1917) or held in private

family hands (Harshberger 1899: 178-180).

Ezra Michener was born Nov. 24, 1794, on a fourth-generation farm in the

small Quaker town of London Grove Township, Chester Co., Pennsylvania.

His early education was in the local schools, and he showed signs of a passion

for botany early in life. Facing the dearth of botanical literature for American

beginners during his first two decades (Harshberger 1899), Michener

overcame that hardship by teaching himself through trial-and-error (Kelly

& Burrage 1920).

12 ... Petersen

At age 21, Michener moved to Philadelphia in 1815 to live with the Dr.

David Jones Davis family. During 1816, after attending botany lectures by Dr.

William P.C. Barton, Ezra was appointed house surgeon at the Philadelphia

Dispensatory (!)—salary $250 per year (Harshberger 1899), and in 1819, he

married Sarah Spooner. By this time, Barton’s FLORA PHILADELPHICAE was

finally published with FLoRULA CEsTRICcA (1826) by William Darlington

(1782-1863) not far behind. At last there was help of a sort, although neither

book dealt in depth with the lichens and hardly at all with fungi.

At first, Michener was interested in higher plants and became involved

with Darlington’s FLORULA. It was probably this interest that led him to Curtis

and later to Ravenel, both of whom had similar avocations. Later, perhaps in

the early 1850s, Michener realized that one plant group missing from the list

was the fungi, a group for which there was also little help in identifications.

By the time Michener wrote to Curtis concerning fungi around 1852, he was

a mature 56-year-old Quaker country physician in Chester County and a

“prominent citizen interested in reform movements” (Shear & Stevens 1917).

Almost all of his life had been (and would be) spent in a small village “almost

entirely isolated,’ as he wrote, “from the seats of learning, from scientific

libraries, and from personal intercourse with the cultivators of science”

(Michener’s autobiographical notes, as quoted by Shear & Stevens 1917).

Michener’s interest in fungi finally fell on welcoming eyes, and Curtis invited

him to send as many specimens as he wished. They would be filtered and

added to those bound for Berkeley in England.

The wanderers

Another actor entered the scene in 1847. Augustus Fendler (1813-1883;

Fic. 6), a Prussian émigré, had been chosen by George Engelmann (St. Louis,

Missouri) and Asa Gray to collect plant specimens in the far reaches of the

southwestern territories (western Texas, New Mexico). After Fendler returned

from the expedition to deliver his plunder, an extremely impressed Gray and

Engelmann tried to send Fendler back again.

Although of humble origin and delicate health, Augustus Fendler emigrated

from Berlin, Germany, to the United States in 1835 (age 22), with almost no

personal belongings. His first job in Philadelphia was in a tannery, after which

he worked at a lamp manufacturing business in New York. [Incandescent

bulbs were not available for some years: lighting was by gas.] He moved on

to St. Louis, where he arrived after a thirty-day trip on a paddle-wheeler up

the Mississippi against the current. Although employed, he soon left for New

Mycology in 19th century North America... 13

Fic. 6. AUGUSTUS FENDLER

[Ann. Missouri Bot. Gard. 76(1). 1989]

Orleans, crossing the Mississippi, and walking through Illinois, Kentucky,

and West Tennessee. Once in New Orleans he decided to continue to Texas,

where he arrived in 1839, only two years after Charles Wright (more below),

and stayed for about a year before returning to Illinois where he taught school

for some time. The fall of 1841 found him on an uninhabited island in the

Missouri River, where he lived in solitude (Canby 1885).

In 1844 he sailed back to Berlin, where he learned that sets of plant

collections (exsiccati) could be valuable, and on his return to St. Louis he

14 ... Petersen

started collecting plant specimens in various expeditions in the United

States, including west Texas and eastern New Mexico. Panama was another

destination. Fendler became known for the quantity and quality of his

collections. Asked by Engelmann and Gray to return to Texas/New Mexico

(Crosby 1973), Fendler flatly refused (Stieber & Lange 1986), but later

emigrated to Venezuela, where he collected fungi and other botanicals

(Caomhanach 2012).

Fic. 7. CHARLES WRIGHT

[Wikipedia]

While many mycologists (and others) enjoy the outdoors and are eager

to explore rarely investigated ecosystems—or well-explored places not

Mycology in 19th century North America... 15

previously sampled for a particular fungal group—there is a special breed

intent on gathering specimens merely for identification and to add to

knowledge of taxonomic variation. Augustus Fendler was one of these and

John Macoun would be another, but Charles Wright (1811-1885; Fic. 7) saw

more of the world than the other two and brought home a greater spectrum of

“plants.” Of the three, he was home-grown, born 29 Oct 1811 in Wethersfield,

Connecticut, on the family homestead where he died.

Early educated in local schools and Yale College (1831-1835), Wright was

attracted to botany, although no particular mentor has been identified. As

another northerner, M.A. Curtis, had moved south, upon graduation Charles

also moved south to Natchez, Mississippi, where he was a family tutor on a

plantation. A general financial depression of 1836 affected his employer and

Wright was out of a job.

A rush of immigration was underway to Texas (then an independent

republic) and Wright joined the migrants in 1837, walking from the

Mississippi to the Sabine River. Wright became part of a surveying crew for

a couple years, but he was always seeking and collecting plant specimens.

In 1844, he established a correspondence with Asa Gray and sent a large

gathering from Texas. Gray was impressed by the quality of the cache and

gladly received Wright's harvest through upcoming years. He (Gray) acted to

portion out specimens according to the expertise of the recipient.

A seemingly unrelated event of no little consequence occurred in the 1840s

and illustrates the times in which American mycologists were working.

William Miller (1782-1849) gained gravitas by making Bible-based

calculations precisely predicting the end of time. In 1844, many adherents

gathered, but the appointed time came and went—the effect was “The Great

Disappointment” that spawned sects known as Millerites, including Advent

Christians (1860) and the Seventh-day Adventists (1863), the latter remaining

with significant parishioners.

A small but diligent network of botanical correspondents operated in

the northeast during the 1840s, but the group was quite willing to co-

opt southerners who could contribute specimens from other regions and

simultaneously reveal new taxa. Gray, Tuckerman, William Oakes (1799-

1848), and Stephan Thayer Olney (1812-1878) were prominent in northern

botany. A small botanical clique was developing in the south, but always in

touch with the northern savants. The chief printed manual for the southerners

16 ... Petersen

was that by Stephen Elliott (1821-1824), for the books by Eaton and Gray did

not adequately treat the southern flora.

Once in touch and focused on mycology, Curtis advised Ravenel on

preservation of fungus specimens (a universal problem). Detailed notes

on pileus color, lamellar attachment, and spore color were needed before

pressing, since all were distorted during the preservation process. To Gray,

Curtis commended Ravenel’s vigor and quantities of specimens. Despite the

yawning discrepancy between Ravenel’s wealth and Curtis's miserly salary

and the manner of their upbringing, the two men did share a passion for

botany and nature.

In March 1848, Ravenel visited Curtis in Society Hill, North Carolina.

During their conversation, Ravenel inquired about establishing direct

contact with Berkeley, as Curtis wished to remain the sole source (and filter)

of American fungi to the Englishman. Eventually relenting, Curtis wrote to

Berkeley to relate Ravenel’s inquiry, but also suggested the French botanist,

Camille Montagne (1784-1866). Berkeley countered, recommending his own

new assistant, Christopher Broome (1812-1886). Berkeley seemed amenable

to direct contact with Ravenel, but Curtis struck an agreement with Ravenel:

all identified specimens could go directly to Berkeley, but unidentified

specimens would go to Curtis for possible forwarding. Curtis stood firm in his

coveted role as conduit. This difference of opinion marked a possible rift, but

Ravenel demurred back into his role as collector. “If Ravenel was agreeable, he

was not cowed” (Haygood 1983: 114).

Up north, Charles Horton Peck (1833-1917; Fic. 8) was born and raised in

“Sand Lake, Rensselaer County, [which] was just a few clearings less than an

unbroken forest.” Peck’s father and grandfather ran a sawmill manufacturing

oak staves for sale in Albany, the nearby New York state capital. In his teenage

years, Peck worked in the sawmill and attended “winter-school? “The school

house that provided shelter for the master and a few children from the nearest

homes was built of logs. The seats were made of saw-log slabs turned flat

side up” (Haines 1978). Peck was described as having a “naturally delicate

constitution.”

In the late 1840s, Ravenel’s confidence built, his library grew, and his circles of

correspondence became substantial, perhaps especially including Tuckerman

who was about to travel to Europe and who agreed to spread Ravenel’s

Mycology in 19th century North America ... 17

Fic. 8. CHARLES HORTON PECK

[Bot. Gaz. 65(1). 1918]

name (and some specimens) to worthy European workers. Ravenel was

delighted and particularly singled out Elias Fries as a possible recipient (and

identifier) of his (Ravenel’s) lichens. 1849 marked Ravenel’s first botanical

publication (on higher plants). By 1850, Ravenel was a respected member of

the local agriculture society, state-wide botanist, and nation-wide botanical

correspondent. Although Curtis was no longer essential, the two continued

to exchange letters and occasional specimens, but Ravenel had already issued

century 1 of FUNGI CAROLINI ExsiccaTI without Curtis's help or credit. Curtis

found little of substance to criticize but disparaged the many typographical

errors in the labels. This needling and belittlement would continue for his

entire life.

18 ... Petersen

During 1845-1850, the European “potato famine” resulted in mass migration

to the US., especially Irish. Unrest across continental European states also

produced mass migrations of German, Swiss, and other German-speaking

people. All groups were generally abjectly poor, illiterate and (especially the

Irish) not Protestant but Roman Catholic. They met with discrimination

including employment, integration into society, and even voter suppression.

One result was ghetto enclaves, especially around disembarkation points,

Boston (Irish) and New York (German). Ravenel considered the south’s

insularity a good thing, not a bad thing. Raw materials from the south (cotton,

rice, sugar, tobacco) were essential to the north, so the north would not risk

war and interruption of this flow.

But the “Trish Potato Famine” also provoked a different, if more esoteric,

interest. In 1845, the popular English GARDENER’S CHRONICLE AND

AGRICULTURAL GAZETTE carried reports and editorials on the devastating

“murrain” as it occurred in country after country. Two theories of its origin

were published, that 1) the extreme weather variation of that growing season

caused degeneration and rotting of plants, including tubers, which then,

once weakened, were prone to molding or 2) the mold was the cause, not

the result, of the disease. The former was a popular notion espoused by John

Lindley, editor of THE GARDENERS’ CHRONICLE; the latter was held almost

solely by Rev. M.J. Berkeley, the clergyman who devoted much time with all

sorts of molds, mildews, mushrooms, and toadstools (Large 1962; Berkeley

1845). Although no remedy could be put forward, the claim of disease caused

by fungi was put forward decades before Pasteur’s germ theory. Some time,

however, would elapse before the media tangle would be resolved, but the

paroxysm spawned an interest in possible links to other plant diseases caused

by microorganisms, including fungi. Simultaneously, Grape Mildew spread

across Europe, again with a fungus suspected as causal agent. Plant pathology

was born and would become a dominant pathway toward mycology as an

independent science.

One victim of the “potato murrain” was Sarah Macoun, who lived with her

sons on the tiny family farm, now failing, in Magheralin, County Down,

Ireland. With her last savings she and sons emigrated to Canada in 1850,

settling in Seymour Township, Ontario, to resume farming.

When the Macoun family migrated, their conveyance might have been a

ship rigged for both sail and steam propulsion. The former saved expense,

using wind power and a rudder for control; the latter demanded fuel (coal)

Mycology in 19th century North America... 19

and a complex, heavy engine. The means of propulsion would have been a

paddle wheel, either posterior to the hull or as a side-wheeler, carefully

designed so the paddle was not immersed too deeply.

One of her sons, John (born 17 April 1831) seemed not interested in

farming, although “a small wiry man, with little formal education and without

capital... John by his own account seems to have been a pugnacious boy who

always came out best in fights by using his left hand. He left school at thirteen

and little is known about him until after he went, at the age of twenty, to

Canada.... He made up his mind to become a teacher in order to have spare

time for study of botany. After studying grammar on his own, he gave up

his job, and walked the forty-three miles from the farm to see the county

school inspector in Toronto, presumably with a view to seeking recognition

for teaching. Three weeks later he gained his teacher's certificate” (McGeown

undated).

Mm. 1849

4 Z angie, ss

Fic. 9. A.C.J. CORDA

[Wikipedia]

In 1849, A.C.J. Corda (Fic. 9) came from the far-off Austro-Hungarian

Empire to the German settlement of New Braunfels, Texas, at least to collect

20 ... Petersen

plants (including fungi) and perhaps to make a home. Asa Gray and Curtis

knew of his presence in America, although somewhat dismissively despite his

impressive five-volume set of ICONEs (Corda 1837-1842) and compendious

ANLEITUNG (Corda 1842). Regardless of Corda’s plans, on August 25, 1849,

the ship carrying him and his specimens away from New Orleans disappeared

and Corda was lost.

Enlarging the circle

In 1850, Ravenel was elected to membership in the Academy of Natural

Sciences of Philadelphia and American Association for the Advancement

of Science (AAAS), both societies considered prestigious in the U.S. The

AAAS planned to hold its 3 annual convention in Charleston (SC) in 1853,

providing an opportunity for Ravenel to meet fellow botanists from all over

the United States and to present research data to this audience. Also in 1850,

the Episcopal diocese of South Carolina held its convention in Charleston;

Curtis attended and Ravenel invited him to visit him at home. In their days

together, they mixed with others of like persuasion—they agreed on the

traditional idea of a single creation of man, rather than multiple human

creations (including color).

Ravenel could also demonstrate his new French microscope. The cost was

$35-40, some $5 more than Ravenel had budgeted. But while he was delighted

at the result, neither he nor Berkeley could budge Curtis into purchase or use.

Nonetheless, the instrument brought Ravenel another rung upward to parity

with Curtis.

In 1848 Curtis published his first paper on North American mycology

in Silliman’s Journal (Curtis 1848). The journal's many typographical errors

prompted Berkeley to publish their subsequent papers in the LONDON

JOURNAL OF Botany and (eventually) in the AMERICAN JOURNAL OF SCIENCE

AND Arts (Shear & Stevens 1919). Three years later, Ravenel (1851) published

on fungi (the last in a series of three papers—mosses, lichens, fungi) and

issued the first century of FUNGI CAROLINI ExsiccarT1 in the following year

(Ravenel 1852).

If Berkeley was irked by typographical errors of American type-setters, he

was in good company. Young Elias Magnus Fries had confronted the same

problem years prior when Hornemann, Fries’s Danish mentor, offered to read

galley proof of Fries’s long OBSERVATIONES MyCOLoGIcae (Fries 1815) but

had difficulty reading Fries’s handwriting (Petersen & Knudsen 2015). For

Berkeley and Curtis, the introduction of the typewriter was still more than

Mycology in 19th century North America... 21

40 years distant (late 1880s) and even then, only a single copy was produced,

with errors creating problems, rectified only by inked-in notes. Complicating

this process, type was set manually, letter by letter, diacritical mark by

diacritical mark. Type-setting of scientific publications took unique skills, not

everywhere available.

Charles Wright had made several forays in all directions within Texas and on

to the Arizona Territory (1845-1852), when he finally started home, stopping

in St. Louis to visit with George Engelmann. His time back home was short,

though, for he signed up for the North Pacific Exploring Expedition under

the leadership of Ringgold and Rogers. Casting off June 1853, the ships made

their way from Norfolk, Virginia, through the Canary Islands and around

the Cape of Good Hope, an area with great botanical richness. After a stop in

Sydney, Australia, Wright sailed on to Hong Kong, the Bonin Islands, Japan,

and the Bering Straits, arriving in San Francisco in October 1855. There,

separated from the Expedition, Wright made his way home via Nicaragua.

Ante-bellum mycology

At 18, Peck entered the [New York] State Normal School in 1851 and took

his first course in botany. He was determined to be a botanist. Upon graduation

in 1852, he returned home to Sand Lake. “After graduation, he taught for a

season in the district school and spent some months as a dry goods salesman

in a general country store, but he quit because of poor health” (Haines 1978).

During 1852-1853, Peck taught 60 students at Schram’s Collegiate Institute at

Sand Lake, and in 1855 entered Union College in Rensselaer Co. One teacher

was Prof. Johnathon Pearson, known for his field trips. After graduating from

Union College, Peck taught for three years at Sand Lake Collegiate Institute.

Focusing his avocation on mosses, Peck established correspondence with Leo

Lesquereux, Sullivant, and others. Soon, he started a friendship with Elliott

C. Howe and Judge George W. Clinton (Atkinson 1918). He earned an M.A.

from Union College in 1862 and started teaching at the State Street High

School at Albany, thence to Albany Classical Institute, known more popularly

>»)

as “Cass’s.

Ravenel’s health began to deteriorate in 1852: chronic dyspepsia and eye

fatigue. To escape the summer heat, Ravenel spent the season in “the cool

hills of North Carolina” (Haygood 1983: 154) (“The Smoky Mountains”).

Returning in October, he resumed work on his 2° century of the FuNa1

22 ... Petersen

CAROLINI ExsIccatI, but poor health returned in a few months. By October

1852, he decided to move to Aiken, South Carolina, a newly “discovered”

resort town with cooler, drier air becoming popular with Charlestonians. His

Northampton estate was sold for $10,000. Part of Ravenel’s move to Aiken

involved selling and gifting numerous slaves: some staying with him and the

family, some to his brothers and three families to his father. Arriving in Aiken

in January 1853, he soon bought a farm and started building a house there. The

cotton raised on Ravenel’s new farm was termed Santee Cotton, of somewhat

lower quality than Sea Island cotton from the coastal islands of South Carolina

and Georgia. But significantly, there were peach orchards and vineyards on the

farm and slaves to do the planting and picking. Parenthetically, improvement

of crop rotation was used to bolster secession sentiments (a steady, profitable

income was necessary for a prospective separate country—even in the 1850s).

Three months later, Curtis visited Ravenel in Aiken. Ravenel was planning

a recreational trip to New York and Boston, and by July he was in Philadelphia.

There he visited the Schweinitz herbarium, where he was invited to take

duplicates, harvesting about 150. The second century of FUNGI CAROLINI

EXsICCATI was issued in late 1853, and in the meantime, Ezra Michener

had established a correspondence with Ravenel. Collecting assistance came

from Thomas Milton Peters (1808-1888) of Moulton, Alabama, and John

F. Beaumont (1825-?1865). By fall 1854, Ravenel was working on the third

century of his exsiccati, but over Christmas his wife Elizabeth suffered a series

of strokes and died in February 1855.

In 1854, a new correspondent, Job Bicknell Ellis (1829-1903; Fie. 10), saw a

notice of Ravenel’s ExstccaTI1 in a bookstore and wrote him. Ellis sought help

with his preoccupation, fungi, especially literature to help with identifications.

In the ensuing correspondence, which extended to Ravenel’s death many

years thereafter, Ravenel recommended Grevilles CRyPTOGAMIC FLORA and

Fries’s SystEMA MycoLoGicum, ELENCHUS FUNGORUM, and “his new work,”

EPICRISIS SYSTEMATIS MYCOLOGICI.

Ellis was a native of Potsdam, New York, almost as far north in the US as

possible and home to a horrendous climate—bitter cold and horizontal snow

off the St. Lawrence River for many months each year. Ellis, of unsubstantial

physique, apparently longed for a kinder climate (which remained a major

theme for years).

At the tender age of 16, the bookish Ellis taught “winter school” in 1845,

at Stockholm, St. Lawrence Co., (even farther north a short distance from

Mycology in 19th century North America ... 23

afi

Gores J

ELL

Fic. 10. JoB BICKNELL ELLIS

[Bot. Gaz. 15(11). 1890]

Potsdam) “... for which service he received ten dollars a month [half in

cash, half in grain, the last installment of which was paid twenty years later]

and “boarded around” [rotating boarding with families of the students]. In

1849, “having completed his academical course,’ Ellis entered Union College

but in his impecunious state also securing a teaching job in Charlton. By

1851, he graduated from Union College (age 22), moving to Germantown,

Pennsylvania, in June. By September he moved to Albany Academy (New

York) as classics teacher but with a growing interest in botany. After starting

24 ... Petersen

his own private school (and failing), he taught at Bartlett's Boarding School

in Poughkeepsie (1853). After three successive boat trips to New York City,

each intended to last three days to “do the city,’ he always returned on the first

train, suffering from a headache from the excitement of the city and the rigors

of his trip.

In his 3% century of FuNGr Caro.ini Exsiccati (spring 1855), Ravenel

included specimens from Charles Wright (1811-1885, specimens from

Texas), Michener (Pennsylvania, apparently having short-circuited the Curtis

connection), and Sartwell (New Jersey). Work began on the fourth century,

with plans for a fifth. That summer, Ravenel contacted Camille Montagne

in Paris, sending free sets of exsiccati. Montagne was effusive in thanks and

praise and (of course) solicited more, sending a collection of his papers in

repayment.

Over the winter of 1858, Augustus Fendler returned from his period in the

mountains of Venezuela with plant specimens including fungi and offered

the fungi to Curtis. Although not the sole plant collector in Venezuela over

the 19 century (Patouillard & Gaillard 1888), Fendler spent four years

(1854-1858) in the small town of Colonia Tovar, settled by immigrants

from Baden in 1843. Colonia Tovar was (and is) enclosed by cloud forests of

enormous biodiversity. Fendler collected avidly and his were among the first

specimens from this area to be studied by North American experts.

Apprised of Fendler’s fungus collections and confessing a shortage of

money, M.A. Curtis wrote several letters soliciting support to purchase

Fendler’s Venezuelan fungi. To Charles James Sprague in Boston: (letter: 11

Feb. 1856) the following: “Do you, or your Society, or anybody (individual

or corporate) feel disposed to take a share in a Mycological venture? Fendler

is now collecting plants in Venezuela, and has got a few hundred Fungi, to

which he will add more as opportunity offers. I can get possession of these,

by promising a remittance immediately upon their receipt. I should be glad

to have the elaboration of them, but am unable to make any offer, having no

extra funds. If individuals or Societies would agree to take sets, I could better

determine how to act in the premises. —Que dites vous?”

Curtis purchased the fungi for $100 before notifying Berkeley. After the fait

accompli and under cover of apologetic intrusion, Curtis inquired whether

Berkeley might be interested in Fendler’s specimens. Berkeley acceded,

and the collections were shipped. Fendler’s sojourn in Venezuela was not

Mycology in 19th century North America ... 25

otherwise exciting to Northern Hemisphere mycologists, and his expedition

was not even mentioned by Chardon and Toro (1934).

Fendler’s Venezuelan collections included about 3000 phanerogams, ferns,

and bryophytes and almost 400 specimens of fungi and lichens. Gray, Curtis,

Tuckerman, and Sullivant (all familiar names) bought sets, and to pay for

them, Gray and Curtis helped distribute specimens to subscribers (Todzia

1989).

During 1864-1871 Fendler returned to Missouri and settled in Allenton

(Spaulding 1909). While there, he resumed plant collecting and became

known to Henry Shaw, who hired him to work at the Missouri Botanical

Garden. Fendler curated the first botanical specimens in the newly organized

herbarium.

After Missouri and a few years in Europe, Fendler moved to Trinidad in

the Caribbean, where his brother lived. He died there in 1883, surely one of

the most intrepid collectors of his century, rivalled only by John Macoun and

Charles Wright.

In 1855, Curtis requested Ezra Michener to collect and send a specimen of

Botrytis (now Phytophthora) infestans, shown by Berkeley to be the cause of

the “potato blight” of the previous decade. Michener attempted to do so but

could not find a specimen. After some time, he wrote Curtis but, in addition

to explaining the lack of a specimen, went to some lengths to tell Curtis why

he was engaged in mycology. Michener wrote, “The inducements which led

me into the study of the fungi are a desire to supply a blank in the catalogue

of Natural productions of Chester County, which some of us contemplated—

and while I could not obtain the means to determine all the species myself —

I vainly hoped that my services as a collector of fungi might entitle me to the

assistance which I stood in need of in determining species. —Disappointed in

this hope, I have but little inducement to prosecute the study much farther—

yet there is a charm attending it which will probably [con]tinue as long as

Iam able to pursue it.” This was followed by: “How is the review of Schweinitz

progressing? I have been watching for it in the Journal of the Academy.”

As might be expected, Michener understood the importance of the

Schweinitz herbarium—first in Phanerogams and later especially the fungi—

and in the winter of 1855-1856 traveled to Philadelphia, not long after Curtis

(followed by Ravenel) had raided the Schweinitz fungi. Knowing that Curtis

had sent many Schweinitz splits to Berkeley, Michener (6 Nov 1855) wrote

Curtis, “In a few days I expect to commence arranging Schweinitz Fungi—

26 ... Petersen

and it would have been a great pleasure to have had the assistance of your

corrections during the examination of his species. —I shall however implicitly

follow his numbering and labeling [two words blurred] out in the SYNopsis

FUNGORUM” (Smith 1954).

Curtis must have been put off at what he thought was Michener’s impatience

for the Berkeley report on the Schweinitzians. Curtis's [lost] answer must have

been peckish, for Michener answered penitently, unctuously using the “Thee”

and Thy” of Quaker plain speech (Birch 1999). Michener took note of Curtis's

words: “You need not trouble yourself by sending me specimens too often.”

Wounded, Michener went further. "It was a kno[w]lidge [sic] of thy numerous

engagements in the Mycological field - apart from “business” proper, which

has frequently led me to feel and to express a fear of trespassing too much

upon thy time—and, so far as I can recollect, has prevented me from making

any claim upon thy attention farther than it may have been convenient to

thee to bestow—nor am I conscious of having manifested that “impatience”

of which thee speak.”

Perhaps to salve the tension produced by his words—or perhaps to rub

some salt in the wound—Michener added, “In working over the first eight

books of Schweinitz’ Fungi, I have been grieved to find a number of the

envelopes either missing or empty in addition to the many which [have been]

rendered valueless by the ravages of insects.—Perhaps the destruction will not

be so great after we get through the Hymenomycetes.” He was to find in the

Hymenomycetes that many of the specimens had been reduced to a minimum,

and he might have surmised that Curtis had taken enough both for himself

and for Berkeley.

Married twice, and having attained 92 years of age, Ezra Michener died

quite near home in Toughkenamon , Chester Co., Pennsylvania, on June 24,

1887.

J.B. Ellis’s search for an ameliorated climate entered a new chapter, and

in February 1855, the 26-year-old Ellis, accompanied by his sister, left

Poughkeepsie, NY, for Charleston, SC, with the intention of teaching school

there. Ellis called on one of the professors in the South Carolina College in

Columbia, SC, to seek information on the subject. Said Ellis, “I told him that

I had come South to teach and make a home there. He at once asked me

whence I came, and when I answered from New York [State], he replied,

while slowly swinging in his revolving office chair: “Well, the state of feeling

Mycology in 19th century North America ... 27

between the North and South is such that I doubt very much whether you will

succeed,” (Haygood 1983). And he didn’t. In March [1855] he and his sister

made their way from Charleston to Alexander, near Augusta, Georgia, where

he began teaching in an academy that fall. En route, he met with Ravenel,

who hosted him graciously. The conditions in rural Georgia were odious—so

much so that he returned to Potsdam (NY) after the first school term. Asked

whether there was one incident he considered interesting during his sojourn

in Alexander, Ellis opined, “No; the most interesting incident was to get away,

that is very distinct, even now,’ (Anderson 1890). By 1856, he was Principal

at Canton Academy, St. Lawrence Co., (a short distance SW of Potsdam). “...

and on the eighth of April, 1856, an event took place which made it possible

for him to do the enormous and valuable work he has since done for American

mycology. This was his marriage to Miss Arvilla J. Bacon, who has been a

faithful partner in all the vicissitudes of life, and a constant and painstaking

assistant in his mycological work for the past thirty-four years,’ (Anderson

1890). In 1863, at the height of the Civil War, Ellis and his wife were teaching

in the public schools of Potsdam village.

Even before the armed conflict of the Civil War, emotional involvement in the

underlying issues was often strong, as shown by the discrimination J.B. Ellis

confronted during his attempt to make a career in the south.

Another example of those who did not fit in was the Macbride family of

Rogersville in East Tennessee, strict abolitionists in a state that was eventually

to secede to join the Confederacy. Their lives were apparently subject to

harassment, and they felt forced to move—not due north, but northwest, to

Iowa. That state, although settled by pioneers from all directions including

south, was predominantly sympathetic to the Union. With an agriculture-

based economy, Iowa furnished foodstuffs as well as 75, 000 volunteers to

the Union army. The Macbrides moved from town to town, finally settling

in Princeton in Scott County. Certainly they did not dream that son Thomas

would become a mycological leading light.

On August 12, 1856 Henry Ravenel (44 years old) married 25-year-old

Mary Dawson. Ravenel’s increasingly weak eyesight made microscope work

distasteful (paralleling the later problems of Curtis). In 1857, Ravenel donated

his higher plant herbarium to the Elliott Society of Charleston. As early as late

1859, Ravenel read Darwin's ORIGIN OF SPECIES, vacillating on the question

of immutability of species, but accepting certain points. In practical terms, in

28 ... Petersen

Ravenel’s mind his work on collecting and cultivating grape vines was more

important than the philosophical premise of natural evolution.

Meantime, Curtis transferred to Hillsboro, NC, where a larger church afforded

less time for botany, and he had thoughts of withdrawing from mycology

altogether.

But Curtis could not divorce himself from floristic botany. Alvan

Wentworth Chapman, M.D. (1809-1899), of Apalachicola, FL, approached

Curtis to write a manual of the southern flora to match Gray’s manual for the

northeast. Many years later, Chapman wrote to Lamson-Scribner, “I believe

the [Chapman] FLora owes its existence to a suggestion of mine to Dr. Curtis

about the year 1856 that we needed for the South a work something like what

Gray had made for the North, and that he (Curtis) was just the man to do it.

But at that time his hands were full of toadstools and he was rusty among the

Phanerogams, and so turned over the matter to me, while promising every

assistance in his power if I would undertake the job,’ (Lamson-Scribner 1893:

331). Chapman's first edition was published in 1860 (Chapman 1860).

During the war (1856, 1857, 1861, 1865) and after (1867), Charles Wright

commuted back and forth to Cuba, each time returning with new collections

of tropical plants and fungi. Gray parceled out the specimens, “As to the lower

cryptogams, Mr. Wright's very rich collections were distributed in sets and

published by specialists: the FuNc1 by Berkeley and the late Dr. Curtis...,”

(Sargent 1889, Berkeley 1863, Berkeley & Curtis 1863). Except for short

forays, Wright spent the rest of his days working with his own specimens in

the Harvard Herbarium. As written by Gray: “By degrees [Wright's] strength

was sapped by some organic disease of the heart, which had given him serious

warning; and on the 11" of August [1885, in Wethersfield, Connecticut] he

suddenly succumbed, while making his usual round at evening to look after

the domestic animals of the homestead.”

Gray summarized Wright's physical appearance: “Mr. Wright was a person

of low stature and well-knit frame, hardy rather than strong, scrupulously

temperate, a man of simple ways, always modest and unpretending, but

direct and downright in expression, most amiable, trusty, and religious. He

accomplished a great amount of useful and excellent work for botany in the

pure and simple love of it; and his memory is held in honorable and grateful

remembrance by his surviving associates” (Sargent 1889). Wright's collections

formed the basis for scores of floras and monographs and offered the botanical

Mycology in 19th century North America... 29

community glimpses of variation attainable in no other way than personal

acquaintance.

The war years

Veterans of the American Revolution still alive into the 1860s were about

to see a new, albeit internal, rebellion.

Ravenel slowly evolved from unionist to southern nationalist, while with

the death of “the great compromiser” (Daniel Webster), polarization and

simmering tensions tightened. The problem of slavery in the territories

became critical. Ravenel believed sentiments were “unanimous” that South

Carolina would secede if Lincoln were elected President, and by November

he was ready to see South Carolina take the lead to secession. Tuckerman and

Gray worried from afar; Ravenel hoped that their friendship could continue

regardless of politics (Stevens 1919).

Early 1861 brought secession after the attack on Fort Sumter in Charleston

harbor. Charleston was blockaded by the Union navy. “The blockade

cut southern scientists off from contemporary European and northern

contributions, and no one could concentrate properly on research when

the fate of the entire country was being played out on the fields of Virginia.”

(Haygood 1983: 236). In fact, the northern mycologist J.B. Ellis was part of

the blockading fleet. No transatlantic communication was allowed. Even

postal service south to north was impossible, and the southern mycologists

were marooned. Ravenel’s health further deteriorated, including periodic

deafness. Nonetheless, the fifth fascicle of the FUNGI CAROLINI ExsICCATI

was completed and with it, “final withdrawal” from botany.

J.B. Ellis served in the navy during the Civil War. The latest innovation

in ship building was the wooden hull covered with a layer of iron plates (i.e.

“ironclad”), coupled with steam-powered propulsion, introduced in sea battle

during the war.

Another fledgling mycologist would see less action. Albert Nelson Prentiss

(1836-1896; Fic. 11), had been born on a farm near Cazenovia, New York, a

location frequented later by Lucien Underwood. A product of local schools,

in 1861 Prentiss graduated in a class of seven from the new Michigan State

University. With war imminent, all seven enlisted in the army and were

assigned to a camp. But in spite of secession and armed violence, the camp

was closed, and group cashiered. The War was fought without them.

30 ... Petersen

Fic. 11. ALBERT NELSON PRENTISS

[Bot. Gaz. 21(3). 1896]

Unbeknownst to Ravenel or Curtis, another young man destined to influence

American mycology had entered the population: Andrew Price Morgan

(1836-1907; Fic. 12). “From this position [teaching in Dayton, Ohio, High

School] he went into the Union army, first in the 84th Ohio Infantry—

carrying a stoutly bound copy of [Asa] Gray’s MaNuat in his knapsack—and

again as First Lieutenant in the Fourth Independent Battalion O.V.C. Being

a member of the Dayton Light Guards he was a well-drilled soldier and did a

great deal of work drilling volunteers. At the close of his army service he was

very ill with typhoid fever” (Kellerman 1907).

In 1862, a bill written by Vermont congressman Justin Smith Morrill

(1810-1898) was made law. The “Morrill (or Land-Grant College) Act”

Mycology in 19th century North America... 31

Fic. 12. ANDREW PRICE MORGAN

[Mycol. Notes (Lloyd) 31. 1908]

granted each state 30,000 acres per congressional seat (thus favoring the

more populous northern states) for establishment of schools specializing in

“agriculture and the mechanic arts.” Some states used the land for its chief

purpose, while others sold the land and turned the funds over to extant state

or private colleges to create colleges of agriculture and engineering*. Military

training was required in the curriculum, not least because the act came in the

midst of the Civil War.

Yet another resident of Chester County, Pennsylvania, was Abraham Robinson

Mcllvaine (1804-1863), a prominent Quaker three-term congressman. In

1840, a son was born to the McIlvaine family: name, Charles (1840-1909;

Fig. 13). Everything written about Charles points to a youth without financial

restrictions, and the word “indolent” perhaps could be attached to him, at

* Numerous present-day colleges and universities still bear this notation (ie. Texas A&M,

Louisiana State University, Michigan State University, North Carolina A&T, etc.).

32 ... Petersen

Fic. 13. CHARLES MCILVAINE

[Harshberger. 1899]

least until he was seized by a life-centering crusade—the edibility of fleshy

fungi. According to McIlvaine himself, the conversion event occurred in 1877

(age 37) as he rode through the forests of West Virginia.

Having been “educated” at the local county schools to the age of 11, Charles

then attended Northwest Grammar School in Philadelphia for two years. This

seems to have been the extent of his schooling. “He was not a university man

and took no academic degrees, which was later to prove an embarrassment to

the acceptance of his work in mycology” (Bowser 1972).

At his father’s urging Charles raised a company of volunteers to support the

Union cause, becoming attached to 97" Regiment of Pennsylvania Volunteers.

For his efforts, he mustered as Captain. Later, he was offered promotion to

Mycology in 19th century North America ... 33

Major of the First South Carolina Cavalry (Colored), but demurred, citing

ill-health. (While African-Americans were allowed to enroll in the Union

military, their officers were strictly Caucasian.)

Called up twice to serve in the military in 1863 and 1864, Ravenel was

excused from service due to poor health. Financially, at war's start Ravenel’s

principal income was from interest-producing investments (bonds of South

Carolina banks). After secession, as these debts were converted into inflated

Confederate dollars, Ravenel repeatedly put the income back into Confederate

bonds; as it turned out, an unwise decision.

During the War, with foodstuffs becoming scarce, Ravenel extolled

mushrooms as a food source. He and Curtis both enjoyed mushrooms

pre-war, but now they became more exigent. Ravenel brought wild edible

mushrooms to monthly meeting of the St. John’s Agricultural Society; others

brought fruits or vegetables.

The Curtis family was also in dire straits. Not only was the Union General

Sherman's March to the Sea within a few miles of Hillsborough, but two sons

were in the Confederate army. One, nicknamed “Non” came home for a brief

leave, but upon returning to duty, was killed at the battle of Bentonville NC.

Mary Curtis had become a gold star mother before the term was established,

but her grief had to be contained as she ministered along with Moses to their

congregation, also suffering their own calamities.

Charles Edwin Bessey (1845-1915; Fic. 14, next page), a farm-boy, was

home-taught but received a certificate to teach. While he attended Seville

Academy (Seville, OH), his father died in 1863 during the midst of the War,

and Charles (age 18) began teaching in Wadsworth, Ohio. In 1866, Bessey

entered Michigan Agricultural College, graduating in 1869. His advisor was

Prof. Albert N. Prentiss, later Botany Department Head at Cornell University,

and Bessey turned from engineering to botany. In 1869, Bessey became an

instructor at Iowa State College of Agriculture in Ames, Iowa. Awaiting him

there was a young student, Joseph Charles Arthur, eager to study botany and

destined to play a major role in mycological research.

In 1864, Yankee infantry approached Charleston and St. John’s Parish was

abandoned by Confederate forces. Ravenel remained at his father’s Pooshee

plantation to help the family prepare for the Union army. All valuables were

buried (some were stuffed in mattresses) and four months of provisions were

34 ... Petersen

Fic. 14. CHARLES EDWIN BESSEY

[Amer. J. Bot. 2(10). 1915]

given to the slaves. On March 1, raiding parties reached Pooshee: “Horses,

wagons, buggies, harness, the contents of the smoke house and root cellar,

wine, guns, and poultry were all taken, but no one was hurt, no buildings

were burned, and Ravenel was able to find comfort in the loyalty exhibited by

the Pooshee slaves during the incursion” (Haygood 1983: 249). The home in

Aiken was raided by renegade Confederate troops but largely left intact.

In spring 1865, the Confederate Army surrendered, and the War ended.

General Sherman's march left a swath of destruction through Georgia and

South Carolina. Confederate money was plentiful but useless. Hampton Hill

(Ravenel’s Aiken home) was intact and the orchards and vineyards untouched

(Haygood 1983: 257-259). Other complications abounded, however: the

Mycology in 19th century North America... 35

peach crop was bountiful, but with the obliteration of transportation, the crop

could only be used locally. The situation extended to the crops of 1866 and

1867. By 1868, the crop could be shipped, but picking was desperate, and the

crop arrived in New York bruised and low-priced. As early as 1865, Ravenel

decided he should pay his newly freed slaves a salary, not “in-kind” housing,

food, medical treatment, etc., but he had no money to pay them. So even

though he and his family were safe, relatively healthy, and with a roof over

their heads, life was not as comfortable as pre-War and useable money was

tight. In the fall of 1865, Ravenel offered to sell important works from his

library and gave his microscope to the family doctor in lieu of payment for the

year, while spare fascicles of the exsiccati went up for sale.

Termination of hostilities

At War’s end, American mycology remained in the hands of its two

leading lights, Curtis and Ravenel. A third man, J.B. Ellis, was enthusiastic

but otherwise occupied as a teacher. Born before or during the War, over the

upcoming quarter-century, several others would emerge. A review of their

circumstances indicates two major pathways to their careers: 1) almost all

were born on family farms, anticipating family inheritance in their future; and

2) most made their way through a series of teaching jobs—public schools or

academies—probably without a premonition of their eventual fate. Whether

through a farm-boy’s exposure to nature or an acquired interest in plants,

interest likely progressed from nature to plants, thence to cryptogams or

phytopathology and on to mycology.

It was in these years that electricity for lighting rapidly replaced gas, at least

in the industrialized north.

Canada, of course, was little involved in the “War Between the States.” In

Ontario, John Macoun had been captivated by plants as a youth, collecting

and identifying as many as possible without guidance. “Although almost

entirely self-taught, his reputation as a botanist grew, and just ten years after

leaving the farm, in 1869 he was offered the Chair of Natural History in

Albert College, Belleville”? (McGeown undated). By this time he was married

(1 January 1862) with a family. Nonetheless 1872 found Macoun on a

“botanical holiday” steaming across the Great Lakes. By chance, he met Sanford

Fleming, chief engineer for the proposed Canadian Pacific Railway. Fleming

offered Macoun a place on an upcoming expedition to lay out the route for the

coast-to-coast route, which he gladly accepted (there is no record of his wife's

36 ... Petersen

reaction). “Despite the privations of this journey, Macoun fell in love with the

wilderness, and over the next ten years [1872-1881] made a number of long

and strenuous journeys into the North-West Territories, the last two on behalf

of the Canadian government. He published the observations made on these

journeys in MANITOBA AND THE GREAT NORTH-WEST in 1882 (McGeown

undated). In 1881, Macoun was appointed Botanist to the Geological and

Natural History Survey of Canada and moved to Ottawa in 1882. “He became

a charter member of the Royal Society of Canada and wrote a CATALOGUE

OF CANADIAN PLANTS and later a CATALOGUE OF CANADIAN Birps. In 1886,

Macoun was sent to England as a Canadian representative to the Colonial

Exhibition. “He suffered a stroke in 1912, but fortunately recovered and later

that year moved to Vancouver [British Columbia] .... He was engaged in

writing his autobiography when he died of heart failure at Sidney, Vancouver

Island, 18 July 1920” (McGeown undated).

John Macoun must take his place as the dean of collectors. His journeys

preceded those of Augustus Fendler, Charles Wright, and others by some

years, as an individual seemingly not fit to settle down for prolonged quiet

but instead given to exploring hitherto unexplored and often uncivilized

wilderness.

The War was over, and A.P. Morgan returned to teaching in Dayton, Ohio,

but over the ensuing years, he suffered from “confinement in the school

room” (Kellerman 1907). This seeming claustrophobia (who is to say that it

was not the PTSD of 150 years later?) became debilitating and forced Morgan

to change his life. He became a book-publisher’s representative in school

systems in Wisconsin and Minnesota, far away from home. During this time

(1870, age 34) he met and married Laura Vail of Pomfret, Vermont. In his

seventh year with the firm, he suffered “a partial paralysis [which] closed his

career [with the book firm]. He could neither read nor write for two years”

(Kellerman 1907). He and Laura moved to her family home between the

Green and White Mountains of New England.

The circumstances ofhis “more than three years of ... delightful recuperating

of health” are unknown, but it was then that became fascinated with fungi.

Surely with some mentoring, “he procured a copy of HyYMENOMYCETES

EuropaE! by Elias Fries” for $8.25 (Kellerman 1907) and began collecting

agarics in earnest. His early gatherings were identified by C.H. Peck in

Albany, NY, a fine person on whom Morgan could rely. Tuckerman and Frost

also helped. He traveled to Kentucky for more collecting (Underwood 1899).

Mycology in 19th century North America ... 37

Around 1877, again feeling fit, Morgan (and wife) returned to Dayton (Ohio)

where he became principal of the District High School. In the same year he

published his first contribution in mycology (Morgan 1877).

C. L. SHEAR

WITH JOHN DEARNESS

Gatlinburg, Tennessee, August, 1939

Fic. 15. CORNELIUS LOTT SHEAR with JOHN DEARNESS

[Mycologia 49(3). 1957]

Another New York State farm-boy, Cornelius Lott Shear (1865-1956; Fic. 15)

matriculated at Albany State Normal School (graduating in 1888), where one

38 ... Petersen

of his instructors was E.A. Burt, with whom Shear would again interact some

years later. While at Albany State, he became acquainted with C.H. Peck, by

then well ensconced as the New York State Botanist. They remained friends

through Pecks life.

As a young adult, Shear held numerous teaching jobs but never felt any as

a calling—only as a means to a “very unsatisfactory” living. Married in 1890,

he taught in Alcove, New York. Avid collecting produced material for three

centuries of exsiccati, NEw YORK FunGI. In 1894, he moved with his family to

Osborne, Kansas, the home of his in-laws.

Mrs. FLora W. PATTERSON IN HER LABORATORY

Fic. 16. FLORA WAMBAUGH PATTERSON

[Mycologia 21(1). 1929]

As 19" century mycologists’ timelines go, Flora Wambaugh Patterson's

(1847-1928; Frc. 16) will seem familiar. Although born in Columbus, Ohio

(a town rather than on a farm), she was educated in public schools, but at

an age when most women undertook either a life of marriage and family or

a “womans” career (i.e. nurse, librarian, school teacher), Flora Wambaugh

continued her education, graduating first from Antioch College (a small

liberal-arts institution) in 1860 before earning her B.A. from Wesleyan

College in 1865 and M.A. (also from Wesleyan) in 1883. Marriage to Capt.

Mycology in 19th century North America... 39

Edwin Patterson in 1869 not only punctuated the interim between degrees

but also added obligatory migration with husband and rearing their family of

three boys. A few years after marriage, Capt. Patterson was seriously “injured

in a steamboat explosion from which he never recovered but lived ten years,

a helpless invalid” (Charles 1929).

Fic. 17. THomas Huston MACBRIDE

[Mycologia 26(5). 1934]

Thomas Huston Macbride (1848-1934; Fic. 17) received his B.A. (1869) and

M.A. (1873) from Monmouth College, Illinois. From 1870-1878 he taught at

Iowa's Lennox College, and in late 1875, married his former Lennox student,

AO ... Petersen

Harriet Diffenderfer. In 1878, he moved to Iowa State University in Ames,

where he stayed for 56 years.

During his lengthy Iowa State career, Macbride published on myxomycetes,

but his research time must have been limited, for he served Iowa State

University as Professor, President, and President Emeritus. In his later years,

the faculty was joined by George Willard Martin (1880-1971) who carried

the myxomycete torch into another generation.

“An eminent scientist, a profound scholar, an inspiring teacher, a brilliant

public speaker, a public-spirited citizen, a lover of the beautiful in everything

material as well as moral and ethical, a man of high character and a true

friend, Dr. Macbride presented a rare combination of high qualities which

made his well-rounded life an inspiration and a benediction.” (Shimek 1934)

Fic. 18. GEORGE ENGELMANN

[Neilans. 1963]

Mycology in 19th century North America... 41

In spring 1866, a sliver of light shone for Henry Ravenel. George Engelmann

(1809-1884; Fic. 18) in St Louis was engaged in a monograph of Juncus and

asked Ravenel for specimens, agreeing to pay Ravenel for weeks of collecting

around Charleston. Engelmann, a highly credentialed physician and German

émigré, was also an educated botanist of considerable financial means.

Although his medical practice took time, he traveled in pursuit of his avocation

and published taxonomic papers on selected plant groups (Humphrey 1961).

Fic. 19. MORDECAI CUBITT COOKE

[English. 1981]

Engelmann’s request and offer struck a spark in Ravenel, whose experience

and expertise could finally be put to financial use. Ravenel undertook to

write to Gray, Curtis, Engelmann, Berkeley, and others about collecting

42 ... Petersen

phanerogams and seeds for sale. Berkeley spread the word, and Mordecai

Cubitt Cooke (1825-1914; Fic. 19) was enthusiastic. Cooke had already

published three books: MANUAL OF BOTANIC TERMS (1862), PLAIN AND Easy

ACCOUNT OF BRITISH FUNGI (1862), INDEX FUNGORUM BRITTANICORUM

(1865). Also, in July came a letter from Thomas Meehan (1826-1901), a

Philadelphia nurseryman. He desired seeds of native southern forest trees;

Ravenel inquired about selling seeds to European individuals and gardens.

Meehan responded positively, but also encouraged Ravenel to consider

starting a nursery garden along the lines of André Michaux. Ravenel (after

some qualms) agreed and received a large assortment of root stock from

Meehan and a collection of rose cuttings from a local neighbor. He was on the

cusp of yet another new career.

In 1865, only months after cessation of armed conflict between North and

South, Curtis took up his pen to attempt to resurrect pre-War contacts: Asa

Gray, M.J. Berkeley, Spencer Fullerton Baird (1823-1887) of the Smithsonian

Institution, and others. He soon caught up with news of botany and botanists.

Gray’s response was especially anxiously awaited, for Curtis could not be sure

of his welcome in Cambridge. Not only had Gray news, but he also reported

on Charles Wright's fungus collections from Cuba (1865, but back there again)

and other Mexican collections from Orizaba. All this was promptly shipped,

and Curtis spent many hours examining, glad to be involved once more

in fungi (Berkeley & Berkeley 1986). Baird brought news of the proposed

transfer of the Smithsonian's herbarium to the Department of Agriculture, a

move with which Curtis disagreed. (Petersen 1979).

Curtis's mind was still on edible fungi, however, and “In a letter to Berkeley,

of England, he wrote: ‘In October 1866, while on the Cumberland Mountains

in Tennessee, although with little leisure for examination during the two days

spent there, I counted 18 species of edible fungi. Of the four or five species

that I collected for the table, all who partook of them declared them most

emphatically delicious. On my return home, while stopping a few hours at a

station in Virginia, I gathered eight good species within a hundred yards of the

depot. And so it seems to be throughout the country. Hill and plain, mountain

and valley, fields and pastures, swarm with a profusion of good nutritious

fungi which are allowed to decay where they spring up, because people do not

know how, or are afraid, to use them.” (Lamson-Scribner 1893: 323).

In November 1866, former Confederate General Daniel Harvey Hill

(1821-1889), then editor of a magazine in Charlotte (NC), wrote to inquire

Mycology in 19th century North America ... 43

whether Ravenel would write articles on a southern agricultural theme at

$3 per page. Given this added opportunity to make ends meet, Ravenel

penned several articles until the magazine was sold in 1868. Nonetheless, in

late 1867, he started digging up his peach trees for sale. At the same time, a

complicated arrangement for sale of fungi and phanerogams to M.C. Cooke

was struck (English 1987). In his outgoing letters, a thick religious veneer

concealed Ravenel’s post-war anger and humiliation (to Curtis 11 Jun 1868:

“In a word, I feel contented, satisfied and trusting.”), but his journal revealed

his rancor at his lifestyle change.

A decade or more after the Civil War, mycology in the fledgling United States

consisted of a handful of men for whom the fungi were an avocation: North

Carolina clergyman, southern plantation owner, Pennsylvania country

doctor, and New England shoemaker. The last was Charles Christopher

Frost (1805-1880) of Brattleboro, Vermont, a third-generation cobbler.

After receiving corporal punishment from a teacher in 1820, Frost abruptly

gathered his books and departed school, ending his formal education at age

15. But his curiosity and mental acuity were not satisfied, and over the years he

ravenously digested books on mathematics, chemistry, physics, meteorology,

geology, and botany and gathered collections of insects, shells, and plants.

The Berkeley and Cooke manuals served as mycological foundation, but

neither took up the American mycota. He and his friend, Charles J. Sprague

(later to become the Curator of Botany at the Boston Society of Natural

Science), together bought a French microscope. By middle age, Frost could

be counted as an authority on local plants of southern Vermont and adjacent

Massachusetts, and somewhat later, he began to look at and study fungi.

Even before Charles Horton Peck was hired by the New York State

Museum, Charles Frost was mentoring him in mycology. In August 1866, a

61-year-old Frost wrote the fledgling Peck: “There is no elementary work on

the fungi published in this country as yet that I am aware of. For their study

you will have to depend on foreign wholly and they will necessarily be in

Latin French & German & if you are acquainted with these languages I can

recommend the following: Corda, EINLEITUNG; Nees, SYSTEM; Fries, SYSTEMA,

EPICRISIS, MONOGRAPHIA; Rabenhorst, DEUTSCHLAND CRYPTOGAMEN

FLora; Berkeley, BRiTIsH FuNGI.’ Frost then added his recipe and method

of preserving fleshy fungi. This from a third-generation shoemaker from

Brattleboro, Vermont! Of course, having taught classics, including Latin (and

probably Greek), Peck could use Corda, Nees, and Fries with minor effort.

44 ... Petersen

The following year, Peck was hired by New York State to “fill the herbarium,”

and Frost continued his advice. Peck, in turn, was in touch with the aging

Moses Curtis, who professed thoughts of writing his own mycological manual

(Burnham 1919).

Strangely, none of Peck’s biographers mention the Civil War, which raged

during the years of Peck’s teaching career. The immunity of Peck remains

unexplained.

Charles Edwin Bessey (1845-1915), with degree in hand in 1869, became an

Instructor at lowa State College of Agriculture at Ames, Iowa, one of the early

Land-grant schools. Perhaps pivotal to his career and philosophy, he spent

three months with Asa Gray at Harvard in 1872-1873 and again in 1875-

1876. In 1874, Bessey gave a series of lectures at the University of California,

and in 1879, he earned a Ph.D. from Iowa State. Bessey was recognized for

his well-known diagram depicting the evolution of angiosperms (known as

“Bessey Cactus”). His contribution to mycology was minor (with the major

exception of the guidance he gave to student J.C. Arthur) but he was in touch

with all the botanical leaders of the time. A second notable achievement was

Charles's son, Ernst Athern Bessey (1877-1957), who in the next century

would become one of North America’s leading mycologists.

William Ashbrook Kellerman (1850-1908; Fic. 20) was another farm-boy

from the American mid-west, this time from Ashville, Ohio. Of his early

life little has been written, but the Civil War was fought during his formative

years. He progressed through “country schools” and at 17 was a teacher

(Humphrey 1961). How he came to matriculate at Cornell cannot be related,

but he graduated in 1874, precisely at the time that Albert Prentiss arrived to

chair the Botany Department, and Kellerman was likely to have had classes

from him. How he secured a teaching position at the State Normal School

in Oshkosh, Wisconsin, is similarly unreported, but he remained there for

five years (Anonymous 1908). Married in 1876, Kellerman determined

that he needed European exposure and embarked on a two year-residence

(1879-1881) in Switzerland, earning a Ph.D. from the Polytechnic in Ziirich

(Willard 1909). Upon return, Kellerman taught at the State Agricultural

School (later the University of Kentucky) in Lexington.

Later in 1881 an unusual event transpired. Ohio State University inaugurated

a professorship in biology. At least three applicants were considered: Andrew

P. Morgan, Joseph C. Arthur, and William A. Kellerman. Of the three,

Mycology in 19th century North America ... 45

Fic. 20. WILLIAM ASHBROOK KELLERMAN

[J. Mycol. 14(2). 1908]

Morgan, who had the thinnest credentials, was selected but—mysteriously—

withdrew after three months under now-unknown circumstances but with a

commendation from the Board of Trustees (Ellett 1970). The most probable

46 ... Petersen

cause may have been poor psychological health, but in the meantime,

Kellerman had moved to Kansas and Arthur to Purdue. Kellerman’s seven

years in Kansas, three on faculty and four as Kansas State Botanist, included

teaching and research, the latter almost strictly phytopathological.

Fic. 21. ELAM BARTHOLOMEW

[Mycologia 27(2). 1935]

Perhaps a stereotype of the rugged, stalwart, quiet but competent, studious

American homesteader of the late 19th century is Elam Bartholomew

(Fic. 21). Although he is now known chiefly as a taxonomist of plant

pathogens, his influence was much more widely felt as a gatherer and publisher

of exsiccati and for, as others of his ilk, his wide correspondence with the

small group of established mycologists. Born on 9 June 1852 in Strasburg,

Pennsylvania (situated between the Lancaster County Amish and Chester

Mycology in 19th century North America ... 47

County Quakers), Bartholomew soon moved with his family first to Ohio and

shortly thereafter to Illinois. In his teens, Bartholomew decided to become a

teacher, but one pre-requisite for a teaching certificate was a general knowledge

of botany - something theretofore outside his worldview. He obtained a

copy of Asa Gray’s Lessons IN Botany and before long was able to pass the

qualifying examinations. Having taught for only a single school year, in 1874

Bartholomew moved to Kansas where he homesteaded a patch of land in the

north-central portion of the state, but not before he was betrothed to Rachel

Isabel Montgomery back in Illinois. In a whirlwind trip, they were married

in Illinois in 1876 and immediately returned to Kansas. Their existence was

without frills—a sod house where they raised seven children—but in addition

to farming, Bartholomew was a school and town officer, active in Republican

Party politics, an elder in the Presbyterian Church, and a district court clerk

(University of North Carolina Herbarium 2018).

Despite his many other activities, his brush with botany had established

roots. He became an earnest collector and made a herbarium with very nearly

all the green plants of his neighborhood. He corresponded with several

established botanists, in particular W.A. Kellerman, then of the Kansas State

Agricultural College. For the rest of his life, Bartholomew would relate the

following. “One day in July 1885, Dr. Bartholomew, like Cincinnatus of old,

was plowing in his field when W.A. Kellerman, then professor at the Kansas

State Agricultural College, came to see him. The two had never met but had

corresponded much concerning phanerogamic matters. After some time

Dr. Kellerman stooped down, pulled a leaf from a weed, and straightening up,

said ‘Bartholomew, why don't you study something that is really interesting?

Look at this leaf? It was an Amaranthus leaf and was covered with white

pustules (Albugo) on the lower surface. Bartholomew did become interested

and many times in later years as he developed his mycological herbarium, he

was heard to remark: “The plucking of that leaf by Professor Kellerman from a

weed in my cornfield marked a turning point in my life” (Bartholomew 1935).

In one of his columns for General Hill, Ravenel had extolled the use of the

legume Lespedeza striata as a rotation crop for its ability to return fertility

to the soil. Over the winter of 1868-1869, Ravenel received correspondence

from Dr. John Shaw Billings (1839-1913) of the library of the federal

Surgeon General, forwarding an offer for Lespedeza seed of six pounds at

$5 per pound. But Billings was also interested in cryptogams and he and

Ravenel discussed the contemporary French and German ideas about

48 ... Petersen

causation of disease by fungi. “They [the ideas] are very amusing certainly—

and if one were to believe Hallier [Ernst Hallier 1831-1904]—a Penicillium

spore and the Darwin theory will account for pretty nearly every natural

Phenomenon hitherto observed” [Billings to Ravenel, 6 Jan 1869; Ravenel

papers, Clemson].

Billings also informed Ravenel of a small expedition to Texas to investigate

the cause of a cattle disease. The sponsoring veterinarian wanted a botanist

to look at possible fungal causes. Ravenel accepted the appointment at $300

per month plus expenses. His itinerary started in Washington, D.C., thence to

New Orleans and Galveston to Houston. He finally returned to Aiken in May

of the following year (1870). Ravenel’s Texas plant specimens went to Torrey

and others; his fungi, as pre-arranged, went to Curtis and Berkeley.

Evidence that all was not amicable in the wake of the War is reflected in a

remark by Elliot C. Howe (1828-1899) of New York State (another mentor to

Charles Horton Peck), who had been approached by Ravenel in 1868. Howe

confessed to Peck, “I will send [specimens for identification] to Ravenel this

week, though I confess to a feeling akin to horror at the thought of addressing

a South Carolinian! But our science must not suffer though we are compelled

to invoke the aid of Le Diable himself” (Haygood 1983: 344; Petersen ined.).

Lucien Marcus Underwood (1853-1907; Fic. 22) was born in Woodstock,

New York, not far from the homes of J.B. Ellis and C.H. Peck. As a child he

made collections of grass leaves and of rocks and minerals (Hesler 1975). “As

he learned to read and write, he became interested in collecting papers and

documents of all kinds and would prepare lists of celebrities and of events.

Later, when his school days brought him in touch with natural science subjects,

the house became the repository of rocks and minerals and the laboratory for

such physical and chemical experiments as his ingenuity could devise” (Curtis

1908). He was clearly too young to be affected by the Civil War. From age

5 to 11, Underwood attended summer sessions at “the brick schoolhouse,” and

at 11 he became a farm hand. At age 15, he attended Cazenovia Seminary.

The school was three miles from home and, of course, he walked both ways.

In 1868 or 1869, he was “forced to take over the home farm” (Hesler 1975):

typical|ly] farming in summers, lumbering in winters (Curtis 1908).

The year 1873 found him back at Cazenovia Seminary. Although he had yet

to overcome difficulties in public speaking and writing, a teacher suggested

college and later that year Underwood enrolled at Syracuse University. During

summers he worked on farms in the area, saving enough money to visit

New York City and the Philadelphia Centennial Exposition, his first peek at

Mycology in 19th century North America... 49

hirer ine Ub, Uudrrumre§

Fic. 22. LUCIEN Marcus UNDERWOOD

[Bull. Torrey Bot. Club 35(1)]

the outside world (Curtis 1908). At Syracuse, Underwood “... soon became

dissatisfied with the scientific course because [it was] deficient in the amount

of work required and after seriously considering the advisability of completing

the college work in three years, he finally decided to enter the Latin-scientific

course although this necessitated the preparation of six books of Virgil, four

orations of Cicero, Sallust, Roman History, and Latin Prose.” (Curtis 1908) [His

cause for change must have been rare in that day and even rarer in 2019.]

50 ... Petersen

Upon graduation in 1877, Underwood accepted principalship of the

Morrisville Union School for an annual salary of $700. In 1878, he completed

one year of post-graduate classes at Syracuse and purchased Hooker's Synopsis

Fiticum. During the 1878-1879 school year, Underwood taught natural

science back at Cazenovia Seminary. In winter sessions he taught 14 classes a

day, receiving a salary of $600 for the year. In these years His interest, which

at the time focused first on ferns and then on hepatics, gradually migrated to

fungi. 3

‘

- .

a ,/

“—

Fic. 23. FRANKLIN SUMNER EARLE

[Mycologia 2(1). 1920]

Born 4 Nov. 1856, on a large fruit farm near Dwight (now in Livingston Co.,

Illinois), Franklin Sumner Earle (1856-1929; Fic. 23) showed interest in

plants and animals early on, supported in his hobby by his mother. At 16 he

prepared for college and attended the University of Illinois, but his studies

were interrupted periodically by exigencies at the family farm, and Earle spent

four years assisting his father uninterruptedly. Again in 1878 he attended the

Mycology in 19th century North America... 51

University and came under the guidance of Prof. Burrill but had to withdraw

within a year. (Shear 1929) This brush with college-level education was to end

his formal education.

In 1892, Earle was appointed superintendent of one of the branches of the

Mississippi Experiment Station, but his penchant for research led to a position

as assistant pathologist in the USDA herbarium in Washington (Chardon

1929). In this position, Earle was primarily a phytopathologist, a discipline

that engaged him for the rest of his life. He had also been introduced to

tropical climate and flora with their pathological susceptibilities.

Curtis Gates Lloyd (1859-1926; Fig. 24) was also born just before the war on

17 June 1859 in Florence, Kentucky, moving with his family to Crittenden

_—

—

Fig. 24. CURTIS GATES LLOYD

[Photo origin unknown]

52 ... Petersen

when he was nine. Typical for the era, he attended country schools and never

enrolled in any college or university. “Whether or not this latter fact influenced

him to chide the professors or not can hardly be established” (Hesler 1975)

As he neared puberty, Lloyd became acquainted with the medicinal plants

surrounding him and discovered that his harvest, when correctly prepared,

could be sold to a pharmacy. Lloyd and his brothers established a successful

pharmacy of their own and expanded the business into nearby Cincinnati,

Ohio, as the Lloyd Brothers Pharmaceutical Company. The company grew

and prospered (thanks largely to the efforts of his brothers) while Curtis Gates

devoted time to avocations, including global travel.

Fic. 25. JoHN DEARNESS on his 80" birthday

[Mycologia 47(1). 1947.]

Mycology in 19th century North America ... 53

In “far-off” Canada in 1872, John Dearness (1852-1954; Fics 15, 25) passed

a three-month course at Toronto (Ontario) Normal School and became

Principal at Lucan village school. The following year saw him Principal of

Strathroy Public School and in 1874, he was appointed Public School Inspector

of East Middlesex (a position he held for 25 years). Much later (1899-1914)

he was Professor of Biology, Medical School of Western University. He was to

become an “early” Canadian mycologist.

Fic. 26. JOSEPH CHARLES ARTHUR

[Phytopathology 32(10). 1942]

Although born in rural New York State on 11 January 1856, Charles Joseph

Arthur (1856-1942; Fic. 26) and his family moved to northeast lowa when

he was six. As could be expected, Joseph (as he was known to distinguish

54 ... Petersen

him from his father, Charles) passed through public schools through high

school in Floyd County. Embarrassed by mix-ups in his name in school, he

informally changed it to Joseph Charles in adulthood. Upon graduation,

Arthur was sent to the newly created Iowa State University (1869) in Ames

where he already planned a career as a botanist. He was disappointed that

there were no botany courses in the new school, but coincidentally, Charles

E. Bessey arrived in 1870, and a match was made. Graduation soon followed

with a B.S. and after a short pause, an M.S. in 1877.

Arthur's obduracy with what he considered senseless university

requirements became troublesome, specifically, “In 1872, the American

Association for the Advancement of Science [AAAS] met at Dubuque, Iowa.

The great botanist, Asa Gray, was to be a speaker. Of course, a budding

botanical student desired to be there. In fact, he just had to go. But the

college authorities did not think it wise and would not give their approval

to his petition to be absent long enough to attend the meeting. No amount

of persuasion changed their verdict. What did our independent, ambitious

student do? He went. And he graduated a little later, but many amends had to

be made.” (Kern 1942)

Enter a powerful player in politics but also a small but pivotal role in mycology.

George William Clinton (1807-1885; Fic. 27) was born of an illustrious

family: his father, Dewitt, was a famous inventor and recent governor of New

York State. George had a bent for natural history and as a student of 19, he was

permitted to join an academic journey through the full length of the brand-new

Erie Canal, opened in 1825. The teacher for this endeavor was Amos Eaton,

author of several botany texts. The experience reinforced Clinton’s natural

history leanings. Time passed: Admitted to the bar in 1831, Clinton forged a

career as a prosecutor and judge. Always involved with natural history, he was

elected the first president of the Buffalo (NY) Society of Natural History, and

later a member of the New York State Board of Regents, one duty of which

was to oversee the State Museum. All the while, he maintained an enormous

correspondence with naturalists both domestic and foreign, one of whom was

Charles Horton Peck.

In about 1878, Andrew and Laura Morgan moved to “the little farm in Preston

[Ohio] for twenty-three years” (Kellerman 1907). His collecting, mushroom

studies, and interaction with Lloyd continued. “In the case of the more fleshy

fungi it is Mr. Morgan's practice to preserve only a painting. His collection

Mycology in 19th century North America ... 55

Fic. 27. GEORGE WILLIAM CLINTON

[Neilans. 1963]

contains many hundreds of these all made by the hand of Mrs. Morgan who

has aided in bringing together a most valuable collection” (Underwood 1899).

During 1867, Peck contributed his botanical skills as a volunteer part-time at

the State Museum. He was distraught when Cass’s Academy announced that it

would close at the end of the term due to financial difficulties. How would he

support his family? He appealed to Judge Clinton to promote a government

allocation in the state budget for a botanist. Clinton did so, and in 1867, a

56 ... Petersen

position was created for a worker to help “fill the herbarium.” Peck, of course,

was pleased to be able to pursue botany full-time, but the position was an

annual item—easily erased by omission or reallocation of funds. Not until

1883 was the title of “State Botanist” made permanent (Haines 1978; Petersen

1980b; Lloyd 1912).

Not immediately did botanist Peck understand that of all the plants of New

York, the least represented in the herbarium were the fungi. It took a couple

years and urging by a learned amateur mycologist and mentor, Elliot Howe,

before he began to investigate the group, and it was in that period that Frost

also lent his advice.

Although born in Vermont, Elliot Calvin Howe (1828-1899; Fic. 28) came

to New York State early in life. Educated in Troy and Lansingburgh, he had

strong interests in natural history and music (not unlike William Farlow a

generation later).

Early in life he gave indications of a love for natural science and turned

his attention to the study of geology, zoology, and botany. Music (piano) also

received a significant share of his attention. Pharmacy was also attractive to

him, which soon led him to the broader field of physiology and medicine.

While engaged in medical studies in New York City, Howe did literary and

reportorial work for the NEw YorRK TRIBUNE, then under the management

of its celebrated editor, Horace Greeley. After receiving the M.D. he returned

to Troy and commenced the practice of medicine. Music remained a strong

avocation and Howe wrote numerous pieces of popular music, of which one,

“The Wanderer’s Dream,” was played by the musicians of both armies during

the Civil War, a unique contribution to the war effort.

After a few years, Howe was induced to leave Troy to enter the large and

flourishing Charlotteville (NY) seminary as teacher of music, physiology, and

botany. His job was abruptly terminated when the seminary building burned

to the ground, but he quickly moved to Fort Edward Institute where he taught

music, botany, and German. There he met Miss Emily Z. Sloan, also a teacher

at the institute, who afterward became his wife.

While at Fort Edward Institute, Howe also began the study of fungi and

entered into correspondence with M.A. Curtis of North Carolina, who at that

time was the chief devotee and exponent of American mycology. Also during

this period, “he directed the attention of [C.H. Peck] to the interesting features

of mycology and induced him to enter this field of botanical investigation,

which at that time was almost a terra incognita in this country” (Peck 1899).

Mycology in 19th century North America ... 57

Fic. 28. ELLIOT HOWE

[Neilans. 1963]

After seven years Howe left Fort Edward to renew a medical practice in New

Baltimore (NY), but the field there was limited, and he soon moved to Yonkers

(NY), just north of New York City. His biographer (Peck) does not explain

the circumstance, but seven years before death, Howe became a quadriplegic,

and thereafter was confined to his home, a helpless invalid. On March 2, 1899,

“he fell asleep and a varied and useful life was closed” (Peck 1899).

In 1869, John Torrey informed Ravenel of two possible academic positions

(University of California and Washington University in Kentucky). Ravenel

demurred from both possibilities (and no offers were made), using deafness

58 ... Petersen

as an excuse. By the end of 1869, moreover, he was more sanguine about his

life and circumstances—certainly, life had improved substantially since the

end of the war. But former botanical relationships based on free exchange

now had to turn financial. This was difficult in some instances.

By 1872, he resumed correspondence with J.B. Ellis, who apparently

connected Ravenel with E von Thtimen (initially in Germany; then from

1876, in Austria), who desired specimens from America for his (Thtiimen)

exsiccati, MYCOTHECA UNIVERSALIS (Stevenson 1971: 448-459). Ravenel

expected financial return for the specimens submitted through Ellis ($6.00

per hundred). Ravenel specimens appeared in centuries VI (1876) through

XXII (last, 1884). Decades later, Stevenson (1971) identified several type

specimens from among Ravenel collections.

On April 10, 1872, Moses Ashley Curtis died suddenly, bringing finality to

the long correspondence between Curtis and Ravenel (and numerous others).

Curtis's post-war financial existence had been somewhat precarious, especially

considering the expenses of specimen shipments to Berkeley. Although the

congregations around Hillsboro, North Carolina, that were serviced by Curtis

did their best, there certainly was little money left over to pay their minister.

In contrast, his friend and coworker, Ravenel, had significantly changed his

lifestyle from plantation owner with numerous slaves to entrepreneur selling

his botanical expertise and experience.

Reconstruction

Abraham Lincoln issued the Emancipation Proclamation during the Civil

War, and the War had put the proclamation into practice. The result was

anything but smooth. A holistic set of laws and unwritten etiquette conspired

to mire Blacks in the lowest caste position. Although usually remembered

as laws that excluded Blacks from public transport and facilities, juries,

jobs, and neighborhoods, passage of the 13th, 14th, and 15th Amendments

to the Constitution actually afforded Blacks the same legal protections as

whites and women. However, after 1877, and the election of Republican

President Rutherford B. Hayes, southern and border states began restricting

the liberties of Blacks. Unfortunately, the Supreme Court helped undermine

the constitutional protections of Blacks with the infamous Plessy v. Ferguson

(1896) case, which legitimized “Jim Crow” laws and the “Jim Crow” way of life.

In 1877, Ravenel (in his journal; Arney 1947) planned a trip to Georgia and

Florida to collect all cryptogams for subscription, the proceeds of which

Mycology in 19th century North America... 59

would add to the household income. M.C. Cooke in London advised Ravenel

(in detail) on methods of preservation. “Cooke will attend to the fungi,

Tuckerman will take lichens.” Cooke's involvement was in answer to Ravenel’s

inquiry about the possibility of selling fungus collections from the southern

United States, suggesting that Cooke might have been the instigator and

executor of the exsiccati plan. Ravenel’s trip took place, and Cooke announced

the exsiccati in December 1877. “The series will be edited by M.C. Cooke,

and each century will be issued when it is ready. As only a small number of

sets will be made up, early application to the Editor of this Journal [Cooke,

GREVILLEA|] is desirable” (Stevenson 1971: 307). Indeed, century 1 of FUNGI

AMERICANI ExSICCATI was issued from London in 1878. The series would

extend to 1883 (century 8), and the final payment to Ravenel came promptly

that year—$42.

Although the decade from 1865 to 1875 saw the emergence of William Gilson

Farlow (1844-1919; Fic. 29) as a cryptogamist, only at the end of that time did

Farlow begin to emphasize the fungi. His overall contribution to mycology in

America was (and is) so significant that a somewhat lengthy sketch of his

timeline seems warranted.

A venerable and well-endowed institution, Harvard University was not

a place for second-echelon students and/or impecunious families unable

to pay the tuition. But Harvard also had an appreciation of prominent

families of means and when above-average intellect was matched with

financial resources, at least admission, if not graduation, was probable.

So when John S. Farlow, the successful businessman (Linder 1945), avid

prize-winning horticulturalist, prominent politician, and President of the

Boston Handel and Haydn Society recommended his son, William, to the

University, Harvard welcomed the young man, even though he had received

his previous education from public schools. William, like his father, had two

strong interests: botany and music. The first was exhibited in William's courses

with Asa Gray, who singled him out and co-opted his help in the classrooms.

The second prompted another teacher to suggest a professional career in music.

As an undergraduate, William's years at Harvard (1862-1866) continued to

impress Gray, especially as Farlow’s focus was on the “lower” cryptogams—

algae, mosses, fungi. [Setchell (1927) sketches the state of botany in the US.

during this time period.] But when William began planning a career in

botany, Gray forcefully advised a more dependable, lucrative profession to

which botany could be appended as an avocation. Examples were Torrey and

60 ... Petersen

Catntitl _ Moro,

Fic. 29. WILLIAM GILSON FARLOW

[Mycol. Notes (Lloyd) 40. 1916]

Gray, both with medical degrees. Following Gray’s advice, Farlow earned a

medical degree from Harvard in 1870. Instead of entering practice, however,

Farlow returned to Gray’s laboratory as an official assistant.

Just then, Farlow’s interest was focused on marine algae. The summer of

1871 was spent at Woods Hole (Massachusetts), the marine biology station,

in the company of outstanding algologists from all over the nation. At the

Mycology in 19th century North America... 61

same time, Charles Wright brought a cache of algal specimens from Cuba,

which Gray passed on to Farlow and which formed Farlow’s first publication

(Farlow 1871).

Shortly thereafter, Gray again persuaded Farlow, this time to consider a

European exposure to the leading botanists and laboratories. There ensued a

two-year grand tour through (at least) “England, the Scandinavian countries,

Russia, Germany, France, Italy, and Switzerland” (Harris 1945, Weston 1945).

There was no anxiety about finances, and Farlow bought numerous rare

books, returning also with an armful of specimens, mostly algae. During his

absence, in 1872 Gray purchased Curtis's herbarium on behalf of Farlow. Upon

return, Farlow was appointed Professor of Cryptogamic Botany, with duties

split between Cambridge and the Bussey Institution (Harvard's subsidiary of

Agriculture and Horticulture) in nearby Jamaica Plains.

Soon, Farlow was relieved of his Bussey duties and the Professor took

charge of teaching Cryptogamic Botany. He was able to introduce teaching

methods observed in Europe, including those of H.A. de Bary: microscopes

used in class and plant specimens inspected in lab—new to botanical teaching

but soon tried in other schools.

“In [physical] stature, Farlow was decidedly below the average, a matter

concerning which he was somewhat sensitive, especially when associated with

one which was taller. He seldom, however, made reference to it except through

some witticism” (Setchell 1927). Otherwise, his energy and volubility made

his lectures enjoyable, and he spent long hours sorting specimens together

with graduate students and hangers-on. Farlow’s focus shifted to fungi and

he began especially to consider parasitic fungi, especially plant pathogens.

He professed that there were three requirements for teaching cryptogamic

botany: 1) an herbarium of authentic reference specimens, 2) an extensive

and complete reference library, and 3) an index to keep track of the first two.

With student help he began projects in all three.

Elsewhere, the year 1874 saw a catalog of New England “boleti” by the learned

shoemaker, Charles Frost (Frost 1874). Although one of only a handful of Frost

publications, the paper must have had the imprimatur of Peck, considering

their friendship and the Buffalo journal in which it appeared.

Spurned from a hitch in the army in the Civil War, Albert Prentiss pursued a

career through educational jobs in public schools until he was appointed to

the faculty of the University of Nebraska for some years. Among his students

62 ... Petersen

were C.E. Bessey, W.P. Wilson, B.D. Halsted, and S.M. Tracy, all of whom went

on to produce meaningful mycological contributions. In 1868, not only did

Cornell open to students, but Prentiss made his way there as Head of the new

Department of Botany and director of campus grounds. The first course was

offered to an enrollment of four, but by the next term, attendance swelled to

144, a token of his teaching abilities.

Far from a country bumpkin, Prentiss was refined, sophisticated—and

widely traveled. In 1870, he banded some students together to join “The

Cornell Exploring Expedition,’ investigating the natural history of the

Amazon River in Brazil for six months. Two years later, he traveled to Europe

to study in the herbaria at Kew and Paris. During these years his health began

to deteriorate, and he was obliged to reduce his duties supervising campus

grounds. But he remained engaged in the classroom and with graduate

students who were numerous, bright, and destined to careers in botany. After

his retirement in 1885, he was succeeded by George Atkinson. Prentiss died in

Ithaca on August 14, 1896 (Atkinson 1896; from an entry apparently written

just before Prentiss’s death).

Mycologically, all was not dark in Europe while the Americans increased

their contributions. After all, Fries (the “father of mycology”) was still alive

and almost fully productive right up to his death in 1878. But even during

his lifetime other European workers proposed new schemes of fungal

classification. Léveillé (1846), well-known for his work in the South Seas,

produced a new organization. De Bary (1866), with whom Farlow studied,

added a new classification. In Great Britain, Berkeley’s OUTLINES OF BRITISH

FUNGOLOGY (1860) purported to be aimed at the British Isles but in many

ways actually covered the whole mycological world. Berkeley was succeeded

by M.C. Cooke, whose INTRODUCTION TO THE STUDY OF FUNGI (Cooke

1895) was even more comprehensive and popular. Cooke’s multivolume

ILLUSTRATIONS OF BRITISH FUNGI (Cooke 1881-1891) brought excellent

images to those who could afford them.

Although his work was hardly known in English-speaking North America,

Carlos Luis Spegazzini (1858-1927; Fic. 30) was busy collecting fungi and

describing what he found in Argentina. Born in Acosta, a small town near

Turin, Italy, Spegazzini took special interest in fungi and was friends with

Beccari and Pier Andrea Saccardo. He would follow Saccardo’s system of

classification for his lifetime.

Mycology in 19th century North America ... 63

Fic. 30. CARLOS LUIS SPEGAZZINI

[Origin unknown]

In 1879 (age 21), armed with a diploma from Collegio Real de Viticultura

y Enologica, Spegazzini embarked on a trip to South America where

settlements of Italians already existed. After a stop in Brazil, he disembarked

in Buenos Aires on Rio de La Plata, Argentina. The locality was to be his

home for life.

Repeated expeditions to remote parts of Argentina resulted in lengthy

descriptive publications of the fungi found wherever. In several ways,

Spegazzinis role in Argentina mirrored Peck’s role in the United States—

each a pioneer in his field, each describing hundreds of new species, and each

remaining in place and productive for many years. Had Spegazzini written

in English, perhaps his work would have been appreciated in the Northern

Hemisphere, but he was greatly celebrated in Argentina and his herbarium

is preserved at the Spegazzini Institute in La Plata (Anonymous undated,

Murrill 1924, Anonymous 2010, Macjus 2013).

64 ... Petersen

In a day when support groups were not formalized, family was of supreme

importance when death overcame a spouse. After the death of her invalid

husband, Flora Patterson moved to Ames, Iowa, where her brother taught

at Iowa State University and she earned a second M.A. in 1895. As her

boys grew, her brother was offered a position at Harvard Law School. The

circumstances of her migration are lost, but in 1896, she was teaching in a

Boston, Massachusetts, private school. Flora took the opportunity to study in

the Gray Herbarium at Harvard.

Stationed in Boston and her boys coming of age, Patterson was determined

to continue her studies at nearby Yale University in Connecticut. “Thinking all

conditions for pursuing work at Yale had been arranged, she went there only

to find the doors closed against her, women not being eligible at that time.

In spite of the keen disappointment and inconvenience she persisted in her

desire to continue botanical investigations, and, [returning] to Cambridge,

registered for work at Radcliffe College. There she remained for three years,

1892-1895, taking courses in botany and working as an assistant in the Gray

Herbarium” (Charles 1928).

Ellis & Everhart

During his hitch in the navy, J.B. Ellis had, as usual, complained about

the climate in Potsdam, New York. A chum suggested the unlikely town of

Newfield, New Jersey, as possessing a pleasant climate. Ellis investigated the

place and wound up settling there for the rest of his life. During 1877-1878,

Ellis decided to drop all other pursuits to issue an exsiccati entitled FUNGI

Nova-CAESAREENSES, comprising ten sets of New Jersey fungi. He sold

two sets, one to Farlow (Harvard), the other to Mr. Isaac C. Martindale. At

the urging by Martindale these sets were recalled and renamed as NoRTH

AMERICAN FunGI (Anderson 1890, Stevenson 1971).

With these words as entré, some additional thoughts on exsiccati seem

appropriate. The idea was early developed that specimens (of anything) could

be exchanged among like-minded enthusiasts. But when a single specimen

proved interesting to more than one worker, the idea of sets of labeled

specimen multiplicates, which could be sent (gratis) or sold (a derivative idea)

became attractive, and these compilations took on the name “exsiccati.”

In all exsiccati with named specimens, the specimens are only as valuable

as the identifications appended to them. Thus, specimens of powdery mildews

identified (or named as new) by Ellis (the American expert at the time) take

Mycology in 19th century North America... 65

on more “authenticity” than those of most agarics (some, originating from

North America, were identified by M.C. Cooke from England without

any geographical caveats and assuming that American taxa and European

taxa could be labelled with the same names—all too common even now).

Therefore, exsiccati specimens are candidates for basic taxonomic research—

not to be treated as “deo volenti.”

Ellis had been exposed to the concept (and, as it turned out, to mycology

as a whole) when he saw a broadbill announcing Ravenel's exsiccati. Years

later he exploded the concept to theretofore unheard-of size. The issuance

of exsiccati multiplied apace and by the last years of the 19" century there

were scores of exsiccati available from and representative of several European

countries and American states. The specimens were heavily of fungi that could

be pressed and dried, including gasteromycetes and manageable polypores

(those which could be pressed or sliced so as not to take up excessive space).

Hymenomycetes were more difficult to preserve and lost most of their shape,

stature, and color. Ancillary notes on these characters must accompany such

specimens.

In Ellis’s case, between NORTH AMERICAN FUNGI [36 centuries in two

series: as sole author centuries 1-15 (starting in 1885) and with B.M. Everhart

centuries 16-36 (1886-1896); assuming c. 50 subscriptions totaling 180,000

specimen packets] and FUNGI CoLUMBIANI [centuries 1-51 (starting in

1891; and with century 14 by C.L. Shear), and centuries 15-51 (ending in

1917) by Elam Bartholomew in Kansas; using the same calculation totaling

255,000 specimen packets], a total of 435,000 specimens were added to North

American and European herbaria.

In acameo performance, Benjamin Matlock Everhart (1818-1904) was not

a mycologist—at least not by education or profession—although that opinion

was not universal. “He is one of the best mycologists that America has ever

produced, having done much good work with Mr. Ellis in the description

of new and rare plants” (Harshberger 1899). Born May 24, 1818, in Chester

County, Pennsylvania, Everhart and his family were neighbors of (and perhaps

fellow Quakers with) of Ezra Michener and Dr. William Darlington. After the

usual rudimentary education at West Chester Academy, Benjamin worked

in the family’s general store (eventually succeeding his father) and built a

“comfortable fortune” (Harshberger 1899, Kellerman 1904, Hesler 1975).

A glimpse of Everhart was offered by Harshberger (1899), who visited him in

1895. “Having kept a general store in West Chester, in which he took much

66 ... Petersen

pride, Mr. Everhart amassed a considerable fortune, and is looked upon as

one of the wealthiest men in West Chester. By his neighbors he is considered

rather unapproachable and peculiar, as a man ready to push a close bargain.

In person Mr. Everhart is a man of striking appearance; his nose is aquiline,

his forehead low, his beard is scant and fringing, his eyes are bright, and his

smile pleasant. At the age of seventy-seven he is still a man of considerable

activity, although slightly deaf” (Harshberger 1899: 227). Everhart had a

strong avocation—natural history—and was a collector of plants, birds, and

insects.

The details of how J.B. Ellis first interacted with B.M. Everhart have been

lost, and their initial correspondence dealt only with the cost of specimens.

Most likely Everhart heard of or saw a century of Ellis’s exsiccati and therefore

initiated contact. Nonetheless, as their association developed, Ellis may have

told Everhart of the financial obligations inherent in issuing large numbers of

exsiccati specimens. Everhart, the businessman, perhaps saw an opportunity

to underwrite Ellis’s efforts for the good of natural history and perhaps receive

the products, and it has been said that thereafter Ellis provided the specimens

while Everhart provided the money. He also collaborated in the publication

of Ellis's major monograph on Pyrenomycetes in 1892, and his name is on the

title page as co-author.

Because of the heavy emphasis on those fungi which Ellis knew well, the

exsiccati include many type specimens as well as contributions from numerous

collectors across the continent. Attribution for identifications included most

of the leading mycologists of the day, including European, and specimens

represent all classes of fungi (of the time).

In 1899, Arvilla, Ellis’s wife died, putting responsibility of total exsiccati

work on J.B’s shoulders, and the task soon proved overwhelming. As Ellis’s

biographer wrote: “Many years ago she began aiding her husband, Mr. J.B.

Ellis, in the arduous labor of preparing and mounting the specimens for the

NorTH AMERICAN Funai and later for the FUNGI COLUMBIANI, and with her

own hands bound the books in which these were delivered to subscribers.

Had it not been for her help, the first of these great distributions (numbering

3,600 specimens) would have been suspended early in its history, and the

second (numbering 1,400 specimens) would never have come into existence.

To her deft fingers, which wrought so patiently, botanical science is indebted

for the more than two hundred thousand specimens of the fungi which Mr.

Ellis distributed to the botanists of the world” (from Science Aug 11, 1899;

Kellerman 1906).

Mycology in 19th century North America ... 67

It must have been sometime after 1878, around the time Andrew P. Morgan

and wife Laura moved to a farm in Preston, Ohio, that C.G. Lloyd began

interacting with Morgan, to whom Lloyd attributed his introduction to the

study of fungi.

The details of Lloyd’s “retirement” from the family business are hazy, but

he withdrew from active service only to continue receiving the income of

a partner. This enabled him to travel at will, including trans-Atlantic tours,

and to begin production and dissemination of his own publications on fungi,

which began in 1898 and filled seven volumes to 1925 (Stevenson 1933).

It can be reported that Lloyd was mycologically self-taught, but more than

that, Morgan, his mentor, was also self-taught. This inevitably left lacunae in

Lloyd's “education,” but how this affected his personality cannot be known.

Elam Bartholomew's interaction with W.A. Kellerman kindled a fresh fire

in Bartholomew. Soon he was collecting and identifying plant pathogens

and, as was typical, began a herbarium. His circle of correspondents

changed somewhat, expanding to include the mycological luminaries

of the day: Peck, Ellis, Holway, Arthur, Bethel, Dearness, and others.

Introduced to the concept of specimen exchange through exsiccati, in 1901

Bartholomew endeavored to publish and disseminate the exsiccati series

FuNGI COLUMBIANA originated by Ellis and Everhart (36 centuries by 1917).

In 1911, he began his own exsiccati, NORTH AMERICAN UREDINALES, which

over the years extended to 35 centuries. Work on his Kansas farm went on

unabated, but as his reputation spread, Bartholomew received invitations to

join faculties of colleges and universities. These were consistently refused.

He gave as his reason for refusal that he preferred to work in private on

the farm which he had homesteaded in the waning days of the buffalo and

antelope (Bartholomew 1935). Eventually in 1929, however, he and Rachel

moved to Fort Hays where he became curator of the herbarium. There they

remained until his death, 18 Nov. 1934 (Kansas Historical Society 2018).

Not a prolific publisher, Bartholomew contributed some summaries.

His specialty became the rust fungi, which were common and diverse in

Kansas. THE PLANT RUSTS OF KANSAS and especially his HANDBOOK OF THE

NorTH AMERICAN UREDINALES were comprehensive, with the HANDBOOK

announced and reviewed by the leading uredinologist of the day, J.C. Arthur

(Arthur 1934a). Bartholomew filled two important roles in the mycology of

the late nineteenth and early twentieth century: 1) his ardent collecting and

exsiccati exchange furnished specimens to a wide audience, and 2) his hitherto

68 ... Petersen

unexplored Kansas location gave a hint at the plant-fungus associations of

the Great Plains.

1896 found ES. Earle, now Biologist and Horticulturist of the Alabama

Experiment Station [to become the Alabama Polytechnic Institute in 1899

and in 1960 Auburn University], “and it was here that Professor Earle came

into close personal touch with the eminent mycologist George F. Atkinson”

(Chardon 1929). Almost surely untrue (or misleading), for Atkinson served

on the Alabama faculty during 1889-1892 and by 1896 had returned to

Cornell. But as Atkinson also collected in Alabama on subsequent field trips,

interaction might have been possible at those times. Lucian Underwood, also

at Alabama during 1895-1896 before migrating to Columbia University in

New York, probably also met Earle.

Except for his 1901-1904 sojourn at the New York Botanical Garden as

curator of the mycological collections (where his interest turned briefly to

agarics), Earle never again deviated from plant pathology. With a rich library

and time for research, Earle amassed what became a seminal work, albeit

largely overlooked for many years, THE GENERA OF THE NORTH AMERICAN

GILL FUNGI, which appeared in print after his departure as a Bulletin of the

Garden (Earle 1909). Previous more focused papers (Earle 1902a,b,c) also

reflect some interest in agarics.

“Being himself a member of the old school, he ventured to cover too

wide a field in taxonomy. Thus, while specializing on the Agaricales on one

hand he also tried to cover such widely diverse groups as the tropical species

of Meliola, the Hypocreales, the Xylariaceae, and all the groups of the Fungi

Imperfecti’ (Chardon 1929).

Born during the Civil War, but far away from the front, Robert Almer Harper

(1862-1940; Fic. 31) came into this world the son of a minister in Le Clair,

a tiny town on the shore of the Mississippi River, just inside the lowa border

with Illinois. His parents apparently had a high regard for education, and

Robert graduated from Ohio’ Oberlin College in 1886 at the age of 21.

For two years Robert Harper taught Latin and Greek at Gates College,

Nebraska, but moved on to study at Johns Hopkins University in Baltimore,

Maryland. His formal education sent him to the Strassburger laboratory in

Bonn, Germany where he was awarded a Ph.D. degree in 1896 (age 34).

Harper's credentials, including a German doctorate, earned him an

appointment as Professor and Head of the Department of Botany at

Mycology in 19th century North America ... 69

Fic. 31. ROBERT ALMAN HARPER

[Bull. Torrey Bot. Club 69(5). 1942]

Wisconsin University, where he served for a dozen years (1896-1911).

His enthusiasm for field trips was evident and he reveled in the richness of

the local flora.

In 1885, George Francis Atkinson (1854-1918; Fic. 32) at 31 years of age,

emerged from Cornell University with a B. Phil. (now considered a B.A.),

ultimately to be his highest earned degree. He had been born in mid-winter

1854, in the hamlet of Raisinville, Michigan, not far from Maybee and

southeast of Ann Arbor. The Civil War had been fought far away during his

early years, and his teenage activities are no longer known, but in his 24" year

(1878) he enrolled in Olivet College (Eaton Co, Michigan), a small liberal

arts abolitionist institution founded in 1845 to educate students regardless

70 ... Petersen

Fig. 32. GEORGE FRANCIS ATKINSON

[Cornell Plant Pathology Herbarium]

of gender, background, or financial need). There he took some courses but

was soon after admitted to Cornell as a junior (Humphrey 1961). Possibly

these undergraduate experiences prompted him later to write several books

intended as an introduction of botany to fledgling students.

Mycology in 19th century North America... 71

Armed with his formal degree, Atkinson embarked on a whirlwind

revolving-door tour of southern educational institutions. The first was

the University of North Carolina (three academic years, 1885-1888). Next

was the University of South Carolina (a single academic year, 1888-1889),

then to a newly created position as Professor at the Alabama Agriculture

and Mechanical Institution (now Auburn University) for three academic

years, 1889-1892). In all these positions he focused professionally on plant

pathology, but he was able to insinuate some publications on much wider

subjects, including entomology, zoology, ornithology, and botany. Fresh

from Alabama (where he had been awarded an honorary M.A.) Atkinson

accepted a position as Assistant Professor of Botany at his alma mater, Cornell

(1892-1918), replacing William R. Dudley who migrated to Stanford

University in California. Atkinson was promoted the following year. In 1896,

upon the death of Albert Prentiss, Atkinson succeeded as Department Head.

Although he might not have known, Cornell was to be his final career home.

Educationally, Flora Patterson's credentials were superb, but the opportunities

for using her extensive training equal to men with less education were slim to

none. Her own boys fledged from the nest, she was appointed in 1896 to the

staff of the (then) Division of Vegetable Pathology and Physiology, then under

the direction of B.T. Galloway, also interim editor of JouURNAL OF MycoLoey.

In this position she excelled and eventually was chosen as Mycologist in

Charge of Mycological and Pathological Collections in Beltsville, Maryland.

In this capacity she published several bulletins of the Division of Plant

Industry (Galloway 1928). She also gathered and disseminated multiplicates

of over 1400 excess specimens, not as centuries in the usual exsiccati format,

but simply packeted, labeled, boxed, and shipped to institutions in all corners

of the United States. In one large assemblage she was aided by W.W. Diehl and

Edith Cash of the same department.

Connected with many scientific organizations, Mrs. Patterson was a

Fellow of AAAS and member of the Botanical Society of America, National

Geographic Society, Washington Botanical Society, Biological Society of

Washington, American Phytopathological Society, and American Association

of University Women (Charles 1929). Flora Wambaugh Patterson died in

Brooklyn, New York, near her sons on February 5, 1928.

Traveling to a job in New York City, J.C. Arthur stumbled into an internship at

the newly formed Johns Hopkins University in Baltimore, and upon meeting

72. ... Petersen

William G. Farlow, a guest lecturer, spent a semester at Harvard. Positions

followed at the University of Wisconsin (1879-1881) and University of

Minnesota (1882) before he was appointed to the New York Agriculture

Experiment Station at Geneva, New York in 1884. Arthur easily traveled

the short distance to Ithaca, site of Cornell University, to earn Cornell's first

Doctorate of Science in 1886. His dissertation was a treatise on pear diseases.

In 1883 Arthur and two other scientists founded the BOTANICAL GAZETTE,

destined for a long and prestigious run.

In 1887, Arthur moved to Purdue University, the state's land-grant

institution in West Lafayette, Indiana, to begin what evolved into his final

position, and he married to Emily Stiles Potter of Lafayette in 1901.

Arthur had kept in touch with Edward Willet Dorland Holway

(1853-1923) since their days at Iowa State, and in 1894, like so many

mycological colleagues, the two began issuing their exsiccati, UREDINAE

ExsIccaTI ET Icongs. Arthur furnished the specimens and identifications,

while Holway assumed the actual presentation of the specimens, presumably

underwriting the costs as well as collecting in Iowa, California, and Mexico

(Stevenson 1971). The number of specimens per fascicle was low compared

to other compendia (e.g., Ellis & Everhart), totaling only 60 specimens in

four fascicles, the last of which appeared in 1902.

Among Arthur's many students at Purdue was Fred J. Seaver, then beginning

a long mycological career at the New York Botanical Garden. “The writer's

[Seaver's] first intimate knowledge of Dr. Arthur's work was gained during

the spring of 1903 while a student in the State University of Iowa, having been

delegated by Professor T. H. Macbride to assist Dr. Arthur in his rust work for

that season, at Purdue University. The ten weeks spent in Indiana were both

profitable and interesting. Although not particularly interested in the rusts,

I was much impressed by the methodical manner in which the culture work

was conducted and the great care in handling all details connected with his

rust herbarium” (Seaver 1928).

One of his more dispassionate biographers relates that George Atkinson's

lectures were dull and boring (Thom 1956) but that frequent fieldtrips were

exciting and fun. Students worked into the night arranging and studying

the fungi collected during the day. Atkinson’s course syllabus was a set

of mimeographed detailed keys to the fungi covered in lecture (limited to

25 hard-bound copies with instructions that they were not to leave the lab;

Fic. 33), often annotated with personal taxonomic comments. Although

Mycology in 19th century North America ... 73

~ Ma ae

Bi. Ark nied oe, Coch nro

Mtr, make notes Le,

hes oii wk iene Lar

Wikeat tha pec pevmisoden fn a a

by, £ (Chinern—

Fig. 33. GEORGE F. ATKINSON

[Inscription from laboratory syllabus in Atkinson’s hand. Ined.]

carefully compiled, it was difficult to determine the taxonomic arrangement

of the fungi.

Adding to the attraction of field trips was Atkinson’s growing attention

to fleshy fungi, commencing about 1900. It must have coincided with his

discovery of photography as a means of representing organisms accurately

and relatively quickly. The equipment was delicate, cumbersome, complicated

to assemble and required lengthy (often a minute or more) exposure (FIG. 34).

The final images were produced on thin glass sheets, easily scarred or broken,

and faithfully capturing fingerprints when carelessly handled. He introduced

such photos as early as 1897 (Atkinson 1897). Encouragement must have

come from an unexpected source—C.G. Lloyd, who introduced photographs

in his writings as early as 1898 (Lloyd 1898), and Charles McIlvaine was to

include photographs in his magnum opus in 1900.

V7"

Fic. 34. George F. Atkinson with bellows camera and mushroom subject

[Cornell Plant Pathology Herbarium]

7A ... Petersen

In addition to considering himself mycologically educated enough to

summarise some fungal groups (notably polypores, phalloids, and puffballs),

C.G. Lloyd also had the hubris to criticize two aspects of the profession:

[1] the excess of species being proposed (Lloyd maintaining that there was

a limited number of species of fungi and the same organism was being

described repetitively) and [2] despite the nomenclatural code’s rules for

citing originating or transferring authorities, presentation of author names

constituted blatant self-advertising and must be eliminated. But instead of

merely adhering to his own biases, Lloyd used his pen to chide those who

did not conform to his dicta. Not subject to the review process or journal

regulations, he used his own self-funded publications for his criticisms.

Two devices were conjured for this purpose: [1] “Dr. McGinty” (obviously a

pseudonym), who took taxonomists to task but described fictitious species of

his own (bringing a problem to the accepted nomenclature regulations), and

[2] a small Buddhistic efhgy dubbed “Tso Kay,’ who signaled an attack on

“Kuntzeism” (referring to deceased taxonomist Otto Kuntze, who zealously

appended his own name after each proposed binomial) and “self-advertising.”

Transgressors were named, and the offending publication cited. Nevertheless,

Lloyd’s own taxonomic proposals were legion and filled 290 pages (Stevenson

& Cash 1936).

In “The Myths of Mycology,’ published in 1917, Lloyd wrote, “the mistakes,

blunders, and personal foibles of mycological writers have been my chief

source of pleasure. I have never failed to express myself plainly, and have

spared neither friend nor antagonist. I have always tried to be good-natured

in my comments, and as a general thing the parties affected are taking it more

as a joke on themselves and an idiosyncrasy of myself. . .. Nor am I deceiving

myself into the belief that I will accomplish what Iam apparently trying to bring

about, the abolition of personal advertisements in mycology” (Fitzpatrick

1927). Not all recipients, however, took them as light. Some, notably Lee Oras

Overholts and George F. Atkinson, were wounded. In Overholts’ case, Lloyd’s

comments caused an unfortunate end to the exploration of “split genera”

of polypores and return to Polyporus as a large and unwieldy assemblage.

Atkinson was merely offended, and his self-confidence shaken. He despised

Lloyd, but the reverse was not true (Hesler, pers. comm.).

Lloyd established “headquarters” in London, Paris, and Uppsala—also

sporadically in Berlin—and knew and was known to many of the active

workers of Europe. Their opinions of Lloyd were not recorded, but in far-off

New Zealand, frequent correspondent G.H. Cunningham (1892-1962) left

Mycology in 19th century North America ... 75

some acerbic comments as marginal notes in his copies of Lloyd publications

(S.R. Pennycook, 2019 pers. comm.).

His writings went on unabated until shortly before his death in Cincinnati

on 11 November 1926 (age 67) (Fitzpatrick 1927). He had amassed a large

herbarium—specimens were received in trade for his writings—and a large

library at his home in Cincinnati. Hesler (1975) related the following story:

“Shortly after coming to Tennessee [1919], I took occasion to ship a few

polypores and related things to [Lloyd]. He responded promptly, and asked

for more, which I continued to send for some time. One hot September

day, I had stopped overnight in Cincinnati, and went to see Mr. Lloyd at his

herbarium. I found the place, stepped to the front door and rang the bell. I

saw an older man sitting very close to, but with his back to the open door. He

was attired in only blue-jeans and an undershirt, and no shoes. I continued

to ring, but he seemed to pay no attention. Finally, the housekeeper answered

with a ‘ye-e-e-ess?’ I said a wished to see Mr. Lloyd. She replied: “Right here’

And pointed. Finally, I gave her my name, whereupon he wheeled around,

greeted me heartily and then took me (he bare-footed) all over the herbarium.

I asked what he intended to do with his enormous collection. He said: ‘Some

good institution will get it.” That institution turned out to be the Smithsonian

Institution and his collections reside at Beltsville, Maryland.

If mycology at the turn of the 20" century had been a court, Lloyd would

have been the court jester.

Letter from Father A.B. Langlois (1832-1900 (Fic. 35), in far-off Louisiana,

to Lamson-Scribner (1893): “In 1884, through the kindness of Mr. Lehnert,

of Washington, I began the study of mosses, liverworts and lichens, and in

the latter part of 1885, at the suggestion of Mr. Scribner, I began the study of

fungi. I soon acquired a deep interest in these plants, and have been greatly

aided in their study by Prof. Ellis, of New Jersey. The mycological flora of

Louisiana being so rich and at the same time so poorly known, I have for

the past three years given almost my entire attention to it. Every day I make

new discoveries, and I am yet far from having exhausted this intensely

interesting part of the Louisiana flora... Mr. Langlois has now an herbarium

containing some 5000 species of North American plants, including 1214

species of Phanerogams and vascular cryptogams of his State. So far as his

State is concerned, this work has been done single-handed. About a year ago

[~1892?], Langlois published a catalogue of Louisiana plants which embraced

the fungi he had found, now numbering 1200 species. Langlois’ collections

76 ... Petersen

. ec a se

Fic. 35. FATHER A.B. LANGLOIS

[Courtesy Dr. Meredith Blackwell]

are widely distributed in the herbaria of this country and in France, and his

specimens are highly valued by all who possess them.”

Edward Angus Burt (1859-1939; Fic. 36) is best-known for his series on

the Thelephoraceae, but his background is also elucidating. Born in Athens,

Pennsylvania, a small settlement on the Susquehanna River, his early years

are unrecorded. One position held (1880-1885, credentials unknown) was at

Albany [New York] State Normal College, where he knew Peck, and where

C.L. Shear was one of his students. After this practical experience, Burt was

awarded a Harvard B.A. in 1893 at age 34, followed by an M.A. in 1894. There

he was a product of William G. Farlow and Farlow’s assistant and successor,

Mycology in 19th century North America ... 77

Roland Thaxter. Burt’s next appointment was at Middlebury College in

Vermont (1895-1913). Finally, in 1913, he was employed as Mycologist and

Librarian at the Missouri Botanical Garden in St. Louis. He held a concurrent

position at the Henry Shaw School of Botany at Washington University, also

in St. Louis. Coincidentally, C.L. Shear was again a Burt student. When the

Farlow watercolor illustrations of mushrooms by Bridgham and Krieger were

being compiled into book form (see below), Burt wrote descriptions for each

species, including some nomenclatural and/or experiential annotations. It

was also during this period that he published the series on Thelephoraceae

as the family was understood at that time. While its content was as detailed

nomenclaturally as the times would permit, descriptions and illustrations

Fic. 36. EDWARD ANGUS BURT

[Mycol. Notes (Lloyd) no. 47. 1917]

78 ... Petersen

were not so, and the series met with mixed reviews (Lloyd 1917). He died in

Ballston Spa, New York, in 1939 at age 80.

In July 1887, Henry Ravenel suffered a serious of small strokes, but death

was caused by failure of vital organs—in short, by old age. His herbarium

was sold piecemeal—a substantial portion of the cryptogams to the British

Museum for $600. Long after his death, a large part of his phanerogams was

sold to George Washington Vanderbilt (1862-1914) for his planned Biltmore

estate and ultimately were ruined by a flood. Another part went to Converse

College, SC.

Mycology in the last quarter of the 19" century is a piecemeal story. Several

young men were in various stages of their careers, most as teachers in

high-school situations. Their careers can be traced, and their interactions

considered, but information on their personal lives and personalities depend

largely on obituaries, some of which are eulogies and others dry and skeletal.

Lucien Underwood's days as an Illinois school principal began an all-too-

common tour of teaching jobs—1878-1879 back at Cazenovia, 1879-1880

a professor of Natural Sciences at Hedding College, and 1880-1893 at

Illinois Wesleyan University (in 1881 marrying Marie A. Spurr, who would

accompany him on all subsequent career moves). In 1883 he became

Instructor at Syracuse (New York), a less peripatetic position lasting until

1890 during which Underwood introduced the “new” method of teaching

biology with using microscopes and living plants, introduced at Harvard

some years previously.

One of William Kellerman’s major contributions to mycology per se came

with the start of the JoURNAL OF Myco.oecy with J.B. Ellis and B.M. Everhart

as co-editors. With volume two, however, Kellerman was sole editor. Volumes

1-5 (1885-1889) were decidedly mycological in content but included reviews

of literature and mycological news. Financial difficulties grew, however, and

volumes 6 & 7 (1891-1892) were issued by the USDA with the 26-year-

old Beverly Thomas Galloway (1863-1935) as editor. Content was almost

exclusively phytopathological, in keeping with Galloway’s interest and

Galloway’s position as Chief of the brand-new USDA Division of Vegetable

Pathology and Physiology. After a ten-year publication hiatus ensued, in

1902 Kellerman (by then at Ohio State University) took back the JoURNAL OF

Mycology in 19th century North America... 79

Myco.oey and maintained it until his death in 1908. The journal, he testily

stated, separated mycology from phytopathology, emphasizing the former. In

doing so, it created a focal point for American mycologists who could read

each other’s research and keep track of positions and locations. The journal

fulfilled a necessary nucleus around which the discipline could gather, but

with Kellerman’s death in 1908, it ceased, leaving North American mycology

without such a publication for many years.

Although described here in only a few words, the country shivered through

an economic crisis in the mid-1890s. Blamed by some on failed investments

in far-off Argentina, a Euro-American wave of stock sales into currency

backed by gold & silver spurred failures of 500 banks with insufficient gold

reserves, over 15,000 private businesses, hundreds of over-extended farms,

and the bankruptcy of the country’s three largest railroads. Soup kitchens

emerged in big cities, presaging the Great Depression some 30 years distant.

Based on shrinking tax revenue throughout the country, administrations

from municipal to federal cut back on employees and salaries. Reduction

in expenditures (including salaries) rippled through college and university

budgets, while tuition revenue decreased. One small corner of this trauma

affected individual government employees whether at the USDA, the newly

founded New York Botanical Garden, or the Land-Grant universities

across the agrarian states. Although not everyone was put on “reduced

rations,’ anxiety over the future of capitalism and democracy was rampant

as unemployment reached 25% in Pennsylvania, 35% in New York, and

45% in Michigan (compared with <10% nationwide unemployment during

the 2008-2011 recession). Until some order was restored in 1897-1898,

mycological progress, like everything else, stood still and many of the

characters introduced here must have had life-altering circumstances

seemingly unrelated to their mycological activities.

After his wife’s death, J.B. Ellis redoubled his taxonomic research and in 1892

again teamed with Everhart, this time to publish THE NorTH AMERICAN

PYRENOMYCETES. As usual, it would appear that Everhart'’s philanthropy made

publication possible. It was to be Ellis’s last major mycological contribution.

Everhart passed away 22 September 1904 in West Chester, PA, with Ellis dying

a few months thereafter at home in Newfield on 30 March 1905.

Ellis, for all his productivity, was a simple, unassuming person from early

age onward. Unlike some other mycological associates like Farlow at Harvard

80 ... Petersen

or Atkinson at Cornell, Ellis had no students to mentor and no immediate

colleagues with whom to share opinions. Two quotes from others sum him up.

Written 15 years before Ellis’s death: “His fellow-botanists feel his influence

and recognize the value of his work but wonder why they never see his kindly

face at any of the botanical meetings of the country. It is simply because his

health at all times precarious, demands constant quietude coupled with strict

simplicity and regularity in his daily life. A thorough scholar and quite a

linguist, he is perfectly familiar with Latin, Greek, German, and French and

has also a good practical knowledge of Polish, Swedish, Italian, and Spanish”

(Anderson 1890).

From Curtis Gates Lloyd (1915), not easily given to kind words about

fellow mycologists: “He [Ellis] was exceedingly timid and shrinking, but

possessed of a charming personality, and by his lovable disposition endeared

himself to all who knew him.

“While I was aware of many mistakes that he made, I never, during his

lifetime, mentioned one of them in print. So earnest and honest was he in his

work, and possessed of such a lovable disposition, that it was impossible for

me to say a word that might have hurt his feelings in any way. I do not know

of any other person towards whom I have felt exactly as I did towards Mr.

Ellis, but consideration for his sensitive disposition and the high regard in

which I held him personally, prompted me thus to make of him an exception”

(Lloyd 1915).

Home-grown mycology

In the years from 1875 until shortly before his death, Farlow assiduously

gathered the tools of his discipline, cryptogamic botany: reference specimens,

reference library, indices to the field. The first were obtained by purchase of

numerous exsiccati, both domestic and foreign. These were kept intact but

indexed, so specimens could be retrieved easily. The second meant purchase of

domestic and foreign books and journals resulting in the largest mycological

library in North America. Third, several major indices were developed, most

of them unpublished (Setchell 1927). In one (Farlow & Seymour 1891),

Farlow wrote: “...believing that an approximately complete list of our parasitic

species and their hosts would aid materially in the advance toward a more

accurate study of our mycological flora and would tend to lessen the amount

of indiscriminate species making which has already become a serious evil,

the present index, the result of work extending over several years, has been

prepared for publication.” Although written for parasitic fungi, the words

Mycology in 19th century North America... 81

echo those written over the years by Lloyd for some of the same reasons.

In response to a swelling body of mycological literature, Lindau and Sydow

(1908) began their multi-volume bibliographic compilation, THESAURUS

LITTERATURAE MYCOLOGICAE ET LICHENOLOGICAE.

Married late (1900 at age 56), there were dinners at the Farlow home for

students and staff and the establishment of a summer home at Chocorua,

New Hampshire, on a lakeshore that he made famous for its wealth of

fleshy fungi. Setchell (1927) alluded to Farlow’s quiet financial support for

impecunious students, and Stevenson (1971) reported that Farlow may have

helped Ellis to launch his exsiccati, NORTH AMERICAN FunGI. The cover of

Century III indicates so.

At Farlow’s death (1919), his entire gathering of books and specimens made

its official way to Harvard where it formed the bulk of what became known

as the Farlow Herbarium and Library, the cryptogamic analog to the Gray

Herbarium and Library. Farlow’s roster of students was Malthusian: leading

lights included Blakeslee, Duggar, Pierce, Oakes Ames, Faull, Schenk, Fink,

Riddle, Burt, Jackson, and Thaxter, each becoming a leading contributor but

mostly after the time treated here.

In 1884 and again in 1885, Lucien Underwood had traveled to Harvard

to study with Asa Gray. This experience doubtlessly opened his eyes to

the greater transatlantic connections possible for a botanist. An AAAS

meeting in Ann Arbor, Michigan, presented an introduction to J.C. Arthur,

already becoming a recognized authority in the rust fungi. In the summer,

Underwood collected plants in Tennessee, Georgia, and Virginia for the

Smithsonian Institution, and in 1886, he was promoted to professor at

Syracuse. Throughout this period, his interests were drawn increasingly

to the fungi, and in 1889 Underwood and his student Orator Fuller Cook

issued their exsiccati, A CENTURY OF ILLUSTRATIVE FUNGI WITH GENERIC

SYNOPSES OF THE BASIDIOMYCETES AND MyxoMYCcETES (Stevenson 1971).

The long introduction (27 pp) treated fungal classification in general and

then provided family and generic keys for the covered groups. Fungi were

arranged taxonomically: 1-59 Basidiomycetes, 60-85 Ascomycetes, 86-88

Fungi Imperfecti, 89-93 Phycomycetes, and 94-100 Myxomycetes (Stevenson

197A)s

But Underwood's travels were not yet over. In 1890, he was Professor at

De Pauw University, Greencastle, Indiana, but the winter of 1891-1892 was

spent in Florida studying diseases of orange for the USDA, and he visited

82 ... Petersen

Cuba in 1892. His growing reputation prompted election to the committee on

nomenclature for the Rochester botanical meeting in 1891 and as a delegate

to the Genoa Botanical Congress in 1895. This trip had the “... advantage

of the opportunity to examine the famous herbaria and become acquainted

with the European botanical leaders in his line of study, such as Prantl,

Strassburger, Ascherson, Magnus, Ward, Chodat, Saccardo, De-Toni, Baillon,

and others. The interest thus aroused led him repeatedly to visit England and

the Continent, in all making eight trips for the purpose of comparison and

study at various botanical centers” (Curtis 1908).

As though in direct contradiction to Atkinson’s and Lloyd’s photographic

depictions, in 1889, Farlow hired John Bridgham to illustrate fresh fungi. His

watercolors were careful, attentive to detail under Farlow’s eye, and quite lovely.

His fee was $5.50 per diem (computed at about 75 cents per hour for a full

day’s work). He continued for 12 years (1901) before retiring. Bridgham was

replaced by the already recognized L.C.C. Krieger, who moved to Cambridge

for this singular purpose. Krieger continued in Farlow’s employ for a decade

(1912), his aquarelles every bit as fine as those of his predecessor. During his

tenure with Farlow, Krieger developed interest in presenting fungi, including

slime molds, to the public, publishing in NATIONAL GEOGRAPHIC MAGAZINE

(Krieger 1920) and a handbook for the public (Krieger 1935; financially

supported by Dr. Howard A. Kelly, his long-time patron). In the magazine

and handbook, Krieger's watercolor illustrations were supplemented by

photographic images. The work of both Bridgham and Krieger was gathered

posthumously, paired with descriptions by Farlow’s former student E.A. Burt,

and published as ICONES FARLOWIANAE (1929, with notes by Farlow written

in 1892; Pfister 1975).

The brief (April-December 1898) Spanish-American War ended with the

Treaty of Paris, which exchanged Cuba, Puerto Rico, the Philippines, and

Guam to the United States in return for $20 million. During the three and

a half years before Cuban independence in May 1902, Cuba was a USS.

protectorate.

ES. Earle was apparently attracted to tropical and/or subtropical climates

and their mycological possibilities, for in 1904, he accepted directorship of

Estacion Agronomica, Santiago de Vegas, Cuba (by then under the Cuban

government). The station soon closed, receiving inadequate governmental

support. For several years, Earle was agriculturist to the Cuban-American

Mycology in 19th century North America ... 83

Sugar Co. as well as President of the Cuba Fruit Exchange, also investing in

some private fruit-growing enterprises with varying success (Chardon 1929).

His worthy reputation was rewarded as Associate Editor of MycoLocia and

(1908) President of the Botanical Society of America. In 1918, with World

War I directing political affairs in the United States, the USDA appointed

Earle as specialist in sugar-cane culture and commissioned him to visit

Puerto Rico with the purpose of studying a very severe disease of sugar cane.

Earle's genetic immunity experiments restored sugar cane production, and

his reputation grew accordingly. Over the following years, he moved from job

to job, eventually associating with the Tropical Plant Research Foundation

until a few months before his death in Herradura, Cuba, on 31 January 1929

(Chardon 1929).

Were it not for Earle’s contribution on agarics, he would not be considered

for this sketch, for his entire career was centered on phytopathology, especially

diseases of sugar cane. He was surely a leader in carving a career in the south

and in his case, the Greater Antilles.

As the 20" century unfolded, an important document was issued from

Washington which, while surely not aimed at mycology, nevertheless

encouraged mycological discovery. Theodore Roosevelt signed a letter of

transmittal “To the Senate and House of Representatives” dated 19 December

1901 (Anonymous 1902b). According to the book-long report, there had

been an “uprising of the American people” over “an interest in practical

forestry, notable and commendable,’ having been preceded by establishment

of forest preserves in the western states. “A movement has grown to establish

a forest preserve in the southern Appalachians. The states which include

this region do not own appreciable land and do not have the revenues to

purchase it. It is therefore the federal government's responsibility to purchase

the forested lands needed as a preserve.” The movement for “preservation

by the Government of the hardwood forests on the slopes of the Southern

Appalachian Mountains” led to the National Forest system up and down the

Appalachian chain from New England to Florida, large patches of which were

to become National Parks.

Europe, of course, had not been still during the latter quarter of the 19"

century. M.C. Cooke (1893) wrote a textbook in which he outlined his

classification scheme. The giant score-keeper of mycology was Pier Andrea

Saccardo (1843-1920), who only meant to track the Pyrenomycetes in SYLLOGE

84 ... Petersen

FUNGORUM (1882) but wound up with 25 monumental volumes extending

past his death to 1931 (Petersen & Hawksworth 2016). In original research

the French became pivotal: Lucien Quélet (1886), and especially Narcisse

Patouillard (1887, 1900). Patouillard’s incessant use of the microscope was

establishing cracks in the Friesian edifice of classification, and the Saccardo

scheme, while practical, was totally artificial. As Patouillard’s work became

better known (French works took some time to become “discovered” by non-

Francophones), anatomical observations made fungal grouping different

than that based solely on basidiomatal macromorphology (Petersen 1971).

Meanwhile in Germany, a father and son team, Paul (father, 1851-1925) and

Hans (son, 1879-1946) Sydow, endeavored to start a new mycological journal.

Although, by now, some mycological journals already existed (i.e. BULLETIN

DE LA SOCIETE MYCOLOGIQUE DE FRANCE; JOURNAL OF MYCOLOGY), none

was German or Germanic. In 1903, the team launched ANNALES MYCOLOGICI,

which was to run through World War II.

Underwood's trans-Atlantic travels and time at Harvard convinced him

that a flora of North America, long dreamed by Gray and Torrey, should

be resuscitated. “[E]arly in 1893 he wrote a letter to Professor Britton

[New York Botanical Garden] proposing the formation of a body to organize

a general descriptive work on the flora of North America. This resulted in the

creation of a standing board of editors of the SYSTEMATIC BOTANY OF NORTH

AMERICA, which was subsequently transferred to Underwood and Britton

under the new title, NorRTH AMERICAN FLoRA, to be published by The New

York Botanical Garden” (Curtis 1908). Nathaniel Lord Britton of Columbia

University had been instrumental in organizing a movement which resulted

in the 1891 establishment of The New York Botanical Garden (BRITTON’S

BOTANICAL EMPIRE: THE NEw YORK BOTANICAL GARDEN AND AMERICAN

BoTANY. 1889-1920).

Then at De Pauw University, institutional financial difficulties forced

Underwood again to change jobs. The details of his migration are not known,

but 1895 found Underwood at the Alabama Polytechnic Institution (later to

become Auburn University). George Atkinson, who had been on the Alabama

faculty during 1888-1892 before returning to his alma mater (Cornell), was

followed by a zoologist on staff. Underwood came to Alabama for 1895 to

be followed by ES. Earle from 1896 to 1902. All three mycologists were well-

reputed: Atkinson and Underwood moved to more prestigious universities

while Earle moved to the Caribbean as a leading phytopathologist.

Mycology in 19th century North America... 85

In the spate of mushroom books emerging toward the end of the century

came two dealing with small regions, both largely overlooked in subsequent

years. The first on the Miami Valley of Ohio penned by A.P. Morgan came in

several parts (Morgan 1883-1888), while the second (Herbst 1899) covered

the Lehigh Valley of Pennsylvania. Some words have been devoted above

to Morgan (by this time on the farm with his wife), but William Herbst

(1833-1906) is also deserving, “being the foremost physician of [Berks] county

until his death. He served for thirteen years as physician at the almshouse,

[and] filled the chair of botany at Muhlenberg College.’ “Lutheran by religion,

Democrat by politics, member of the Masonic fraternity” (Roberts & al.

1914). Like so many others, Herbst acknowledged “Professor Peck, [who]

through his annual reports to the State Museum, and correspondence, was

my best teacher for the last twenty years” (Herbst 1899).

Herbst additionally noted, “C.G. Lloyd, of Cincinnati, had the kindness to

visit me for a number of years, giving me a fresh stimulus to this fascinating

study by his assistance in working out some interesting species” (Herbst

1899). In return, Lloyd gave Herbst’s book a complimentary review: “His

work we consider the most helpful single volume that has ever been issued on

American Agarics, because it considers and describes most of the common

plants (those met by everyone). All who are making a study of fungus should

procure it” (Lloyd 1900).

Employed for short periods as a railroad civil engineer, Charles McIlvaine

seemed never to require steady employment and perhaps came to exhibit

a sense of entitlement as a person who was rarely wrong and—to a large

extent—master of his universe. During the post-War years he married Sarah

G. Mcllvain (no relation). After an 1873-1874 tour of Europe, he began

penning contributions to American magazines: whimsical stories, verses, and

occasional scientific op-ed pieces, often written under the nom de plume,

Tobe Hodge.

By his words, he lived in West Virginia in 1880-1883. It was during a

horseback ride in this period that he was abruptly made cognizant of the

mushrooms growing all about him. Searching for written material about

such things, he was drawn to an article in POPULAR SCIENCE MONTHLY

(Palmer 1877). The article offered a wordy mushroom primer, but also

referred to several British books. Whether Mcllvaine searched out any of

them is unknown, but he set out to learn more, apparently not for elucidation

of the organisms themselves but with which to experiment with edibility.

86 ... Petersen

The result was such experimentation on over 800 taxa with notes on many

more (MclIlvaine 1900), earning him the sobriquet, “Iron Guts Mcllvaine.”

Mcllvaine gave generous thanks to C.H. Peck, who had been New York State

Botanist for years and was the unrivaled expert on eastern North American

mushrooms. But the same hubris which allowed Mcllvaine to “experiment”

with edibility also gave him license to identify and to name many of the forms

in the book. This could easily be multiplied by the ignorance of the reader,

which led to critical reviews by full-time mycologists.

One of the less enjoyable tasks of the New York State Botanist was to compile

and submit an annual report on his activities. This Peck did assiduously for all

the years of his tenure. Once out of his hands, his report was joined to others

— zoologist, geologist, etc. - in a year-end report by the State Museum. In

1890, Peck (1890) separately reported on a gift of mushroom illustrations by

Mary Elizabeth Banning (1842-1901), among the very best ever produced in

the United States (Bridgham and Krieger were yet to come some years hence).

Banning’s paintings remain in the New York State Museum.

Import-export mycology

In spite of his job as director of all research at the Cornell University

Agriculture Station at nearby Geneva, New York (Humphrey 1961),

Atkinson's passion for taxonomy of fleshy fungi prompted a trip to Sweden in

1903 in search of the Friesian identification of mushroom species. There he

was welcomed by Lars Romell (1854-1927), almost his exact contemporary,

who shepherded him in the field and furnished names for the collections.

Soon thereafter, frequent Swedish stops were included by C.G. Lloyd, by then

a mycological rival.

It must be remembered that for such international travel, months were

required for securing reservations in agreement with timetables, for stateroom

and venue accommodations, baggage disposition, and ground transportation,

all by time-consuming international post. Trans-Atlantic journeys required

at least two weeks of steamship travel in both directions as well as rail

transportation from embarkation port to and from the desired destination.

Mycology, like almost all other human activities, flourishes not only

through participants - researchers, students, learned amateurs — but by the

financial backing which permits such activities. Often, when some form of

government does not provide needed funds, the philanthropy of individuals

comes to the rescue. Such was the contribution of Howard Atwood Kelly

Mycology in 19th century North America ... 87

(1858-1943; Fic. 37), a well-reputed medical practitioner in Baltimore,

Maryland. Having amassed more than a comfortable fortune by mid-life, Kelly

could indulge his avocational interests in natural history. Time constraints

did not permit personal research, but vicarious satisfaction could come by

gathering the tools of the trade, in this case mycology. He sought and purchased

a magnificent botanical library heavy in mycology and lichenology, including

manuscripts and letters, original images and reproductions, herbaria, and

Fic. 37. DR. HowARD ATWOOD KELLY

[Mycologia 35(1). 1843]

88 ... Petersen

Fic. 38. LEwis CHARLES CHRISTOPHER KRIEGER

[Mycologia 33(3). 1941]

exsiccati. But such acquisition still took time, both in the execution and in the

arrangement and indexing of individual items. For the decade between World

War I and the Great Depression (1918-1928), Kelly employed Lewis Charles

Christopher Krieger (1873-1940; Fic. 38) for just this purpose. As mentioned

earlier, Krieger was a superb illustrator who had worked for William Farlow

at Harvard during 1902-1912, and during his time with Kelly, Krieger also

contributed over 300 original watercolors to the doctor’s cache. The index

alone of Kelly’s collection was book-length and published by Krieger (1924).

Mycology in 19th century North America... 89

In 1928, Kelly donated his enormous collection of printed material,

archival holdings, herbarium, and mycological images to the Herbarium of

the University of Michigan, where they remain available to the mycological

community. The event put Krieger out of work although he did remain

honorary curator of the Kelly collection.

Through several posts in and out of government, all after the period sketched

here, Krieger continued using his artistic skills, but lest he be considered solely

an artistic amanuensis, he also became an expert on mushrooms themselves

and the history and technique of his illustrative art (Krieger 1922, 1926, 1936,

1940), publishing several (some book-long) contributions. In 1929 (a dozen

years after Peck’s tenure), he became Mycologist of the State of New York

and published (tardily) a mushroom handbook (Krieger 1935). Krieger died

shortly after his wife. Of him, the following: “A figure unique in the annals of

American mycology passed from the scene of his earthly labors July 31, 1940.

L.C.C. Krieger was the creator of the finest series of watercolor paintings

of the fleshy fungi yet produced in America.” Further, “such perfection of

illustration has never been reached by anyone else in this country and in

Europe only by Boudier. There may never be another as competent as he”

(Lloyd 1920). “Not only do Krieger's plates approach perfection artistically,

but they are technically correct as well to the most minute detail, a rare but

much to be desired combination” (Stevenson 1941). Meanwhile, Kelley took

his place as mycological philanthropist alongside Farlow (to Harvard) and

Everhart (to J.B. Ellis).

Reduced duties in the Cornell Botany Department gave Atkinson the freedom

to write on botanical education, including several books: BloLOGy OF FERNS

(1894), ELEMENTARY BOTANY (1898), First STUDIES IN PLANT LIFE (1901),

LESSONS IN BOTANY (1901), A COLLEGE TEXTBOOK OF BOTANY (1905), and

BOTANY FOR HiGH SCHOOLS (1910)—a prodigious output, when added to

his STUDIES OF AMERICAN FUNGI, first issued as bulletins 138 and 168 of the

Cornell Agricultural Experiment Station, then as two book editions (1900,

1903) with numerous photos).

In 1911, Robert Almer Harper became Torrey Professor of Botany at Columbia

University, with a close association with The New York Botanical Garden

(NYBG). How different Manhattan must have seemed when compared with

his previous life in rural settings. Over his Columbia career “he spent one day

per week at [NYBG] for many years. During this period, he was interested in

90 ... Petersen

corn genetics: hybrids between sweet corn, flint, dent, flour, and pop-corn”

(Stout 1945), leading him to doubt the prevailing Mendelian genetics of that

day. He conducted his research at the NYBG experimental garden and a farm

in New Jersey bought by him & wife. Harper’s NYBG involvement included

membership on the Board of Managers for 31 years.

Even at urban Columbia, he led numerous field trips for Torrey Club

members and Columbia students. “He bought a Panhard automobile in

which he carried students to the Pine Barrens of New Jersey, to Cold Springs

Harbor, Long Island [and other seaside destinations]” (Anonymous 1942).

[Panhard was a French brand of luxury cars, never overly popular but

innovative; for some years, the steering wheel was positioned in the middle of

the dashboard—between the passengers but making it easier to drive in both

left- and right-hand countries) ].

Robert at Columbia had a brother, Edward T. Harper (1857-1921; Murrill

1921), atheologian. A graduate of Oberlin College (1881), Chicago Theological

Seminary (1886) and a D.D. from Leipzig (1891), Edward became a professor

of Semitics and Comparative Religions at his alma mater (1892-1911). But in

addition to his calling in theology, his strong avocation was botany, especially

fungi (probably influenced by Robert). He published no fewer than six papers

on the fungi of the Great Lakes area and his collections formed the basis for

the mycological herbarium at the Field Museum. Edward's photographic

illustrations were especially admired, and Krieger (1922) used them as an

example of the “heliogravure” process.

During his closing days, Andrew Morgan studied and published on a variety

of mushroom groups, often monographing his current interest. He died in

1907, on the farm in Preston, Ohio. Lloyd (1908), in a eulogy on Morgan,

reported, “[Morgan] pursued the study of mycology solely as a mental

recreation.” “... and it was he who first gave me an introduction to the study

of mycology.’ Considering Lloyd's blizzard of contributions, Morgan might

have been reluctant to claim his mentorship of the acerbic Cincinnati writer-

mycologist.

Some years after being passed over for a position at Ohio State University,

in 1901 Kellerman was appointed to the faculty there and soon had the

JOURNAL OF Myco.Loey running again. His publishing days did not stop

there, however, and in 1903 he started the OH1I0 MycOLoGICAL BULLETIN, a

pamphlet-sized organ aimed more at an amateur audience. It continued for

Mycology in 19th century North America... 91

at least 41 issues. These activities could still be carried on despite his gradual

hearing loss (Griggs & al. 1908).

Also in 1903, Kellerman, like several of his contemporaries, discovered

the tropics and its natural riches. After a preliminary foray to Guatemala, he

organized subsequent class trips with students, reveling in the experience and

its surroundings. The Ohio State administration approved of his plan and

in 1908, he took the class back to Guatemala. But during preparations just

prior to their return, Kellerman was stricken with malaria and after a few

days of incapacitation, died on 8 March 1908 in Zacapa, Guatemala, where

he was buried. With him went the JOURNAL OF MycoLoecy and the OHIO

MYCOLOGICAL BULLETIN. The former was to see a considerable hiatus.

Kellerman’s life was celebrated by many, for his cheerful energy and

published output made his mycological name a common, household

commodity.

Quite probably based on his relationship with N.L. Britton (Merrill 1938,

Mickulas 2007), in 1896, Lucien Underwood was appointed to the faculty

at Columbia University in New York City. There he was within a half-day’s

buggy ride to the newly founded New York Botanical Garden. Looking

back over his experiential shoulder, he published with ES. Earle a list of

Alabama fungi (Underwood & Earle 1897). In 1899, Underwood's “amateur's

guide to mycology’, MouLps MILDEws AND MusHRooms was published

(Underwood 1899). Hardly an amateur guide, the book provided genus-

by-genus summaries and a fairly extensive history of mycology, mostly

in Europe. Even a list of collectors who had sent specimens to Curtis was

inserted (p. 161). A careful inspection of the state-by-state list of literature

would reveal numerous references to places where Underwood himself had

collected or resided: Alabama, California, Connecticut (several summers in

the northwestern corner), Florida, Indiana, central New York. Together with

an honorary LL.D. from Syracuse, the book would crown a long career in the

fungi (Barnhart 1908, Howe 1908). Underwood died at home in Redding,

Connecticut, in November 1907.

If one could use the term “retirement” for a man of means and no regular

employment, Charles McIlvaine retired to Cambridge, Maryland, where

he died on August 4, 1909, a widower without heirs. He was active in the

Chautauqua movement, lecturing often on his favorite topic, but also on more

scientific planes. His most significant production was popularly nicknamed

92 ... Petersen

“The Mushroom Bible,’ (Bowser 1972) and had met with mixed reviews but

also commanded a “cult following” which existed well into the 20" century.

Peck published his annual reports from 1868 until his retirement in 1915

(Petersen 1980b). Established as the country’s mushroom expert, Peck

received specimens from all over the states. More or less in return, he

contributed specimens to at least 16 centuries of von Thiimen’s exsiccati,

MycoTHECA UNIVERSALIS. None of the specimens, however, represented

types (Stevenson 1971)

The annual reports were interspersed by occasional papers on extralimital

taxa in either the BurFALO NATURAL SCIENCE SOCIETY BULLETIN or

the BULLETIN OF THE TORREY BOTANICAL CLuB. One of his biographers

(Burnham 1919) estimated Peck’s output at 2700 new taxa, most of them

hymenomycetes. Two indices summarize his output: one by Peck (Bull. N-Y.

State Mus. 131: 51-190), the other by Gilbertson (1962).

In 1915, Peck suffered a series of small but debilitating strokes, necessitating

his retirement from active duty; in 1917, he died of a massive stroke in

Menands, near his family home.

Atkinson's growing reputation in research and education led to his election as

the first President of the BOTANICAL SOCIETY OF AMERICA in 1907. He might

have served equally in the mycological community had an organization of

mycologists existed at the time, but that was some years away.

Atkinson made the laborious trip to the International Botanical Congress

in Vienna in 1905, only to have mycological topics such as starting date for

fungal names and the type method postponed until the scheduled Brussels

Congress in 1910. Together with William Farlow and J.C. Arthur, Atkinson

attended the Brussels Congress, where botanical (and therefore mycological)

nomenclature was a significant topic. One can imagine lively conversations

among the trio, perhaps including the use of photographic images versus

watercolor paintings.

When Atkinson, Farlow, and Arthur “sailed” to Europe for the Brussels

Congress they would have been passengers on a steamship with steel

hull and rigged with a screw—a propeller under the posterior of the hull.

Although some cargo might have been loaded, above-deck passengers

would have had staterooms (often fitted with a porthole), while below-—deck

“accommodations” were dubbed “steerage” for their proximity to the engine

room. Masts and sails would have been jettisoned decades earlier. Fuel was

Mycology in 19th century North America ... 93

coal (probably bituminous), producing billows of ashen smoke for anyone

downwind from the funnels. Diesel engines in large ships were decades away.

In Brussels, mycological subjects would be discussed and committee

reports circulated prior to the Congress proper. Although not on the “official

commission” of the AAAS Botanical Club, Atkinson was a strong proponent of

the resultant AMERICAN CODE OF BOTANICAL NOMENCLATURE (Anonymous

1907) which enshrined these principles. Duly, Atkinson shepherded many

of the principles of the AMERICAN CODE into the INTERNATIONAL CODE. At

Brussels, he was not only a member of the nomenclature commission but

represented The University of Chicago, Cornell University, University of

Nebraska, AAAS, University of Vermont, and the American Phytopathological

Society. At the nomenclature sessions he introduced several motions already

in the American Code, of which all but one was accepted (the last being

postponed in the rush to adjourn) (Wildeman 1910).

The closing chapter

Throughout his career in phytopathology, C.L. Shear persisted in studying

the fungal pathogens themselves, more than just the diseases they caused.

His more outstanding mycological contributions were THE GENERA OF

Func1 with EE. Clements (1931) and, with B.O. Dodge (1927), “Life

histories and heterothallism of the red bread-mold fungi of the Monilia

sitophila group” (Jour. Agric. Res. 34: 1019-1042). This latter paper launched

a career for Dodge, put Neurospora on the genetic map, and got Dodge

an acknowledgement in Edward Lawrie Tatum’s Nobel Lecture. Shear’s

voluminous report on cranberry diseases served as his thesis for his 1907

Ph.D. from Washington University (St. Louis), where his faculty advisor was

E.A. Burt.

Also a student of mycological nomenclature, Shear was a delegate to the

international botanical congresses held in Vienna (1905), Brussels (1910),

Ithaca (1926), and Cambridge (1930), as well as to the 1950 Stockholm

Congress, which he did not attend because of ill health.

Retiring in 1935, Shear maintained his Washington, D.C., office while

spending winters in Florida. His MycoLoGicaLt Notes were published

during 1937-1940 (Cash 1954). After his wife died in 1950, Shear moved

to live with his daughter in Monroe, Louisiana, where he died in 1956

(Stevenson 1957).

Shear’s career balanced phytopathology and mycology, and his major

mycological contributions were made after the time period of this sketch.

94 ... Petersen

It would be easy to label J.C. Arthur a phytopathologist and thus peripheral to

this sketch, but his work on rust fungi, starting with their complex life cycles,

included “sowing” rust spores on possible alternate, greenhouse-grown host

plants. His own report (Arthur 1921) listed some 2300 collections involving

3750 pairings (Mains 1942, Hesler 1975) Thus, fungal genetics and physiology

were intricately involved.

Among his well over 200 published reports, a few are truly stellar. Early

on, Arthur (1905) used analysis of spore stages in the rust life cycle as the

basis for a new classification (Petersen 1974). Almost simultaneously, Arthur,

together with students and collaborators, started treating the rusts for NorTH

AMERICAN FLora (Arthur 1907). His students over the years included G.B.

Bisby, FD. Fromme, H.S. Jackson, E.D. Kern, E.B. Mains, and C.R. Orton. Well

after retirement in 1915, Arthur (1934b) published his MANUAL OF THE RUSTS

FOR NorTH AMERICA, aided by George Cummins.

“He [Arthur] was naturally much concerned with the rules of nomenclature,

not only as they applied to his group of rusts, but also as they applied to the

host plants. He was active in the Congresses at Vienna (1905), Brussels (1910),

and Cambridge (1930). In 1925 a trip was planned specifically to confer with

European mycologists who had special interests in the rusts. It was the privilege

of the writer to accompany Dr. Arthur on this trip. Visits were made with

Klebahn, Sydow, Dietel, Kniep, Lagerheim, Eriksson, Juel, Jorstad, Gaumann,

Ramsbottom, Butler, and Wakefield. A paper in SCIENCE (Vol. 43, pp. 558-560),

entitled “Conversations with European Mycologists,’ illustrates the value of

exchanging opinions with fellow botanists in Germany, Sweden, Norway,

Switzerland, and England. It was a strong link in a chain of cosmopolitan

activities forged by Dr. Arthur throughout his long life” (Kern 1942).

Finally, Hesler reported on an event that brought Arthur's iconoclastic

streak full circle. “After retirement, Arthur became upset when something

happened to displease him. In his miff, he engaged a moving-van on a Sunday,

and moved the entire rust herbarium to his home in Lafayette. The Purdue

community, naturally disturbed, persuaded the President of the University

to pay a personal call on Dr. Arthur at his home. After some diplomatic

exchanges, Arthur engaged the van to haul it all back to its rightful place the

next day” (Hesler 1975; confirmed by Cummins).

J.C. Arthur died on 30 April 1942 in Brook, Indiana. Although mycology

lost the worlds’ expert on rust fungi, the Arthur Herbarium lives on, and his

students (and their students) spread mycological research and pedagogy to

this day.

Mycology in 19th century North America... 95

After retirement from Columbia in 1930, R.A. Harper remained active as

Professor Emeritus until 1938, when the Harper family (Robert, wife Helen,

their son and grandson) moved to Bedford, Virginia, on a former plantation

of about 435 acres in the mountains between Lynchburg and Roanoke (Stout

1946). In 1942, the Torrey Botanical Club celebrated his 80" birthday. Harper

died on his Bedford farm 12 May 1946.

Hesler wrote: “About 1928, I visited Harper at Columbia. I found him an

imposing figure, over six feet high and weighing perhaps 220 pounds. He

had an unusually pleasing voice, was distinguished in appearance, and had

great personal magnetism” (Hesler 1975). Further: “He had a sense of humor:

Harper admired a (then) young physiologist, W.J. Robbins, later Director of

The New York Botanical Garden, and took Robbins on a mushroom hunt.

Once they gathered Polyporus sulphureus which Harper cooked. On tasting,

Harper remarked: “Well, I believe that it is fully equal to a good grade of filter

paper cooked the same way” (Hesler 1975).

It might be asked why Harper is included in a sketch of mycology. At

Wisconsin, Harper included fungi with all other botany and his students

were well-trained therefore. In his last decades at Columbia, his interests

were focused on the heredity and cytology of ascomycetes and myxomycetes,

passing his information to his students. He maintained a mycology genealogy

at Columbia (which also included Lucien Underwood) and at The New York

Botanical Garden, where ES. Earle spent a short but meaningful period.

By the time of Farlow’s death in 1919, Harvard had become the Kew of North

America, surely on a par with the other United States mycological centers.

The combination of Smithsonian Institution, Library of Congress, and

Department of Agriculture could have made Washington a contender had the

collections and libraries been centralized, but The Missouri Botanical Garden

(opened in 1859) and The New York Botanical Garden (founded in 1891)

were vying for recognition.

In his final year of life, with clouds of WW I darkening, Atkinson was

honored by election to the National Academy of Science. Funded by a USDA

grant to collect Amanita and Clavaria, he embarked upon a lengthy (albeit

truncated) field trip to Florida, Georgia, western Virginia, and North and

South Carolina. The man Atkinson had hired as an assistant was abruptly

drafted into the army, but Atkinson opined that two women might be able

to handle the job, and so it was (Haines 1987). Destinations included Chapel

96 ... Petersen

Hill, a former academic stop in Atkinson's career and now home to William

Chambers Coker (1872-1953), whose taxonomy was inferior in Atkinson’s

eyes (see Haines 1987). Later in the year he traveled to the Pacific Northwest,

where he was first ill with “Spanish flu” (an early victim of the 1918-1920

influenza pandemic), and then, no doubt weakened, in Tacoma, Washington,

fatally succumbed to pneumonia at age 67. His last photo—of Mt. Rainier—

survives in the Cornell Plant Pathology Herbarium.

Atkinson’s 1903 European travel coincided with the first powered flight

at Kitty Hawk. By the time of his death fifteen years later, biplanes, now a

common sight, were about to be put to use in World War I. In fact, during

Atkinson's lifetime, Google tells us that Bessemer forged steel from iron,

Nobel invented dynamite, Pasteur produced the first vaccines, Bell invented

the telephone, Edison introduced the incandescent light bulb, the Brooklyn

Bridge was opened in 1883 to horse-drawn and pedestrian traffic, and Henry

Ford produced Model T Ford cars on an assembly line (1903). Mycology was

keeping pace with civilization.

Estey (1994) reported that Canadian mycology was late to develop. Among

the earliest plant collectors were John Macoun (1831-1920; see above),

“who collected thousands of fungi from coast to coast in Canada and was

the first federally employed botanist to publish on fungi” (Estey 1994).

Somewhat later, James Fletcher (1852-1908) migrated from England and was

appointed Dominion Entomologist and Botanist in 1884, shortly before the

Central Experimental Farm in Ottawa became a reality. He was instrumental

in establishing the National Collection of Vascular Plants, the National

Herbarium, the National Arboretum and Botanic Garden. He was also an avid

field researcher who personally collected thousands of plant specimens. He

conducted successful research on effective treatments of various agricultural

pests, but this involved experimenting with toxic materials such as arsenic

and lead powder in the days before people took modern precautions. He died

of cancer at age 56, and there is a significant chance that his work with toxic

chemicals was to blame. As one commentator remarked: “Dying young, at

56, was a bad PR move, however.’ Fletcher never had a chance to write his

memoirs, and his name faded from public view

C.G. Lloyd (1908) mused: “As we look back over the past few years, it is

appalling to consider the inroads that have been made among our American

mycologists by death. First, we lost J.B. Ellis, then Dr. Wm Herbst, then Professor

Mycology in 19th century North America ... 97

A.P. Morgan, then Professor L.M. Underwood and finally W.A. Kellerman”

Together with individuals in this sketch who were still living, most of these

individuals had personally contributed to mycology and surely through their

exsiccati, publications, journals, teaching laboratories and especially students,

had set mycology on an upward curve into the 20" century. But that’s another

story...

Epilogue

In ephemerata accompanying Hesler’s (1975) compilation he kept cross-

indices on particular datum points. One dealt with mycologists’ birth- and

death-dates and the result is telling. From 1800 through 1820, three mycological

births were recorded. In the 1820s, births = three; in the 1830s, five; in the

1840s, seven; in the 1850s, 19; in the 1860s, 15 (surely a reflection of war-time);

in the 1870s, 20 (see TaBLE 1). Births after 1880 meant lives after mycology had

reached critical mass.

Ihave augmented Hesler’s original to include the names of ten (*) mycologists

(or influential botanists) treated here but missing from the 1975 list. Those not

mentioned or treated in depth here are denoted by “.

TABLE 1. North American mycological timeline

Pre-1800s

Lewis Davip DE SCHWEINITZ 1780-1834

EzRA MICHENER 1794-1887

*JOHN TORREY 1796-1873

1800 — 1810s

*GEORGE ENGELMANN 1809-1884

CHARLES CHRISTOPHER FROST 1805 — 1880

GEORGE WILLIAM CLINTON 1807 — 1885

*MosEs ASHELY CURTIS 1808-1872

*Asa GRAY 1810-1888

*CHARLES WRIGHT 1811-1884

* AUGUSTUS FENDLER 1813-1883

*HENRY WILLIAM RAVENEL 1814-1887

BENJAMIN MATLOCK EVERHART 1818 — 1904

1820s

GEORGE MARTIN 1826 — 1886 [Not G.W. MarTIN]

ELLIOTT CALVIN HOWE 1828 — 1899

JoB BICKNELL ELLIS 1829 — 1905

98 ... Petersen

1830s

*JOHN MACOUN 1831-1920

*Fr. A.B. LANGLOIS 1832-1900

CHARLES HorTON PECK 1833 — 1917

WILLIAM HERBST 1833 — 1906

Mary ELIZABETH BANNING 1832 -1901

ALBERT NELSON PRENTISS 1836 — 1896

ANDREW PRICE MORGAN 1836 — 1907

1840s

CHARLES MCILVAINE 1840 — 1909

WILLIAM WIRT CALKINS 1842 - 1914

WILLIAM GILSON FARLOW 1844 - 1919

CHARLES EDWIN BESSEY 1845 — 1915

FLORA WAMBAUGH PATTERSON 1847 - 1928

ASAMUEL MILLS TRACY 1847 - 1920

THOMAS HUSTON MACBRIDE 1848 — 1934

1850s

WILLIAM A. KELLERMAN 1850 — 1908

GEORGE EDWARD MASSEE 1850 — 1917

JOSEPH CHARLES ARTHUR 1850 — 1942

AJAMES FLETCHER 1853- 1908

ELAM BARTHOLOMEW 1852 — 1934

ABYRON DAVID HALSTED 1852 — 1918

JOHN DEARNESS 1852 — 1954

LucIEN MArcus UNDERWOOD 1853 — 1907

EDWARD WILLET DORLAND HO.Lway 1853 — 1923

GEORGE FRANCIS ATKINSON 1854 — 1918

FRANKLIN SUMNER EARLE 1854 - 1929

AERWIN FRINK SMITH 1854 — 1927

CHARLES EDWARD FAIRMAN 1856 - 1934

*EDWARD T. HARPER 1857-1921

NWILLIAM TRELEASE 1857 — 1942

HowarD ATWOOD KELLY 1858 - 1943

AROLAND THAXTER 1858 — 1932

A ARTHUR BLISS SEYMOUR 1859 — 1933

CurRTIS GATES LLOYD 1859 — 1926

EDWARD ANGUS BuRT 1859 - 1939

1860s

ASAMUEL MCCUTCHEN BAIN 1860 - 1915

ROBERT ALMER HARPER 1862 — 1946

LOUIS HERMAN PAMMEL 1862 — 1931

Mycology in 19th century North America ...

ELLSWORTH BETHEL 1863 — 1925

ABEVERLY THOMAS GALLOWAY 1863 — 1938

AGEORGE GRANT HEDGCOCK 1864 — 1946

CORNELIUS LOTT SHEAR 1865 — 1956

AMASON BLANCHARD THOMAS 1866 — 1912

“HOWARD JAMES BANKER 1866 — 1940

NHENRY CURTIS BEARDSLEE 1865 — 1948

GEORGE PERKINS CLINTON 1867 - 1927

WILLIAM HENRY LONG 1867 - 1947

AELIZABETH EATON MORSE 1864 - 1950

WILLIAM ALPHONSO MURRILL 1869 - 1957

CALVIN H. KAUFFMAN 1869 - 1931

1870s

ADavipb Ross SUMSTINE 1870 — 1965

AELIAS JUDAH DURAND 1870 — 1922

ROBERT HAGELSTEIN 1870 — 1945

GEORGE RICHARD LYMAN 1871 — 1926

AFRANK LINCOLN STEVENS 1871 — 1934

AWILLIAM STURGIS THOMAS ~1871 — 1941

CHARLES THOM 1872 — 1956

AGERTRUDE SIMONA BURLINGHAM 1872 — 1962

WILLIAM CHAMBERS COKER 1872 — 1953

ABENJAMIN MINGE DUGGAR 1872 — 1956

ANEDWARD MARTINIUS GILBERT 1872 — 1956

Louris CHARLES CHRISTOPHER KRIEGER 1873 — 1940

AFREDERIC EDWARD CLEMENTS 1874 — 1943

ARTHUR HENRY REGINALD BULLER 1874 — 1944

ERNST ATHERN BESSEY 1877 - 1957

AVERA K. CHARLES 1877 -— 1954

AFRED JAY SEAVER 1877 — 1970

LEIGH HUMBOLDT PENNINGTON 1877 — 1929

‘HERBERT HICE WHETZEL 1877 — 1944

Cognizant of his pride of alma mater, Cornell, Hesler also compiled a list

of schools that produced mycologists who were eligible for inclusion in

his compilation (i.e. death well before 1975, and chiefly mycologists, not

phytopathologists). Not surprisingly, Cornell led the list with Harvard a

distant second. Personages who acted as mycological magnets were easy

to identify. A short table showing school name, student name, date of

degree and major professor follows (TABLE 2 arranged, as Hesler had it,

alphabetically).

100 ... Petersen

TABLE 2. American mycologists: educational origins and professors

ALBANY STATE NORMAL SCHOOL

C.H. Peck

C.L. Shear

UNIVERSITY OF CHICAGO

EL. Stevens

G.W. Martin

COLUMBIA UNIVERSITY

H.J. Banker

G.S. Burlingham

R.W. Benham

ED. Kern

CORNELL UNIVERSITY

W.A. Kellerman

G.E Atkinson

J.C. Arthur

E.J. Durand

B.M. Duggar

W.A. Murrill

H.H. Whetzel

V.K. Charles

HS. Jackson

C.W. Edgerton

H.M. Fitzpatrick

L.R. Hesler

A.J. Mix

C.E. Chardon

J.H. Miller

HARVARD UNIVERSITY

W.G. Farlow

B.D. Halsted

A.B. Seymour

Roland Thaxter

C.H. Kauffman

E.A. Burt

B.M. Duggar

G.P. Clinton

W.H. Weston

D.H. Linder

L.M. Ames

Undergr.

PhD

M.A.

PhD

B.A.

D. Sci.

PhD.

PhD

B.A.

PhD

M.S.

PhD

B.A.

M.D.

D. Sc.

1883-1884

PhD

B.A.

PhD

M.A.

D.Sci.

PhD

1841

1888

1900

1928

1901

1908

1931

1921

1874

1885

1886

1895

1898

1900

1906

1903

1905

1908

L913

1914

1916

1921

1928

1866

1870

1878

Farlow

1888

1895

1902

1916

1926

1933

Burt

Underwood

Prentiss

Atkinson

Whetzel

Reddick & al.

Fitzpatrick

Asa Gray

Farlow

Thaxter

Mycology in 19th century North America... 101

UNIVERSITY OF ILLINOIS

A.B. Seymour M.S. 1886 Burrill

ES. Earle M.S. 1886 Burrill

G.P. Clinton M.S. 1894 Burrill

L.R. Tehon PhD 1934 FEL. Stevens

IOWA STATE UNIVERSITY

Flora W. Patterson M.A. 1895

ED. Kern M.S. 1904

FJ. Seaver PhD 1912

JoHNS HOPKINS UNIVERSITY

W.C. Coker PhD 1901 D. Johnson

UNIVERSITY OF MICHIGAN

E.F Smith PhD 1889

C.H. Kauffman PhD 1907 Newcomb

L.H. Pennington PhD 1909 Newcomb

E.B. Mains PhD 1916 Kauffman

D.V. Baxter PhD 1926 Kauffman

MICHIGAN STATE UNIVERSITY

A.N. Prentiss B.S. 1861

M.S. 1864

B.D. Halsted M.S. 1874 Prentiss

D.Sci. 1878 Farlow

C.E. Bessey 7 : Prentiss

S.M. Tracy : ¢ Prentiss

UNIVERSITY OF MISSOURI

Charles Thom PhD 1899

MONMOUTH COLLEGE

T.H. Macbride B.A. 1869

M.A. 1873

UNIVERSITY OF NEBRASKA

E.A. Bessey LL.D. (Hon.) 1946

C.L. Shear M.A. 1898 CE. Bessey

FE. Clements PhD. 1898

UNIVERSITY OF NORTH CAROLINA

W.H. Ravenel LL.D. (Hon.) 1886

PURDUE UNIVERSITY

ED. Kern MS. 1907. = Arthur

RUTGERS UNIVERSITY

EL. Stevens M.S. 1897 Halsted

102 ... Petersen

UNIVERSITY OF SOUTH CAROLINA

W.H. Ravenel B.A. 1832

SYRACUSE UNIVERSITY

L.M. Underwood M.A. 1877

LL.D. (Hon.) 1906

UNIVERSITY OF TEXAS

W.H. Long M.A. 1900

UNIVERSITY OF TORONTO

H.W. Groves PhD. 1935

UNION UNiversiTy (Schenectady, NY)

J.B. Ellis B.A. 1871

C.H. Peck B.A. 1859

M.A. 1862

D. Sc. (Hon.) 1908

WASHINGTON UNIVERSITY (St. Louis)

L.H. Pammel PhD. 1896

L.O. Overholts PhD. 1915 Burt

S.M. Zeller PhD. 1917. Burt

WILLIAMS COLLEGE

M.A. Curtis M.A. 1827

UNIVERSITY OF WISCONSIN

E.M. Gilbert PhD. 1907 R.A. Harper

YALE UNIVERSITY

N.E. Stevens PhD. 1921 Evans

G.L. Zundel PhD. 1928 G.P. Clinton

Acknowledgments

In the fall of 1975, Lexemuel Ray Hesler, then retired from botany department head

and dean of liberal arts, strolled into my office with a thick volume informally held

together by a plastic claw-like device. “You might be interested in this,” he declared.

“Tve been working on it for some time.” And with that, as was his wont, he left. The

cover did not deal with some fungal genus as I might have expected. If so, he would

have sent it to Alexander Hanchett Smith, his mentor and alter-ego who would have

been co-author. Instead, the title started with “Biographical sketches of deceased

North American mycologists...” I was surprised, for there had been no sign that he

had, in fact, been working on such a volume, but most of the names featured were

only names for me and as I read one or two entries I was not seized with any particular

passion for his subject.

Among individuals to whom he had written for information was Clark T.

Rogerson at the New York Botanical Garden, to whom he confided his intent. [Letter

Mycology in 19th century North America ... 103

dated Dec. 24, 1974, Christmas Eve]: “What I have long wanted to do is to have handy

for graduate students a series of short biographies ... At last, I have decided to bring

together what I can, compiled from the published biographies, for our graduate

students here.”

In conversation later, Hesler opined that the volume of 124 thumbnail biographies

ranging from a half-—page to five pages, was not ready for publication because he had

not exhausted available background material.

Here we are, some 43 years later, and I am at the same point in life as Hesler was

then. A longtime interest has been mycological history, and it may be time to bring

some of Hesler’s work into a context. This paper is aimed in that direction with thanks

for the inspiration and contribution.

Meredith Blackwell and Lorelei Norvell acted as outside reviewers and Lorelei

spent many hours putting the manuscript in order in addition to ordinary editorial

tasks. Shaun Pennycook reviewed a late version of the paper and made constructive

comments.

Thanks are offered to Dr. Beth Mullen for information concerning Quaker “Plain

Language.’ Dr. Teresa Iturriaga, Cornell University Plant Pathology Herbarium,

generously furnished material on Augustus Fendler. Ms Ann Viera of the Pendergrass

Agricultural Library at the University of Tennessee provided digitized copies of

obituaries in early volumes of PHyTOPATHOLOGY.

Many thanks are due to Mrs. Dorene Neilans Setliff, who, in a term paper for a

natural history course, included several of the photo illustrations co-opted here. Little

did she know in 1963, that over a half century later her thoroughness would still be

appreciated.

In juxtaposition to Farlow’s purchase of rare mycological literature and my many

joyous hours in the stacks of The New York Botanical Garden's library, acknowledgment

and thanks must be extended to online sources of entire runs of mycological journals

(and many others), search engines, WIKIPEDIA (although with due caution), and

especially JStor and CyBERLIBER for hosting a plethora of literature. INDEX OF FUNGI

contributes other data of great use. The research for this sketch was completed in

months hardly leaving my office (although with some grieving over the inability to

travel as widely as ever).

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MY COTAXON

January-March 2019—Volume 134, pp. 111-117

https://doi.org/10.5248/134.111

Coronosporidium ecuadorianum gen. & sp. nov.

from submerged decaying leaf from Ecuador

DAYNET SOSA??’, ADELA QUEVEDO!', FERNANDO ESPINOZA’,

LIZETTE SERRANO’, FREDDY MAGDAMA’, MARCOS VERA’,

SIMON PEREZ-MARTINEZ’”, ELAINE MALOSSO?;, RAFAEL E CASTANEDA-RUvUIZ?*

"Escuela Superior Politécnica del Litoral, ESPOL, (CIBE),

Campus Gustavo Galindo Km. 30.5 Via Perimetral,

PO. Box 09-01-5863, Guayaquil, Ecuador

? Universidad Estatal de Milagro (UNEMI), Facultad de Ingenieria,

Cdla. Universitaria Km. 1.5 via Milagro-Km26. Milagro 091706, Guayas, Ecuador

* Centro de Biociéncias, Departamento de Micologia, Universidade Federal de Pernambuco,

Avenida da Engenharia, s/n Cidade Universitaria, Recife, PE, 50.740-600, Brazil

‘Instituto de Investigaciones Fundamentales en Agricultura (INIFAT),

Tropical ‘Alejandro de Humboldt’, OSDE, Grupo Agricola,

Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200

* CORRESPONDENCE TO: dasosa@espol.edu.ec

ABSTRACT—A new genus and species Coronosporidium ecuadorianum are described and

illustrated. The fungus, found on a submerged decaying leaf of an unidentified plant, is

distinguished by blastic production of stauroconidia, obpyramidal coronate, 1-septate,

and inequilateral sympodial extensions of the conidiogenous cells after rhexolytic conidial

secession.

KEY worpDs—asexual fungi, freshwater, hyphomycetes, taxonomy, tropics

Introduction

The diversity of microfungi in the Ecuadorian Rainforest has received little

attention, especially in freshwater habitats. During a survey of hyphomycetes

associated with plant litter submerged in streams from rainforest and cacao

plantations near the Patul river, La Aurora (Fic. 1), Azuay province, southwest

112 ... Sosa & al.

“a

al ae

Pes

. ii)

Fic. 1. Streams in La Aurora rainforest, Azuay Province, Ecuador.

Coronosporidium ecuadorianum gen. & sp. nov. (Ecuador) ... 113

Ecuador, we collected a fungus that differs remarkably from all previously

described genera (Seifert & al. 2011) for which we propose a new genus and

species.

Materials & methods

Samples of decaying plant materials were collected and placed in plastic bags for

transport to the laboratory, where they were washed, treated according to Castaneda-

Ruiz & al. (2016), and placed in humid chambers. Several attempts to obtain this

species in pure culture were unsuccessful after using a flamed needle to transfer

conidia to corn meal agar mixed 1:1 with carrot extract and incubating at 25 °C.

Mounts were prepared in PVL (polyvinyl alcohol, lactic acid) and measurements were

made at 1000x magnification. Microphotographs were obtained with an Olympus

BX51 microscope equipped with bright field and Nomarski interference optics.

The type specimen is deposited in the Herbarium of Universidade Federal de

Pernambuco, Recife, Brazil (URM).

Taxonomy

Coronosporidium R.F. Castafieda, Quevedo & D. Sosa, gen. nov.

INDEX FUNGORUM IF 554557

Differs from Coronospora by its polyblastic conidiogenous cells with minute separating

cells attached to the conidia, which secede by rhexolytic fracture.

TYPE SPECIES: Coronosporidium ecuadorianum R.F. Castaneda & al.

EryMoLoGy: Corono- (Latin) meaning crowned + -sporidium (Latin) referring to the

conidia.

Co.tontgs on the natural substrate effuse. CONIDIOPHORES macronematous,

mononematous, unbranched, subhyaline to pale brown. CONIDIOGENOUS

CELLS monoblastic and polyblastic, integrated, indeterminate, with holoblastic

sympodial extension. SEPARATING CELLS broadly cylindrical. Conidial secession

rhexolytic. Conrp1< solitary, acrogenous or acropleurogenous, stauromorphic,

coronate, gray to brown, euseptate.

Coronosporidium ecuadorianum R.F. Castafieda, Quevedo & D. Sosa,

sp. nov. FIGS 2, 3

INDEX FUNGORUM IF 554558

Differs from Coronospora dendrocalami by the rhexolytic conidial secession and the

cylindrical basal frill of the conidia.

Type: Ecuador, Azuay province, La Aurora, 79°21 W 2°34 S, alt. 499 m, on submerged

decaying leaf of an unidentified plant in stream, 4.III.2018, coll. A. Quevedo Pinos,

E. Leiva Pantoja & J. Ochoa Torres (Holotype, URM 91191).

EryMoLoGy: ecuadorianum (Latin) referring to Ecuador, the type locality.

114 ... Sosa & al.

CoLonizs on the natural substrate effuse, hypophyllous, granulose, brown

to golden gray brown. Mycelium superficial and immersed, composed of

septate, unbranched, smooth, 2-3 um diam, pale brown to subhyaline hyphae.

CONIDIOPHORES macronematous, mononematous, erect, flexuous, straight or

geniculate, unbranched, smooth, 1-3-septate, pale brown to brown, 10-30 x

3.5-4.5 um, smooth. CONIDIOGENOUS CELLS polyblastic, rarely monoblastic,

cylindrical, integrated, indeterminate, terminal, becoming intercalary with

frequent sympodial extensions, smooth, pale brown to brown, 8-20 x 3 um.

SEPARATING CELLS broadly cylindrical, pale brown or pale smoked gray brown,

4-6 um long, fracturing and remaining as a persistent short peg. CONIDIAL

SECESSION rhexolytic. CONIDIOGENOUS LOCI a conspicuous minute collar,

slightly ringed after the conidial secession. Conip1a solitary, acrogenous,

mostly acropleurogenous, broad pyriform, broad turbinate, coronate,

inequilaterally 1-euseptate, smoked gray brown to dark gray, smooth-walled,

30-40 x 20-30 um, with a conspicuous basal frill, 2-3 um, and toward upper

part an acuminate to obtuse, subtriangular central protuberance, 5-18 um

long, encircled near the base by others 4-6 obtuse or rounded, erect or oblique,

corniform protuberances, 4-8 um long, disposed more or less radially at the

apical margin.

NoTE: Coronosporidium ecuadorianum is superficially similar to Coronospora

dendrocalami M.B. Ellis in conidial shape and the polyblastic sympodial

extension of the conidiogenous cells, but C. dendrocalami has cicatrized,

slightly melanized loci of the conidiogenous cells (Ellis 1971). Anacoronospora

diversiseptata J.S. Monteiro & al. can also be compared with C. ecuadorianum

in conidial shape, but A. diversiseptata has disto- and euseptate conidia

produced after enteroblastic percurrent (sometimes sympodial) extension, and

its conidiogenous loci are cicatrized and strongly melanized (Monteiro & al.

2016).

Henicospora amazonensis J.S. Monteiro & al. and H. coronata B. Sutton &

P.M. Kirk resemble C. ecuadorianum in conidial shape and rhexolytic conidial

secession, but the Henicospora species are characterized by monoblastic,

determinate conidiogenous cells and the absence of ringed minute collar

remnant on the conidiogenous loci (Monteiro & al. 2014, Kirk & Sutton 1980).

Fic. 2. Coronosporidium ecuadorianum (holotype, URM 91191). A-E. Conidia with a basal frill of

the separating cells. F Conidiogenous cells, separating cells, and attached conidium. G. Rhexolytic

conidial secession in the separating cell. H, I. Conidiogenous cells with ringed collarette-like

conidiogenous loci. Scale bars = 10 um.

Coronosporidium ecuadorianum gen. & sp. nov. (Ecuador) ... 115

116... Sosa & al.

Fic. 3. Coronosporidium ecuadorianum (holotype, URM 91191).. A Conidia. B. Conidiogenous

cells with the ringed collarette-like conidiogenous loci. Scale bars = 10 um.

The rhexolytic fracture and ringed, minute collar remnant on the loci in

C. ecuadorianum are almost the same as observed in Matsushimiella paraensis

J.S. Monteiro & al. and M. queenslandica (Matsush.) R.F. Castafieda & Heredia,

but those species have phragmoconidia, which are distoseptate, ellipsoidal,

obovate, long ovoid, or oblong, and lacking the conidial arms (Castaneda-Ruiz

& al. 2001, Matsushima 1989, Monteiro & al. 2015a,b).

Coronosporidium ecuadorianum gen. & sp. nov. (Ecuador) ... 117

Acknowledgments

We are indebted to Dr. Luis EP. Gusmao and Dr. De-Wei Li for their critical

reviews. The authors thank to Mr. Arnold Quevedo Mora for facilities and Enoy

Leiva Pantoja and Jorge Ochoa Torres for provide collected samples and M. Caraballo

Fernandez for the technical assistance. The authors are grateful to Escuela Superior del

Litoral (ESPOL), CIBE for financial support and the International Society for Fungal

Conservation for facilities. RFCR is grateful to the Cuban Ministry of Agriculture.

We acknowledge the websites provided by Dr. PM. Kirk (Index Fungorum) and

Dr. K. Bensch (MycoBank). Dr. Lorelei L. Norvell’s editorial review and Dr. Shaun

Pennycook’s nomenclature review are greatly appreciated. [Submitted on 18 August

2018, the manuscript was lost in electronic transmission and not received by

MycotTaxon until 22 February 2019.]

Literature cited

Castafieda-Ruiz RF, Heredia G, Reyes M, Arias RM, Decock C. 2001. A revision of the genus

Pseudospiropes and some new taxa. Cryptogamie, Mycologie 22: 1-18.

Castafieda-Ruiz RF, Heredia G, Gusmao LFP, Li DW. 2016. Fungal diversity of Central and

South America. 197-217, in: DW Li (ed.). Biology of Microfungi. Springer International

Publishing. https://doi.org/10.1007/978-3-319-29137-6_9

Ellis MB. 1971. Dematiaceous hyphomycetes. X. Mycological Papers 125..30 p.

Kirk PM, Sutton BC. 1980. Henicospora gen. nov. (hyphomycetes). Transactions of the British

Mycological Society 75: 249-253. https://doi.org/10.1016/S0007-1536(80)80087-8

Matsushima T. 1989. Matsushima mycological memoirs no. 6. Matsushima Fungus Collection,

Kobe

Monteiro JS, Carmo LT, Fiuza PO, Ottoni BMP, Gusmao LFP, Castafieda-Ruiz RF 2014.

New species of microfungi from Brazilian Amazon rainforests. Mycotaxon 127: 81-87.

https://doi.org/10.5248/127.81

Monteiro JS, Gusmao LFP, Castafieda-Ruiz RF. 2015a. A new species of Matsushimiella

from submerged leaves in the Brazilian Amazon Forest. Mycotaxon. 130: 311-314.

https://doi.org/10.5248/130.311

Monteiro JS, Gusmao LFP, Castafieda-Ruiz RE 2015b. Validation of the name Matsushimiella

paraensis. Mycotaxon 130(4): vu. https://doi.org/10.5248/130-4.cvr

Monteiro JS, Gusmao LFP; Castafieda-Ruiz RE. 2016. Anacoronospora diversiseptata gen. & sp. nov.

from Brazil. Mycotaxon 131: 185-192. https://doi-org/10.5248/131.185

Seifert K, Morgan-Jones G, Gams W, Kendrick B. 2011. The genera of hyphomycetes. CBS

Biodiversity Series 9. 997 p.

MY COTAXON

January-March 2019—Volume 134, pp. 119-124

https://doi.org/10.5248/134.119

Trichoglossum tetrasporum, newly recorded from India

SANJIT DEBNATH*™*, KRIPAMOY CHAKRABORTY’,

BADAL KUMAR DaTTA?;, PANNA Das’, AJAY KRISHNA SAHA"

' Mycology and Plant Pathology Laboratory, * Microbiology Laboratory,

and * Plant Taxonomy and Biodiversity Laboratory,

Department of Botany, Tripura University,

Suryamaninagar-799022, India

*CORRESPONDENCE TO: sanjitdebnath2888@gmail.com

ABSTRACT— The uncommon earth tongue, Trichoglossum tetrasporum, is recorded for the

first time from India. Macroscopic and microscopic characters of the species are described

and illustrated along with its world distribution.

Key worps—apothecia, Geoglossaceae, Geoglossales, paraphyses, setae

Introduction

Earth tongues (Geoglossomycetes; Schoch & al. 2009) are a widely distributed

group of fungi that have been under intense investigation since the descriptions

of Geoglossum Pers. in the late 18" century and Trichoglossum Boud. in the late

19" century. The widely distributed group includes such genera as Geoglossum,

Nothomitra Maas Geest., Sarcoleotia S. Ito & S. Imai, and Trichoglossum (Schoch

& al. 2009, Hustad & al. 2011).

Trichoglossum is separated from Geoglossum based on the production of

abundant dark setae (hairs) in the hymenium and on the stipe (Mains 1954,

Sinden & Fitzpatrick 1930). About 19 Trichoglossum species are currently

recognized (Kirk & al. 2008, Hustad & al. 2013), although Index Fungorum

(2018) lists 23 species and 48 varieties, which include a number of synonyms

and dubious taxa (Hustad & al. 2013).

120 ... Debnath & al.

Sinden & Fitzpatrick (1930) described Trichoglossum tetrasporum from

New York State, USA. It has since been reported from China (Tai 1944, Mains

1954), Great Britain (Nannfeldt 1942), Netherlands (Maas Geesteranus 1964),

Denmark (Benkert 1976), Spain (Galan & Rubio 1998), and Italy (Lantieri

2011). Here we extend its distribution range to India. The macro- and micro-

morphological features of the Indian collection are described and illustrated

below.

Materials & methods

Collection area

The samples were collected from Suryamaninagar (23°45.70’N 91°15.79’E; 80 m

asl), West Tripura, Northeast Region, India. Tripura, one of seven states in north-

eastern India, occupies a geographical area of 10,491 km’, of which 6292 km?

(c. 60%) is covered with forest. The state is located at 22°57’-24°33’N 91°10’-92°20’E.

Its climate averages are: temperature 10°C-35°C, relative humidity 50-80%, and

annual rainfall 2100 mm.

Specimen collection and identification

The specimens were collected from natural habitats using forceps and

photographed. The apothecia and ascogenous portions were measured just after

collection from field. Samples were dried for 24 hours at 45-55°C in a ROV/DG hot-

air oven. After drying, samples were preserved for further analysis in a polyethylene

bag also containing 1,4-dichlorobenzene (Debnath & al. 2017). The dried material was

preserved in the Mycology and Plant Pathology laboratory, Department of Botany,

Tripura University, bearing the collection tag MCCT (Mushroom Culture Collection

Tube)-380. A small portion of a hand-sectioned specimen was squash-mounted

in 5% potassium hydroxide (KOH) solution. Slides were also mounted in Melzer’s

reagent, Lugol's solution, and lactoglycerol and Congo Red solution for observation

of anatomical characters under an Olympus CX21i bright field microscope equipped

with a SLI500 camera. Diagnostic features including the asci, ascospores, paraphyses

and setae were studied and compared with different descriptions for identification

(Sinden & Fitzpatrick 1930, Mains 1954, Maas Geesteranus 1965). The collection

was deposited in Central National Herbarium, Botanical Survey of India, Howrah,

West Bengal, India (CAL).

Taxonomy

Trichoglossum tetrasporum Sinden & Fitzp., Mycologia 22: 60 (1930) Fics 1, 2

APOTHECIA Scattered to gregarious, black or brownish black, clavate to

capitate, 10-90 mm long; head compressed, 20-40 x 20-30 mm diam., hirsute

from setae; odourless. STERILE PORTION OR STIPE cylindrical, slender, 2-3 mm

thick, hirsute from setae. Asci clavate, 4-spored, 170-195 x 16-20 um, rounded

Trichoglossum tetrasporum, new for India... 121

20um

Fic. 1. Trichoglossum tetrasporum (CAL 1687): A. Mature fruit body in natural habitat; B. Cross

section of head showing mature asci, setae, and paraphyses; C. 4-spored asci, dark brown setae,

and bulbous paraphyses with curved capitate tips; D. Immature stage of asci with paraphyses;

E. Slightly curved ascospores with 15 transverse septa; F. Irregularly flexuous setae of stipe. Scale

bars: A = 10 mm; B-D, F = 20 um; E = 10 um.

at apex, short-pedicellate. Ascospores parallelly arranged in the upper part of

the ascus; fusoid to subcylindrical, smooth, 130-145 x 5-7 um, with 10-15

transverse septa when mature (fewer when immature), slightly curved, dark

brown when mature; apical ends narrowed and rounded; individual cells

often uniguttulate. HYMENIAL SETAE 180-220 x 5-13 um, smooth, straight to

flexuous. PARAPHYSES 2-4 um wide, sub-cylindrical toward the bottom but

often with some swollen, filamentous, septate, and curved at tips. STIPE SETAE

irregularly waved outline, dark brown, measuring 170-200 x 4-12 um.

SPECIMEN EXAMINED: INDIA, NORTHEAST REGION, West Tripura, Suryamaninagar,

23°45.70 N 91°15.79 E, 80 m asl, saprobic on soil with decomposed plants, 18 September

2017, Debnath & Saha (CAL 1687).

DISTRIBUTION: Trichoglossum tetrasporum has been reported from USA, Great

Britain, Netherlands, Denmark, Spain, Italy, China, and India.

Discussion

Trichoglossum tetrasporum is characterized by a hymenium with numerous

acuminate brown setae, 4-spored asci, and 10-15-septate ascospores. It is

122 ... Debnath & al.

Fic. 2. Trichoglossum tetrasporum (CAL 1687): G. Mature fruit body; H. 4-spored asci, setae,

and bulbous paraphyses with curved capitate tips; I. Immature stage of asci with paraphyses;

J. Slightly curved ascospores with 15 transverse septa; K. Irregularly flexuous setae of stipe. Scale

bars: G = 10 mm; H, I, K = 20 um; J = 10 um.

morphologically quite close to T. hirsutum (Pers.) Boud. and T! velutipes (Peck)

E.J. Durand but is distinguished by the size of asci and ascospores and number

of ascospore septa. Most authors report 15 septa for T: tetrasporum ascospores

(Sinden & Fitzpatrick 1930, Seaver 1951, Ellis & Ellis 1988, Galan & Rubio

1998), which mostly corresponds with our findings. Mains (1954), however,

reported a much more variable number of septa (0-17) for the species.

Tai (1944) described Trichoglossum tetrasporum var. brevisporum EL. Tai and

T. yunnanense E.L. Tai from Yunnan, China. His description of T. tetrasporum

var. brevisporum showed similarities with our specimen but differed by its

smaller ascospores (117-137 x 6-7 um); and his description of T. yunnanense

differed from T. tetrasporum by its longer and narrower asci (237-294 x 19-22

um) and its longer (143-187 x 6-7 um) and differently shaped (cylindrical

clavate) ascospores. However, Mains (1954) considered T: yunnanense to

represent “a longer, more variable, septate-spored variant of T’ tetrasporum,

and recombined it as T. tetrasporum var. yunnanense (F.L. Tai) Mains. Species

Fungorum (2018) accepts both names as varieties of T: tetrasporum.

Trichoglossum tetrasporum, new for India ... 123

A literature survey found no report of T: tetrasporum from Asia other than

Tai (1944); our record from India therefore represents only the second Asian

report in 74 years.

Prabhugaonkar & Pratibha (2017) documented a new record of T. rasum Pat.

from Asia, also providing a preliminary phylogeny of Trichoglossum showing

T. hirsutum as polyphyletic, in concordance with previous findings by Hustad

& al. (2013). Phylogenetic investigations of T: tetrasporum from different

continents may clarify whether the differences in spore size and septation

reported by different authors represent one highly variable species with a wide

distribution or include one or more cryptic species.

Acknowledgements

The authors are grateful to the Head of the Department of Botany, Tripura

University, for laboratory facilities. AKS is gratefulto the Department of Biotechnology

(DBT), Government of India, for the financial assistance received through project

(Sanctioned Order no. BT/463/NE/TBP/2013). The authors acknowledge the help

rendered in pre-submission reviews by Michael Loizides (Limassol, Cyprus) and Dra.

Larissa Trierveiler-Pereira (Instituto de Botanica, SP, Brazil). The authors express

their appreciation to Dr. Shaun Pennycook (New Zealand) for his useful comments

and nomenclatural suggestions and are highly thankful to Botanical Survey of India

(Government of India, Ministry of Environment, Forest Climate Change, Salt Lake

City, Kolkata, India) for providing the herbarium accession number of the studied

fungal specimen. The authors have no potential conflict of interest.

Literature cited

Benkert D. 1976. Bemerkenswerte Ascomyceten der DDR. II. Die Gattungen Geoglossum und

Trichoglossum in der DDR. Mykologisches Mitteilungsblatt 20(3): 47-92.

Debnath S, Roy Das A, Karmakar P, Debnath G, Das P, Saha AK. 2017. Checklist of mushroom

diversity in West Tripura, North-East India. 205-213, in: S Sinha, RK Sinha (eds). Trends in

Frontal Areas of Plant Science Research. Narosa Publishing House Pvt. Ltd.

Ellis MB, Ellis JP. 1988. Microfungi on miscellaneous substrates: an identification handbook.

Slough: Richmond Publishing. 244 p.

Galan R, Rubio E. 1998. Sarcoscypha austriaca y Trichoglossum tetrasporum, dos nuevos

ascomicetos ibéricos, procedentes de Asturias. Belarra 14: 5-9.

Hustad VP, Miller AN, Moingeon JM, Priou JP. 2011. Inclusion of Nothomitra in

Geoglossomycetes. Mycosphere 2(6): 646-654. https://doi.org/10.5943/mycosphere/2/6/5

Hustad VP, Miller AN, Dentinger BTM, Cannon PE 2013. Generic circumscriptions in

Geoglossomycetes. Persoonia 31: 101-111. https://doi.org/10.3767/003158513X671235

Index Fungorum. 2018. http://www.indexfungorum.org/Names/Names.asp

Kirk PM, Cannon, PF, Minter DW, Stalpers JA. 2008. Dictionary of the fungi, 10th edition.

CAB International, Wallingford, UK

Lantieri A. 2011. First record of Trichoglossum tetrasporum Sinden & Fitzp. (Helotiales,

Geoglossaceae) from Italy. Plant Biosystems 145(1): 116-119.

https://doi.org/10.1080/11263504.2010.543788

124 ... Debnath & al.

Maas Geesteranus RA. 1964. De Fungi van Nederland. I. Geoglossaceae — Aardtongen.

Wetenschappelijke mededelingen van de Koninklijke Nederlandse Natuurhistorische

Vereeniging 52: 1-24.

Maas Geesteranus RA. 1965. Geoglossaceae of India and adjacent countries. Persoonia 4(1): 19-46.

Mains EB. 1954. North American species of Geoglossum and Trichoglossum. Mycologia 46:

586-631. https://doi.org/10.1080/00275514.1954.12024398

Nannfeldt JA. 1942. The Geoglossaceae of Sweden (with regard also to the surrounding countries).

Arkiv for botanik 30: 1-75.

Prabhugaonkar A, Pratibha J. 2017. New record of Trichoglossum rasum from Asia. Mycosphere

8(4): 583-591. https://doi.org/10.5943/mycosphere/8/4/7

Schoch CL, Wang Z, Townsend JP, Spatafora JW. 2009. Geoglossomycetes cl. nov., Geoglossales

ord. nov. and taxa above class rank in the Ascomycota Tree of Life. Persoonia 22: 129-138.

https://doi.org/10.3767/003158509X46 1486

Seaver FJ. 1951. The North American cup-fungi (Operculates). New York: Seaver. 284 p.

Sinden JW, Fitzpatrick HM. 1930. A new Trichoglossum. Mycologia 22: 55-61.

https://doi.org/10.1080/00275514.1930.12016981

Species Fungorum. 2018.

http://www.speciesfungorum.org/Names/SynSpecies.asp?RecordID=183377

Tai FL. 1944. Studies in the Geoglossaceae of Yunnan. Lloydia 7: 146-162.

MY COTAXON

January-March 2019—Volume 134, pp. 125-137

https://doi.org/10.5248/134.125

Perenniporia mopanshanensis sp. nov. from China

CHANG-LIN ZHAO!” & XIANG Ma!

‘College of Biodiversity Conservation and Utilization & * Key Laboratory for

Forest Resources Conservation and Utilization in the Southwest Mountains of China,

Ministry of Education, Southwest Forestry University, Kunming 650224, P.R. China

" CORRESPONDENCE TO: fungichanglinz@163.com

ABSTRACT—A new poroid, white-rot, wood-inhabiting fungal species, Perenniporia

mopanshanensis, is proposed based on morphological and molecular characters. This species

from Yunnan Province, China, is characterized by resupinate basidiomes with a cream to

buff to straw pore surface, dimitic hyphal system with strongly dextrinoid, unbranched,

interwoven skeletal hyphae, and ellipsoid, non-truncate basidiospores (5.5-6.5 x 4-5 um)

with hyaline, distinctly thick, smooth, strongly dextrinoid walls. Phylogenetic analyses of

ITS+nLSU sequences showed that P mopanshanensis is a distinct taxon in the Perenniporia

sensu stricto clade and is sister to P. bannaensis.

Key worps—molecular phylogenetics, Polyporaceae, taxonomy, Truncospora, wood-rotting

fungi

Introduction

Perenniporia Murrill (Polyporaceae, Polyporales) is a large cosmopolitan

genus characterized by poroid basidiomata with thick-walled, ellipsoid to

distinctly truncate basidiospores that are cyanophilous and with variable

dextrinoid reactions (Ryvarden 1991). The Perenniporia hyphal system is di- or

trimitic: the generative hyphae with clamp connections and the skeletal hyphae

with walls that are cyanophilous and variably dextrinoid (Decock & Stalpers

2006). About 100 species are accepted in the genus (Gilbertson & Ryvarden

1987; Hattori & Lee 1999; Nunez & Ryvarden 2001; Dai & al. 2002, 2011;

Cui & al. 2007; Xiong & al. 2008; Choeyklin & al. 2009; Decock 2011, 2016;

Zhao & al. 2013; Ryvarden & Melo 2014; Jang & al. 2015; Ji & al. 2017).

126 ... Zhao & Ma

Recently, phylogenetic studies of Perenniporia based on sequences of

the internal transcribed spacer (ITS) region and the large subunit nuclear

ribosomal RNA gene (nLSU) have revealed several well-supported clades that

could be recognized as distinct genera (Robledo & al. 2009, Zhao & al. 2013).

Robledo & al. (2009) demonstrated that Perenniporia was phylogenetically

related to Abundisporus Ryvarden, Hornodermoporus Teixeira, Perenniporiella

Decock & Ryvarden, and Truncospora Pilat, while Zhao & al. (2013) have shown

that Perenniporia itself is polyphyletic. Several new species of Perenniporia

have been described based on ribosomal DNA sequences (Zhao & Cui 2013,

Jang & al. 2015, Ji & al. 2017)

During research on polypore diversity in southern China, an undescribed

species of Perenniporia was found. We present morphological and molecular

phylogenetic evidence that support the recognition of P. mopanshanensis as a

new species.

Materials & methods

The specimens studied are deposited at the herbarium of Southwest Forestry

University, Kunming, China (SWFC). Basidiomatal descriptions are based on field

notes. Special colour terms follow Petersen (1996). Anatomical observations were

obtained from the dried specimens and made using a light microscope following

Dai (2010). The following abbreviations were used: KOH = 5% potassium hydroxide,

CB = cotton blue, CB- = acyanophilous, IKI = Melzer’s reagent, IKI- = both inamyloid

and non-dextrinoid, L = mean spore length (arithmetic average of all spores),

W = mean spore width (arithmetic average of all spores), Q = variation in the L/W

ratios between the specimens studied, n (a/b) = number of spores (a) measured from

given number (b) of specimens.

The Omega EZNA HP Fungal DNA Kit was used to obtain genomic DNA from

dried specimens according to the manufacturer’s instructions with some modifications.

A 30 mg sample from a dried fungal specimen was ground to powder with liquid

nitrogen. The powder was transferred to a 1.5 ml centrifuge tube, suspended in 0.4 ml

of lysis buffer, and incubated in a 65°C water bath for 60 min. After that, 0.4 ml phenol-

chloroform (24:1) was added to each tube, and the suspension was shaken vigorously.

After centrifugation at 13,000 rpm for 5 min, 0.3 ml supernatant was transferred to

a new tube and mixed with 0.45 ml binding buffer. This mixture was transferred to

an adsorbing column (AC) for centrifugation at 13,000 rpm for 0.5 min. Then, 0.5

ml inhibitor removal fluid was added in AC for a centrifugation at 12,000 rpm for

0.5 min. After washing twice with 0.5 ml washing buffer, the AC was transferred to a

clean centrifuge tube, and 100 ml elution buffer was added to the middle of adsorbed

film to elute the genomic DNA. ITS region was amplified with primer pairs ITS5 and

ITS4 (White & al. 1990). Nuclear LSU region was amplified with primer pairs LROR

and LR7 (http://www. biology.duke.edu/fungi/mycolab/primers.htm). The ITS region

Perenniporia mopanshanensis sp. nov. (China) ... 127

was amplified by initial denaturation at 95°C for 3 min, then 35 cycles of 94°C for

40 s, 58°C for 45 s, and 72°C for 1 min with a final extension of 72°C for 10 min. The

nLSU region was amplified by initial denaturation at 94°C for 1 min, then 35 cycles at

94°C for 30 s, 48°C for 1 min, and 72°C for 1.5 min, ending with a final extension of

72°C for 10 min. The PCR products were purified and directly sequenced at Kunming

Tsingke Biological Technology Limited Company. All newly generated sequences were

deposited at GenBank (TABLE 1).

TABLE 1. Species, specimens, and GenBank accession numbers of studied sequences

GENBANK ACCESSION NO.

SPECIES NAME SAMPLE NO. REFERENCE

ITS nLSU

A. sclerosetosus MUCL 41438 FJ411101 FJ393868 Robledo & al. 2009

A. violaceus MUCL 38617 FJ411100 FJ393867 Robledo & al. 2009

Donkioporia expansa MUCL 35116 FJ411104 FJ393872 Robledo & al. 2009

PRT A fo Cui 6625 HQ876604 JE706340 Zhao & al. 2013

atissimus

H. martius Cui 7992 HQ876603 HQ654114 Zhao & al. 2013

MUCL 41677 FJ411092 FJ393859 Robledo & al. 2009

MUCL 41678 FJ411093 FJ393860 Robledo & al. 2009

has nt MUCL 45229 FJ411106 FJ393874 Robledo & al. 2009

Perenniporia africana Cui 8674 KFO18119 KFO18128 Present study

Cui 8676 KF018120 KFO018129 Present study

P. aridula Dai 12398 JQ001855 JQ001847 Zhao & al. 2013

Dai 12396 JQ001854 JQ001846 Zhao & al. 2013

P. bannaensis Cui 8560 JQ291727 JQ291729 Zhao & al. 2013

Cui 8562 JQ291728 JQ291730 Zhao & al. 2013

P. cinereofusca Dai 9289 KF568893 KF568895 Zhao & al. 2013

Cui 5280 KF568892 KF568894 Zhao & al. 2013

P. corticola Cui 1248 HQ848472 HQ848482 Zhao & al. 2013

Dai 7330 HQ654094 HQ654108 Cui & Zhao 2013

Cui 2655 HQ654093 HQ848483 Cui & Zhao 2013

P ellipsospora Cui 10276 KFO018124 KF018132 Present study

Cui 10284 KFO18125 KF018133 Present study

P. hainaniana Cui 6364 JQ861743 JQ861759 Zhao & Cui 2013

Cui 6365 JQ861744 JQ861760 Zhao & Cui 2013

Cui 6366 JQ861745 JQ861761 Zhao & Cui 2013

P. koreana KUC 200932 KJ 156313 KJ 156305 Jang & al. 2015

KUC 2008J-02 KJ156310 KJ 156302 Jang & al. 2015

P. lacerata Cui 7220 JX141448 JX141458 Zhao & Cui 2013

Dai 11268 JX141449 JX141459 Zhao & Cui 2013

P. luteola H 1308a JX141456 JX141466 Zhao & Cui 2013

H 1308b JX141457 JX141467 Zhao & Cui 2013

P. maackiae Cui 8929 HQ654102 JF706338 Zhao & Cui 2013

Cui 5605 JN048760 JN048780 Zhao & Cui 2013

P. macropora Zhou 280 JQ861748 JQ861764 Zhao & Cui 2013

P. medulla-panis MUCL 49581 FJ411088 FJ393876 Robledo & al. 2009

MUCL 43250 FJ411087 FJ393875 Robledo & al. 2009

Cui 3274 JN112792 JN112793 Zhao & al. 2013

128 ... Zhao & Ma

P. mopanshanensis

P. nanlingensis

P pyricola

P. rhizomorpha

P. russeimarginata

P. straminea

P. subacida

P. subadusta

P. substraminea

P. subtephropora

P. tenuis

P. tephropora

P. tibetica

P. truncatospora

Perenniporiella chaquenia

Pe. micropora

Pe. neofulva

Pe. pendula

Pyrofomes demidoffii

Truncospora detrita

T. macrospora

T. ochroleuca

T. ohiensis

Vanderbylia “delavayi’

V. fraxinea

V. “robiniophila”

V. vicina

CLZhao 5145

CLZhao 5152

CLZhao 2404 [T]

CLZhao 2311

Cui 7620

Cui 7589

Cui 9149

Dai 10265

Dai 7248

Cui 7507

Yuan 1244

Cui 8718

Cui 8858

Dai 8224

Cui 3643

MUCL 31402

Cui 8459

Cui 10177

Cui 10191

Dai 10962

Dai 10964

Wei 2783

Wei 2969

Cui 9029

Cui 6331

Cui 9459

Cui 6987

Dai 5125

MUCL 47647

MUCL 47648

MUCL 43581

MUCL 45091

MUCL 46034

MUCL 41034

MUCL 42649

Cui 8106

Dai 11486

MUCL 39726

MUCL 39563

Cui 5714

MUCL 41036

Dai 6891

DP 83

Cui 7154

Cui 8885

Cui 8871

Cui 5644

Cui 7144

Cui 9174

MUCL 44779

MH784912

MH784913

MH784911

MH784910

HQ848477

HQ848478

JN048762

JN048761

JF706330

HQ654107

JQ861750

HQ876600

HQ654104

HQ876605

FJ613655

FJ411103

HQ876606

JQo001852

JQ001853

JQ861752

JQ861753

JQ001858

JQ001859

HQ876601

HQ848473

JE706327

JN048778

HQ654098

FJ411083

FJ411084

FJ411086

FJ411080

FJ411082

FJ411105

FJ411099

JX941573

HQ654105

FJ411098

FJ411097

HQ654103

FJ411096

JQ861738

AM269789

HQ654095

HQ876611

JF706329

HQ876609

HQ876608

HQ876610

FJ411095

MH784916

MH784917

MH784915

MH784914

HQ848486

HQ848487

JN048782

JN048781

JF706348

HQ654117

JQ861766

JF706335

JF706334

JF713024

AY336753

AY333796

HQ654113

JQ001844

JQ001845

JQ861768

JQ861769

JQ001848

JQ001849

JF706339

HQ848484

JE706333

HQ654112

HQ848481

FJ393855

FJ393856

FJ393858

FJ393852

FJ393853

FJ393873

FJ393866

JX941596

JF706349

FJ393865

FJ393864

HQ654116

FJ393863

KF495019

AM269853

HQ654110

JF706344

JF706345

JF706342

JF706341

JF706343

AF518666

Present study

Zhao & Cui 2013

Zhao & Cui 2012

Zhao & Cui 2013

Zhao & al. 2013

Robledo & al. 2009

Zhao & al. 2013

Zhao & Cui 2013

Zhao & al. 2013

Zhao & Cui 2013

Robledo & al. 2009

Zhao & al. 2013

Robledo & al. 2009

Zhao & al. 2013

Robledo & al. 2009

Zhao & al. 2013

Robledo & al. 2009

Zhao & al. 2013

Robledo & al. 2009

Perenniporia mopanshanensis sp. nov. (China) ... 129

Sequencher 4.6 (GeneCodes, Ann Arbor, MI, USA) was used to edit the DNA

sequence. Sequences were aligned in MAFFT 7 using the “G-INS-I” strategy (https://

maftt.cbrc.jp/alignment/server/index.html) and manually adjusted in BioEdit (Hall

1999). The sequence alignment was deposited in TreeBase (submission ID 22900).

Donkioporia expansa (Desm.) Kotl. & Pouzar and Pyrofomes demidoffii (Lév.) Kotl. &

Pouzar were used as outgroup to root trees following Zhao & al. (2013) in the ITS+nLSU

analyses.

ITS+nLSU sequence analyses were performed using maximum parsimony,

maximum likelihood, and Bayesian inference methods. Maximum parsimony (MP)

analyses followed Song & al. (2016), and tree construction was performed in PAUP*

version 4.0b10 (Swofford 2002). All characters were equally weighted, with gaps treated

as missing data. Trees were inferred using the heuristic search option with TBR branch

swapping and 1000 random sequence additions. Max-trees was set to 5000, branches

of zero length were collapsed, and all parsimonious trees were saved. Clade robustness

was assessed using bootstrap (BT) analysis with 1000 replicates (Felsenstein 1985). Tree

length (TL), consistency index (CI), retention index (RI), rescaled consistency index

(RC), and homoplasy index (HI) were calculated for each Maximum Parsimonious Tree

(MPT) generated. Sequences were analyzed using Maximum Likelihood (ML) with

RAXxML-HPC2 through the Cipres Science Gateway (www.phylo.org; Miller & al. 2009).

Branch support (BS) for ML analysis was determined by 1000 bootstrap replicates.

MrModeltest 2.3 (Posada & Crandall 1998, Nylander 2004) was used to determine

the best-fit evolution model for each data set for Bayesian inference (BI), which

was calculated with MrBayes_3.1.2 using a general time reversible (GTR) model of

DNA substitution and a gamma distribution rate variation across sites (Ronquist &

Huelsenbeck 2003). Four Markov chains were run for 2 runs from random starting trees

for 7 million generations (ITS+nLSU), and trees were sampled every 100 generations.

The first 25% of the generations was discarded as burn-in. A majority rule consensus

tree of all remaining trees was calculated. Branches that received bootstrap support

for maximum likelihood (BS), maximum parsimony (BT), and Bayesian posterior

probabilities (BPP) greater than or equal to 75% (BS, BT) and 0.95 (BPP) were considered

significantly supported.

Molecular phylogeny

The ITS+nLSU dataset included sequences from 86 fungal specimens

representing 46 species (TABLE 1). The dataset had an aligned length of 2100

characters, of which 1553 characters were constant, 86 variable and parsimony-

uninformative, and 461 parsimony-informative. Maximum parsimony analysis

yielded four equally parsimonious trees (TL = 1925, CI = 0.375, HI = 0.525,

RI = 0.745, RC = 0.275). Best model for the ITS+nLSU dataset estimated and

applied in the Bayesian analysis: GIR+1+G. Bayesian analysis and ML analysis

produced similar topologies to MP analysis, with an average standard deviation

of split frequencies = 0.006258 (BI).

130 ... Zhao & Ma

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132 ... Zhao & Ma

The ITS+nLSU phylogeny places Perenniporia mopanshanensis in the

Perenniporia sensu stricto clade (Fic. 1), where it forms a strongly supported

monophyletic lineage (BS = 100%; BT = 100%; BPP = 1) and is sister to

P. bannaensis B.K. Cui & C.L. Zhao.

Taxonomy

Perenniporia mopanshanensis C.L. Zhao, sp. nov. Fics 2, 3

MycoBank MB 827580

Differs from Perenniporia luteola by its larger pores, its smaller basidiospores, and its

unbranched skeletal hyphae.

Type: China. Yunnan Province: Yuxi, Xinping county, Mopanshan National Forestry

Park, on an angiosperm trunk, 19 August 2017, C.L. Zhao 2404 (Holotype, SWFC

002404; GenBank MH784911, MH784915).

ErymMoLocy: The specific epithet mopanshanensis (Lat.) refers to the locality

(Mopanshan) of the type specimen.

BASIDIOMATA perennial, resupinate, without odor or taste when fresh,

becoming corky upon drying, <15 x 6 cm, 8 mm thick at center. Pore surface

cream when fresh, cream to buff to straw upon drying; pores round, 3-5 per

mm; dissepiments thin, entire. Sterile margin narrow, cream, <0.5 mm wide.

Subiculum cream, thin, each layer <0.5 mm thick. Tubes cream to buff, corky,

each layer <2 mm long.

HYPHAL STRUCTURE dimitic; generative hyphae with clamp connections;

skeletal hyphae strong dextrinoid, CB+; tissues unchanged in KOH.

SUBICULUM generative hyphae infrequent, hyaline, thin-walled, unbranched,

2-3 um in diam.; skeletal hyphae dominant, hyaline, thick-walled with a narrow

to wide lumen, unbranched, interwoven, 2-3.5 um in diam.

TUBE generative hyphae infrequent, hyaline, thin-walled, unbranched,

1.5-3 um in diam.; skeletal hyphae dominant, hyaline, thick-walled with a

narrow to wide lumen, unbranched, interwoven, 2-3.5 um. Plenty of crystals

present among hyphae. Cystidia absent, fusoid cystidioles present, hyaline,

thin-walled, 13-19 x 4.5-6.5 um; basidia barrel-shaped to clavate, with

four sterigmata and a basal clamp connection, 14-20 x 7-12 um; basidioles

dominant, mostly pear-shaped, but slightly smaller than basidia.

Basipiospores ellipsoid, non-truncate, hyaline, distinctly thick-walled,

smooth, strong dextrinoid, CB+, (5—)5.5-6.5(-7) x 4-5(-5.5) um, L = 6.15 um,

W = 4.6 um, Q = 1.35-1.42 (n = 120/4).

TYPE OF ROT: white rot.

ADDITIONAL SPECIMENS EXAMINED: CHINA. YUNNAN PROVINCE. Yuxi: Xinping

county, Mopanshan National Forestry Park, on the angiosperm trunk, 19 August 2017,

Perenniporia mopanshanensis sp. nov. (China) ... 133

Fic. 2. Perenniporia mopanshanensis (holotype, SWFC 002404).

Basidiomata. Scale bar: 3 cm.

C.L. Zhao 2311 (SWFC 002311; GenBank MH784910, MH784914); 13 January 2018,

C.L. Zhao 5145 (SWFC 005145; GenBank MH784912, MH784916); C.L. Zhao 5152

(SWFC 005152; GenBank MH784913, MH784917).

Discussion

The new species, Perenniporia mopanshanensis, is supported by

phylogenetic analyses and morphological characters. In the ITS+nLSU

analyses (Fic. 1), it forms a strongly supported monophyletic lineage (BS

= 100%; BT = 100%; BPP = 1) where it forms a clade with P. bannaensis.

However, morphologically P bannaensis differs from P. mopanshanensis by its

annual basidiocarps with buff-yellow to pinkish buff pore surface and smaller

pores (6-8 per mm; Zhao & al. 2013).

Morphologically, the presence of non-truncate basidiospores is shared

by several other species in Perenniporia sensu stricto: P africana Ipulet &

Ryvarden, P. ellipsospora Ryvarden & Gilb., P koreana Y. Jang & J.J. Kim,

P. luteola B.K. Cui & C.L. Zhao, P. rhizomorpha B.K. Cui & al., and P. subacida

(Peck) Donk. A morphological comparison between P. mopanshanensis and

these six species is presented in TABLE 2.

134 ... Zhao & Ma

Fic. 3. Perenniporia mopanshanensis (holotype, SWFC 002404). A. Basidiospores; B. Section of

hymenium; C. Hyphae from trama; D. Hyphae from subiculum; E. Rhombic crystal. Scale bars:

a=5 um; b-e = 10 um.

Polypores are an extensively studied group in Basidiomycota (Gilbertson &

Ryvarden 1987, Nufiez & Ryvarden 2001, Ryvarden & Melo 2014), but Chinese

polypore diversity is still not well known, especially in the subtropics and tropics.

The new species, Perenniporia mopanshanensis, was found in the Chinese

subtropics, where many new taxa in the Polyporales and Hymenochaetales have

Perenniporia mopanshanensis sp. nov. (China) ... 135

TABLE 2. A comparison of Perenniporia species with non-truncate basidiospores

PORE PORES SPORES SKELETAL

SPECIES HABIT REFERENCE

SURFACE /mm (um) HYPHAE

africana Annual Pale orange 6-8 4-5 x Branched Ipulet & Ryvarden

to brown 3-4 2005

ellipsospora Annual Whitish 3-4 4-5.5 x Unbranched Gilbertson &

to pale 3-4 Ryvarden 1987

yellowish

brown

koreana Annual Grayish 5-6 6-7 x Rarely Jang & al. 2015

orange 3.9-5.2 branched

luteola Perennial _Buff- 4-6 6-7 x Frequently Zhao & Cui 2013

yellow 5-5.5 branched

mopanshanensis Perennial Cream,buff 3-5 5.5-6.5 x | Unbranched This study

to straw 4-5

rhizomorpha Annual Yellow- 4-6 5.3-6.5 x Branched Cui & al. 2007

buff to 4,1-5.2

yellowish

orange

subacida Perennial Ivory to 5-6 4.5-6 x Unbranched Decock & Stalpers

yellowish 3.5-4.5 2006

been described (Cui & Dai 2008; Cui & al. 2009, 2010, 2011; Du & Cui 2009; Li

& Cui 2010; He & Li 2011; Jia & Cui 2011; Yu & al. 2013; Yang & He 2014; Chen

& al. 2015). We anticipate that additional polypore taxa will be found in China

after further investigation and molecular analysis.

Acknowledgments

Special thanks are due to Drs. Karen K. Nakasone (Center for Forest Mycology

Research, Northern Research Station, U.S. Forest Service, USA) and Jie Song (Research

Institute of Subtropical Forestry, Chinese Academy of Forestry, P.R. China) who

reviewed the manuscript. The research is supported by the National Natural Science

Foundation of China (Project No. 31700023), Yunnan Agricultural Foundation

Projects (2017FG001-042), the Science Foundation of Southwest Forestry University

(Project No. 111715), and the Science Foundation of Education Department in

Yunnan (2018]S326)

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Zhao CL, Cui BK. 2013. Three new Perenniporia (Polyporales, Basidiomycota) species

from China based on morphological and molecular data. Mycoscience 54: 231-240.

https://doi.org/10.1016/j.myc.2012.09.013

Zhao CL, Cui BK, Dai YC. 2013. New species and phylogeny of Perenniporia

based on morphological and molecular characters. Fungal Diversity 58: 47-60.

https://doi.org/10.1007/s13225-012-0177-6

MY COTAXON

January-March 2019—Volume 134, pp. 139-146

https://doi.org/10.5248/134.139

New records of Bilimbia and Toninia from China

MEI-JIE SUN ', SHU-KUN YAN ', RONG TANG +,

CHUN-XIAO WANG *, LU-LU ZHANG

'Key Laboratory of Plant Stress Research, College of Life Sciences,

Shandong Normal University, Jinan, 250014, PR. China.

“ CORRESPONDENCE TO: 675359138@qq.com

ABSTRACT—As a result of our study on the lichen flora of Northwest China, one species of

Bilimbia (B. lobulata) and three species of Toninia (T. coelestina, T. gobica, and T: superioris)

are reported for the first time from China.

Keyworps—Asia, biodiversity, Lecanorales, Ramalinaceae, taxonomy

Introduction

Bilimbia De Not. and Toninia A. Massal. belong to Ramalinaceae C. Agardh

(Lecanorales), a widely distributed lichen family (Ekman 2001, Ekman & al.

2008).

Bilimbia is distinguished from Bacidia, Biatora, and Mycobilimbia by the

stout paraphyses and ascospores often surrounded by a finely warted perispore

(Smith & al. 2009, Veldkamp 2004.).

Toninia was described by Massalongo (1852) originally based on ascospore

morphology as well as on characters of apothecia and thallus. Today it is

additionally diagnosed by the asci and paraphyses. The asci are Bacidia-type,

clavate. The paraphyses are usually straight and rarely branched (except at

the tips), with apical cells that are distinctly swollen and usually capped with

gelatinous pigment. Epithecial pigments that also characterize Toninia include

grey (K+ violet, N+ violet), bright green (K-, N+ violet), olivaceous green

(K+ violet/brown, N—), reddish brown (K+ red, N-), and dull brown (K-, N-)

(Timdal 1991, 2002; Zahlbruckner 1890).

140 ... Sun & al.

Here we report one species of Bilimbia (B. lobulata) and three species

of Toninia (T. coelestina, T. gobica, and T: superioris) for the first time from

China.

Materials & methods

The specimens studied are deposited in the Lichen Section, Botanical Herbarium,

Shandong Normal University, Jinan, China (SDNU), and the Mycological

Herbarium, Institute of Microbiology, Chinese Academy of Sciences, Beijing, China

(HMAS-L). The specimens were examined morphologically under an Olympus SZ

stereo-microscope and Olympus CX21 polarizing microscope. Thalli and medullae

were tested for identification with K (potassium hydroxide, 10% aqueous solution),

C(sodium hypochlorite, saturated aqueous solution), I (potassium iodide, 10% aqueous

solution), N (10% nitric acid), and P (saturated solution of p-phenylenediamine in

95% ethyl alcohol). Lichen substances were identified using standardized thin layer

chromatography (TLC) techniques with system C (Orange & al. 2001). The lichens

were digitally photographed under Olympus SZX16 and BX61 with DP72.

Taxonomic descriptions

Bilimbia lobulata (Sommerf.) Hafellner & Coppins,

Lichenologist 36: 195 (2004) Fic. 1

THALLUS warted-granular to squamulose, squamules <0.6 mm wide,

continuous, toverlapping, yellowish white; APOTHECIA <1 mm diam.,

lecideine, black, sessile, flat to slightly convex, epruinose to faintly pruinose;

PROPER EXCIPLE reddish brown (similar to hypothecium), K+ intensifying

or reddish purple; EPIHYMENIUM <15 um high, olive green, K-, N+ reddish;

HYMENIUM 70-85 um high, colourless to dull reddish brown; SuUBHYMENIUM

25-40 um high, usually colourless; PARAPHYSEsS conglutinated, simple or

slightly branched, 2-3.5 um diam. with apical cell distinctly swollen <5 um

diam.; ASCOSPORES medium fusiform, (0—)2-3-septate, 15-20 x 3-5 um.

CHEMISTRY—Cortex and medulla K-, C-, KC-; Zeorin detected by

LG.

SPECIMENS EXAMINED: CHINA. QINGHAI: Qilian, Niuxinshan, alt. 3500 m, on

soil, 11 Aug. 2007, 11 Aug. 2007, J.G. Liu 20071633 (SDNU); SHANx!: Niingwu,

Luyashan, alt. 2500 m, on soil, 26 Aug. 2011, H.Y. Wang 20121675 (SDNU).

DIsTRIBUTION—Europe, North America, Antarctica, Asia (Smith & al.

2009, Urbanavichus 2010). New to China.

ComMMENTS—The Chinese material closely matches the previously

published description by Hafellner & al. (2004). Bilimbia lobulata is

First reports of Bilimbia & Toninia spp. in China... 141

$ Fg 5.

Fic. 1. Bilimbia lobulata (SDNU 20071633). A. Thallus and apothecia; B. Apothecium section;

C. Apothecium section (showing epihymenium K+ reaction); D. Ascospores with three septa.

Scale bars: A = 1 mm; B, C = 50 um; D = 10 um.

distinguished from B. sabuletorum (Schreb.) Arnold, which has a crustose

thallus, and longer, 3—7(—9)-septate ascospores (Smith & al. 2009).

Toninia coelestina (Anzi) Vézda,

Cas. Slez. Mus., Ser. A, Hist. Nat. 10: 105 (1961) Fic. 2

THALLUS crustose to subsquamulose, continuous to warty, granules

present; upper side pale yellowish brown to greyish brown, epruinose, without

pores and pseudocyphellae; APoTHECIA 0.75-1.5 mm diam., epruinose,

when young flat and mostly marginate, when mature mostly weakly convex

and immarginate; PROPER EXCIPLE grayish black on the rim, dark brown in

inner part, K-, N+ violet; HypoTHEcIum dark brown; HyMENIuM 50-65 um

high; EprrHecium dark olive green, K-, N+ violet; PARAPHYSES simple or

slightly branched at the tip, 2-3.5 um diam. with apical cell swelling <4.5 um

diam.; AscosPoRES narrowly ellipsoid to bacilliform, 3-7-septate, 17.5-37.5

x 2.5-3.5 um.

142 ... Sun & al.

Fig. 2. Toninia coelestina (SDNU 20126519a). A. Thallus with apothecia; B. Apothecium

section; C. Asci: Bacidia-type; D. Ascospore with three septa. Scale bars: A = 1 mm; B = 50 um;

C, D= 10 um.

CHEMISTRY—Cortex and medulla K-, C-, KC-. Terpenoids detected by

EEG.

SPECIMEN EXAMINED: CHINA. XINJIANG: Tianshan, Yihaobingchuan, alt. 2480 m, on

soil, 26 Aug. 2011, L. Li 20126519a (SDNU).

DISTRIBUTION—Europe: Austria, Italy, Norway, Poland, Scotland, Slovakia

(Flakus 2007, Timdal 1991, van den Boom & al. 1996, Vézda A. 1961). New to

China.

ComMENTS— The Chinese material closely matches previously the published

description by Vézda (1961). Toninia coelestina closely resembles T: thiopsora

(Nyl.) H. Olivier, which differs by its 7-septate ascospores and its C+ orange

thallus (Smith & al. 2009).

Toninia gobica N.S. Golubk., Bot. Zhurn. SSSR 58(6): 838 (1973) Fic. 3

THALLUS squamulose, squamules <3 mm diam., scattered to contiguous,

slightly to strongly convex, weakly pruinose, dull yellow, without pseudo-

First reports of Bilimbia & Toninia spp. in China... 143

Fic. 3. Toninia gobica (HMAS-L 121615). A. Thalli with apothecia; B. Apothecium section;

C. Ascospore with three septa. Scale bars: A = 2 mm; B = 20 um; C = 10 um.

cyphellae, mostly rimose on the surface; APOTHECIA <1 mm diam., flat

to strongly convex, usually on the edge of thallus, epruinose to slightly

pruinose; PROPER EXCIPLE dark brown in inner part, K-, N-; HyPOTHECIUM

dull yellowish brown or colourless; HyMENIUM 60-70 um high; epithecium

bright green, K-, N+ violet; Ascosporss acicular, 3-septate, 30-45 x 3-5 um.

CHEMISTRY—Cortex and medulla K-, C-, KC-. Terpenoids detected by

TLC:

144 ... Sun & al.

SPECIMENS EXAMINED: CHINA. NinGx1A: Zhongwei, Shapotou, Cuiliugou, alt. 1200

m, on soil, 9 Apr. 2010, T. Zhang 121615 (HMAS-L); NEIMENGGuU: Wulatehouqi,

Huhebashige, alt. 1600 m, on rock, 19 Aug. 2011, H.Y. Wang 20122753 (SDNU).

DISTRIBUTION—Mongolia (Golubkova 1973, Timdal 1991). New to China.

COMMENTS— The Chinese material closely matches the type description by

Golubk (1973). Toninia gobica and T. lutosa (Ach.) Timdal both have regular

fissures on the thallus, but T. lutosa has deeper fissures and also differs by

its dark reddish brown epithecium and shorter, K+ red, N-, (1-)3-septate

ascospores (Timdal 1991).

Toninia superioris Timdal, Opera Bot. 110: 103 (1991) Fig. 4

= Kiliasia superioris (Timdal) Timdal, in Kistenich & al., Taxon 67(5): 893 (2018)

THALLUS squamulose, squamules <1 mm diam., imbricate or contiguous,

orbicular to irregular, flat to slightly convex, pruina present or absent,

whitish grey, fissures and pseudocyphellae absent; APOTHECIA <lmm

diam., flat to slightly convex, marginate or immarginate, faintly pruinose;

PROPER EXCIPLE grey on the rim, paler grey to colourless in the inner part,

K+ violet, N+ violet; HyPoTHECIUM colourless to pale brown; EPITHECIUM

grey, K+ violet, N+ violet; Ascosporgs narrowly or broadly fusiform, mainly

3-septate, 12.5-20 x 3.5—-5 um.

CHEMISTRY—Cortex and medulla K-, C-, KC-. Terpenoids detected by

TG:

SPECIMEN EXAMINED: CHINA. NEIMENGGU: Keqi, Huanggangliang, alt. 2000 m,

on soil, 13 Aug. 2011, H.Y. Wang 20117397 (SDNU).

DISTRIBUTION—U.S.A.: Michigan (Timdal 1991). New to China.

ComMENTS— the Chinese material closely matches the type description by

Timdal (1991). Toninia superioris resembles T: hosseusiana Gyeln., which

differs by its dark reddish brown epithecium and proper exciple (K+ red,

N-), and its <3 mm diam. thalline squamules covered with farinose pruina

(Gyelnik 1942, Timdal 1991).

Acknowledgements

The authors thank Dr Edit Farkas (MTA Centre for Ecological Research, Vacratot,

Hungary) and Dr Shou-Yu Guo (State Key Laboratory of Mycology, Institute of

Microbiology, Chinese Academy of Sciences, Beijing, China) for presubmission

reviews. This work was supported by the Emergency management project of National

Natural Science Foundation of China (31750001), the National Natural Science

Foundation of China (31400015, 31570017, 31600100), and the Scientific Research

Foundation of Graduate School of Shandong Normal University (SCX201629).

First reports of Bilimbia & Toninia spp. in China... 145

Fic. 4. Toninia superioris (SDNU 20117397). A. Thallus and apothecia; B. Apothecium section;

C. Apothecium section (showing epihymenium K+ reaction); D, E. Ascospores with three septa.

Scale bars: A = 500 um; B, C = 20 um; D, E= 10 um.

Literature cited

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Mycological Research 105(7): 783-797. https://doi.org/10.1017/s0953756201004269

Ekman S, Andersen HL, Wedin M. 2008. The limitations of ancestral state reconstruction and

the evolution of the ascus in the Lecanorales (lichenized Ascomycota). Systematic Biology

57(1): 141-156. https://doi.org/10.1080/10635150801910451

Flakus A. 2007. Lichenized and lichenicolous fungi from mylonitized areas of the subnival belt

in the Tatra Mountains (Western Carpathians). Annales Botanici Fennici 44: 427-449.

Golubkova NS. 1973. New species of the genus Toninia Mass., found in the Mongolian People’s

Republic. Botanicheskii Zhurnal SSSR. 58: 838-840.

Gyelnik V. 1942. Species tres novae ex Argentina. Annales Historico-Naturales Musei Nationalis

Hungarici 35: 98-100.

Orange A, James PW, White FJ. 2001. Microchemical methods for the identification of lichens.

2nd edition. London, British Lichen Society.

Smith CW, Aptroot A, Coppins BJ, Fletcher A, Gilbert OL, James PW, Wolseley PA. 2009. The

lichens of Great Britain and Ireland. 2nd edition. The British Lichen Society, Department of

Botany, The Natural History Museum, London. 1042 p.

Timdal E. 1991. A monograph of the genus Toninia (Lecideaceae, Ascomycetes). Opera Botanica

110. 137 p.

146 ... Sun & al.

Timdal E. 2002. Toninia. 488-501, in: TH Nash & al. (eds). Lichen Flora of the Greater Sonoran

Desert Region. Vol. 1. Lichens Unlimited, Tempe.

Urbanavichus GB (ed.) 2010. A checklist of the lichen flora of Russia. Nauka, St. Petersburg,

194 p. ISBN 978-5-02-025422-0.

Van den Boom PPG, Breuss O, Spier L, Brand AM. 1996. Beitrag zur Flechtenflora Karntens

Ergebnisse der Feldtagung der Bryologischen und Lichenologischen Arbeitsgruppe der

KNNV in Weissbriach 1994. Linzer biologische Beitrage 28(2): 619-654.

Veldkamp JE 2004. Bilimbia (Lichenes) resurrected. Lichenologist. 36(3-4): 191-195.

https://doi.org/10.1017/s0024282904013908

Vézda A. 1961. Systematische Studien tiber die Flechtengattung Bacidia DNot. emend. Zahlbr.

I. Neue oder wenig bekannte Arten. Casopis Slezského Musea, Ser. A, Historia Naturalis 10:

103-113.

Zahlbruckner A. 1890. Prodromus einer Flechtenflora Bosniens. Annalen des Kaiserlich-

K6niglichen Naturhistorischen Hofmuseums, Wien 5: 20-48.

MY COTAXON

January-March 2019—Volume 134, pp. 147-153

https://doi.org/10.5248/134.147

Anamylopsora altaica sp. nov. from Northwestern China

PARIDA AHAT', ANWAR TUMUR”™ , SHOU- YU GUO?”

" Arid land Lichen Research Center of Western China,

College of Life Science and Technology, Xinjiang University,

Urumgdi 830046, P. R. China

’ State Key Laboratory of Mycology, Institute of Microbiology,

Chinese Academy of Sciences, Beijing 100101, P. R. China

* CORRESPONDENCE TO: *anwartumursk@xju.edu.cn ® guosy@im.ac.cn

ABSsTRACT—Anamylopsora altaica, collected from rocks in the Hot Spring Valley Forest Park

in the Altay Mountains, Xinjiang, Northwestern China, is described as a new species, based

on morphological and chemical characteristics, as well as ITS DNA sequence data. The new

lichen species is characterized by its crowded, overlapping squamules with upturned margins,

crowded globose apothecia with eight simple hyaline spores per ascus, and the presence of

psoromic acid. A detailed description and colour photographs are provided.

Key worps—Anamylopsoraceae, Baeomycetaceae, Baeomycetales

Introduction

The lichen biota of Northwestern China is rich, with more than 750 species

belonging to c. 150 genera (Wei 1991, Abbas & Wu 1998, Abbas & al. 2001,

Guo 2005). One lichen from temperate and tropical regions, Anamylopsora

pulcherrima (Vain.) Timdal, type of the monotypic genus Anamylopsora Timdal

(Huneck & Elix 1993, Lumbsch & al. 1995), has been collected in China from

Gansu (Magnusson 1940, Timdal 1991, Wei 1991), Neimenggu (Magnusson

1942, Timdal 1991, Wei 1991) and Xizang (Obermayer 2004). Anamylopsora

pulcherrima was originally described as Lecidea pulcherrima Vain. (Vainio

1888), subsequently combined as Psora pulcherrima (Vain.) Elenkin (Elenkin

1904), and finally transferred to Anamylopsora by Timdal (1991).

A comparison of anatomical, chemical, and ontogenetical characters

initially supported Anamylopsora within a new family Anamylopsoraceae

148 ... Ahat, Tumur, Guo

Lumbsch & Lunke (Agyriineae, Lecanorales; Lumbsch & al. 1995), but more

recent phylogenetic analyses place the genus in Baeomycetaceae, Baeomycetales.

Anamylopsora specimens collected recently from northwestern China have

been examined and are described here as a new species, Anamylopsora altaica.

Materials & methods

Specimens of the lichen Anamylopsora were collected by from rocks in the

Hot Spring Valley Forest Park and in the Two-River Source Nature Reserve, Altay

Mountains, Xinjiang, Northwestern China, and have been deposited in the Lichen

Section of the Botanical Herbarium, Arid land Lichen Research Center of Western

China, Xinjiang University, Urumqi, China (XJU).

Morphology

The specimens were examined using a Nikon Eclipse E200 stereo-microscope.

Sections were cut by hand using a razor blade and were mounted and observed in

water and iodine. The structure and hymenial characters were studied with a Zeiss

Axioskop 2 plus light microscope and photographed using a Nikon Digital Camera

D50. Chemical constituents were identified by colour spot tests and thin-layer

chromatography using solvent system C (Orange & al. 2010).

DNA extraction, amplification, sequencing

Thallus fragments with ascomata were removed from the holotype and two other

specimens for DNA extraction using the DNAsecure Plant DNA Kit following the

manufacturer’s protocol. The nrITS (ITS1+5.8S+ITS2) region was amplified in a

polymerase chain reaction according to Han & al. (2013) as modified in Abbas & al.

(2014) using the primers ITS1F (Gardes & Bruns 1993) with ITS4 (White & al. 1990).

The DNA was amplified in a 30 uL volume containing 0.75 units of TransStart Taq

Polymerase, 3.5 uL buffer, 0.5 ul of a 5 uM primer solution, 2 uL each per 2.5 mM

dNTP solution, and 1 wL of genomic DNA. The PCR mixture was cycled at 95°C for

5 min; then 35 cycles at 94°C for 30 s, 55°C for 45 s, and 72°C for 1 min; with a

final extension of 72°C for 10 min. PCR products were screened on 1% agarose gels

stained with ethidium bromide and sequenced by Shenggong Inc. in Shanghai. Newly

obtained sequences were submitted to GenBank.

We aligned ITS sequences from our three specimens and the 17 GenBank

representatives using ClustalW and Muscle implemented in MEGA7 (Kumar &

al. 2016). The final matrix, submitted to TreeBase, is available by request from the

corresponding authors.

The evolutionary history was inferred by using the Neighbour-joining (NJ) method

implemented in MEGA7 (Saitou & Nei 1987, Tamura & al. 2004, Kumar & al. 2016).

Bootstrap replication was set to 1000, and the evolutionary distances were computed

using the Maximum Composite Likelihood method and were in units of the number

of base substitutions per site. Rate variation among sites was modeled with a gamma

distribution (shape parameter = 5). The Minimum Evolution (ME) tree was searched

Anamylopsora altaica sp. nov. (China) ... 149

99 | AF274085 Trapeliopsis flexuosa

AF274086 Trapeliopsis flexuosa

NR119922 Trapeliopsis flexuosa

AY600082 Trapeliopsis glaucolepidea

AF274087 Trapeliopsis granulosa

AF274088 Trapeliopsis pseudogranulosa Trapeliales

KY797784 Trapelia placodioides

AF274080 Trapelia involuta

AF274081 Trapelia placodioides

AF274083 Placopsis subparellina

AF274084 Placopsis gelida

AF274091 Placopsis gelida

AF274077 Anzina carneonivea

KT601494 Baeomyces rufus

KT601493 Baeomyces placophyllus

100) KRO17064 Anamylopsora pulcherrima from Russia

AF274089 Anamylopsora pulcherrima from Russia

MH513961 Anamylopsora altaica from China

MH513962 Anamylopsora altaica from China

MH513963 Anamylopsora altaica from China

Baeomycetales

0.02

Fic.1 Neighbour-joining (NJ) phylogenetic tree of ITS sequences for Anamylopsora altaica and

similar species retrieved from GenBank. Numbers above branches indicate values >50% from 1000

bootstrap replicates of NJ analyses.

using the Close-Neighbor-Interchange (CNI) algorithm at a search level of 1. The

initial tree was generated by the Neighbor-joining algorithm. The gaps were removed

for each sequence pair.

ITS sequence analysis and phylogeny

The entire ITS region was successfully sequenced for the holotype specimen

and two additional specimens. A Blast search was conducted, and GenBank

sequences from taxa sharing the most similarity with our new sequences were

selected (Timdal 1991, Lumbsch & al. 1995). Partial sequences of the 3’ end in

SSU region and the 5’ end in LSU region were included in the data submitted

to GenBank.

The final nrITS alignment included 20 sequences representing 13 taxa

and comprised 513 sites, of which 441 were constant, 18 were parsimony-

uninformative, and 54 were parsimony-informative. There were no ambiguously

aligned regions.

ITS sequences support our specimens in Anamylopsora with close affinities

to the existing A. pulcherrima sequences in GenBank (91% with KR017064;

90% with AF274089). The evolutionary history was inferred as the NJ tree. The

150 ... Ahat, Tumur, Guo

phylogeny clearly shows the relationship of the new species with closely related

species (Fic.1). Anamylopsora and Baeomyces cluster together in a monophyletic

group within Baeomycetales, which is a sister group to the Trapeliales.

Taxonomy

Anamylopsora altaica Ahat, A. Abbas, S.Y. Guo & Tumur, sp. nov. FIG. 2

FUNGALNAME FN570571

Differs from Anamylopsora pulcherrima by its overlapping squamules, its crowded

globulose apothecia concentrated along the squamule margins, and the presence of

psoromic acid and absence of alectorialic acid.

Type: China. Xinjiang: Altay Mountains, Hot Spring Valley Forest Park, 47°33’52”N

88°41'29’E, alt. 1030 m, 17 May 2014, A. Tumur & A. Abbas Altay-20140041 (Holotype,

XJU; GenBank MH513961; Isotype, HMAS-L).

EryMo ocy: The new species is named after the Altay Mountains, where the holotype

was collected.

THALLUuS irregularly squamulose, usually appressed on siliceous rocks;

SQUAMULES <1(-2.5) mm diam., intensively overlapping, without soredia

or isidia, pale-green to light yellow-green, with or without a light frosting

of pruina on the upper surface; thallus margin entire to subentire, usually

upturned; UPPER SURFACE white to whitish-grey pruinose; LOWER SURFACE

white to dirty white, without rhizines; UPPER CORTEX 55-95 um thick,

brown to light-brown; ALGAL LAYER 135-195 um thick, continuous, algal

cells 8-12 um diam., photobiont Trebouxia sp.; MEDULLA 190-280 um thick;

LOWER CORTEX paraplectenchymatous, 30-65 um thick.

APOTHECIA lecideine, <1(-1.5) mm diam.; often >10 globose apothecia

in clusters <12 mm diam. attached to the squamule margins; pisc brown

to dark brown, dull, epruinose; EPIHYMENIUM faint yellow to light brown,

25-30 um high; HYMENIUM 95-115 um high. PaRApHysEs simple or sparingly

branched. HypoTHALtus colourless, 70-115 um high. Ascr narrowly clavate

to subcylindrical, lacking an ocular chamber, inamyloid, 8-spored, 50-70 x

12-16 um. Ascospore hyaline, simple, ellipsoid, thin-walled, 9-12 x 5-8

um, usually with 1-3 oil drop(s) and a warty surface. PycNipIA occurring

at the squamule margins, rod-shaped, <0.4 mm diam., dark brown to black.

Conrpia hyaline narrowly ellipsoid to shortly bacilliform, 3 x 1 um.

CoLor TEests—Thallus cortex PD+ orange, K+ yellow, C-, KC-, UV-.

Thallus medulla PD-, K-, C-, KC-. Apothecial disks I+ blue, K-, C-, UV-.

SECONDARY METABOLITES—Only psoromic acid detected by TLC.

Anamylopsora altaica sp. nov. (China) ... 151

Fic. 2 Anamylopsora altaica (Holotype, XJU Altay-20140041). A. General habit; B. Apothecia;

C. Interrupted algal layer in thin section of the thallus; D. Section of apothecium; E. Ascus and

spores; F. Conidia. Scale bars: A = 10 mm; B= 1 mm; C = 200 um; D = 50 um; E = 20 um; F=5 um.

EcoLocy—Anamylopsora altaica grows at altitudes of c. 900-1200 m in

montane forests, on semi-shaded granite rocks, in association with Aspicilia

concorta, and Acarospora sp. ‘The species is known only from Xinjiang,

Northwestern China; although rare in the type locality, it may occur in other

regions in central Asia.

ADDITIONAL SPECIMENS EXAMINED—CHINA. XINJIANG: Altay Mountains, Hot Spring

Valley Forest Park, 47°50’23”N 88°42’51’E, alt. 960 m, 26 May 2014, Anwar Tumur &

Abdulla Abbas 20140032 (XJU-NALH, GenBank MH513962); 47°03’41”N 88°67'06”E,

alt. 1022 m, Anwar Tumur & Abdulla Abbas 20140040 (XJU-NALH); Two-River Source

Nature Reserve, 47°45’09”N 88°34’48”E, alt. 1187 m, 20 July 2015, Anwar Tumur &

Abdulla Abbas 20150068 (XJU-NALH; GenBank MH513963).

ComMMENTS—Anamylopsora altaica and A. pulcherrima are distinguished

by thallus color, apothecial characters, secondary substances, and substrate.

Anamylopsora pulcherrima usually has yellow lobes and black apothecia, with

squamules having a strongly fissured upper side, a purplish-red lustrous thallus

with a lacinulate appearance, and produces alectorialic acid; additionally,

A. pulcherrima grows at higher altitudes (c. 2700-3000 m), on calciferous stone

or calcareous soil in association with Phycia kansuensis, Psorotichia nigra, and

several other small lichens (Timdal 1991, Lumbsch & al.1995). Lecidea hedinii

152... Ahat, Tumur, Guo

and L. undulata, which were synonymised with Anamylopsora pulcherrima

by Timdal (1991), also resemble A. altaica, but differ in their ‘flavescenti-

testaceus thallus colour and areolate shape, their K+ (orange) cortex, and their

infrequent, smaller (0.3-0.5 mm), dirty reddish apothecia (Magnusson 1942).

Anamylopsora and Baeomyces both exhibit a similar ascoma ontogeny

and the same type of conidiophores, although these similarities may well

result from convergent evolution (Lumbsch & Huhndorf 2010). However,

although closely related by molecular phylogeny, they differ in the ascus type.

Anamylopsora altaica resembles some species of Trapeliopsis and Trapelia, e.g.,

Trapelia placodioides, but has slightly smaller lobes [<3 mm] and a smooth

upper surface.

Acknowledgements

The authors thank Prof. David H.S. Richardson (Environmental Science, Saint

Mary’s University, Canada), Dr. Troy McMullin (Canadian Museum of Nature),

Prof. Liu-Fu Han (Hebei Normal University, Shijiazhuang 050024, China) and

Dr. Jan Vondrak (Institute of Botany, Academy of Sciences, Czech Republic) for

the pre-submission reviews. We cordially thank Dr. Einar Timdal and Dr. Thorsten

Lumbsch for many additional helpful suggestions and comments. We thank

Greg Baker (Maritime Provinces Spatial Analysis Research Center, Saint Mary’s

University, Halifax, NS, Canada) for helping to improve the resolution of

Figures 1 and 2. A. Tumur is grateful for the Natural Science Foundation of China

grant (No. 31660009).

Literature cited

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Urumdi.

Abbas A, Mijet H, Tumur A, Wu JN. 2001. A checklist of the lichens of Xinjiang, China.

Harvard Papers in Botany 5: 359-370.

Abbas A, Guo SY, Ababaikeli G, Abdulla A, Xahidin H. 2014. Diploschistes xinjiangensis, a new

saxicolous lichen from northwest China. Mycotaxon 129: 465-471.

https://doi.org/10.5248/129.465

Elenkin AA. 1904. Schedae zu Lichenes Florae Rossiae, Fasc.[I-IV. Acta Horti Petropolitani

24: 1-118.

Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes—

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Anamylopsora altaica sp. nov. (China) ... 153

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2009. Fieldiana Life and Earth Sciences 16(7): 1-65. https://doi.org/10.3158/1557.1

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lichenized ascomycetes with stipitate apothecia (Lecanorales: Agyriineae). Plant Systematics

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Expedition. Publ. 22, XI. Botany 2. Stockholm.

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Beijing.

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https://doi.org/10.1016/B978-0-12-372180-8.50042-1

MY COTAXON

January-March 2019—Volume 134, pp. 155-160

https://doi.org/10.5248/134.155

Four Pyrenula species new to China

JrE-MENG Fu *“*, ANDRE APTROOT 3,

ZHONG-LIANG WANG *, Lu-LU ZHANG 7°

' Key Laboratory of Plant Stress Research, College of Life Sciences &

? Institute of Environment and Ecology,

Shandong Normal University, Jinan, 250014, PR. China

°ABL Herbarium, Gerrit van der Veenstraat 107, NL-3762 XK Soest, The Netherlands

" CORRESPONDENCE TO: * jiemengfu11@outlook.com, * andreaptroot@gmail.com,

AsstrRact—Pyrenula brunnea, P. punctella, P. subducta, and P. submastophora are reported

for the first time from China. This increases the number of Pyrenula species known from

China to 46. Descriptions and distribution of the four species are given.

Key worps—Asia, lichen-forming fungi, Pyrenulaceae, taxonomy

Introduction

Pyrenula Ach. (Pyrenulaceae, Eurotiomycetes) comprises crustose lichens

typically growing on smooth, shaded bark (Aptroot 2012) in the tropics

and subtropics (Mendonca & al. 2016). It is characterized by a corticate

(occasionally ecorticate) UV+ yellow thallus with or without pseudocyphellae

and with perithecioid ascomata, an occasionally inspersed hamathecium with

unbranched filaments, and distoseptate, bacillar or (sub)muriform ascospores

that are brown when mature with rounded, diamond-shaped, or elongated

lumina and with or without orange oil when over-mature.

In his world key to Pyrenula species, Aptroot (2012) accepted 169 species

out of the c. 745 named taxa in the genus. With many new species of Pyrenula

*Jie-MENG Fu & ANDRE Aprroorcontributed equally to this research.

156 ... Fu, Aptroot & al.

since described, the genus currently comprises c. 226 species worldwide (Ingle

& al. 2018), of which 42 are known from China (Wei 1991, Aptroot 2003,

Aptroot & Seaward 1999, Aptroot & Sipman 2001, Fu & al. 2018, Seaward &

Aptroot 2005, Wang & al. 2018).

The purpose of this paper is to contribute to the knowledge of Pyrenula

in China and to provide data for the LICHEN FLORA OF CHINA (in prep.).

Here we report four species from China for the first time—Pyrenula brunnea,

P. punctella, P. subducta, and P submastophora—increasing the number of

species known from China to 46.

The specimens studied are from Guizhou province, located in southern

China on the Yunnan-Guizhou plateau (24°37’-29°13’N 103°36’-109°35’E).

The landscape belongs to the plateau mountain region, with an average

elevation of about 1100 m. Guizhou has a subtropical humid monsoon climate

(rainy and hot at the same time) with the average temperature usually 22-25°C

during the warmest month (July); the annual precipitation averages 1200 mm

(rain mostly occurring during June-September).

Materials & methods

The specimens studied are preserved in the Lichen Section of the Herbarium,

College of Life Sciences, Shandong Normal University, Jinan, China (SDNU). Their

morphology was examined under a COIC XTL7045B2 stereo-microscope and

Olympus CX4lcompound microscope. Unless otherwise indicated, sections were

mounted and measured in tap water or enhanced with KOH. The lichen substances

were studied by standardized thin layer chromatography (TLC) using solvent system

C (Orange & al. 2010). Photos were taken with Olympus SZX16 and BX61 with DP72.

Taxonomy

Pyrenula brunnea Fée, Essai Crypt. Ecorc. 2: 81 (1837) FIG. 1A-D

MorpHoLocy—Thallus corticate, brownish, without pseudocyphellae,

UV-. Perithecia simple, solitary, often partly covered by the thallus, 0.4-0.5 mm

diam. Ostioles black, apical. Perithecial wall <150 um thick. Hamathecium not

inspersed with oil droplets. Ascospores 8/ascus, pale brown, 3-septate, with

rounded ends, 12.5-15 x 7.5 um, lumina in a straight line, somewhat rounded,

end lumina with a thick apical endospore layer, over-mature ascospores

shriveled, without red-orange oil.

CHEMISTRY—No chemical substances detected by TLC.

SPECIMEN EXAMINED: CHINA. GUIZHOU, Tongren, Jiangkou, Mt. Fanjing, alt. 1300 m,

on bark, 1 Nov. 2009, Zhao-Jie Ren 20101318 (SDNU).

DISTRIBUTION—Pantropical (Aptroot 2012). New to China.

New Pyrenula records for China... 157

ComMENTS—Pyrenula brunnea is similar to P. aspistea (Ach.) Ach., which has

narrower ascospores (4-6 um; Aptroot 2012). Our material fits the description

of P. brunnea specimens from northeast India quite well (Upreti 1991).

Pyrenula punctella (Nyl.) Trevis., Conspect. Verruc.: 13 (1860) FIG. 1E-H

MorpuHotocy —Thallus corticate, yellowish, with white pseudocyphellae,

UV-. Perithecia solitary, aggregated only by chance when crowded, 0.3-0.4

mm diam. Ostioles pale, apical. Perithecial wall <140 um thick. Hamathecium

not inspersed with oil droplets. Ascospores 8/ascus, brown, 3-septate, fusiform

with pointed ends, 37.5-40 x 12.5-15 um, lumina angular to usually somewhat

rounded, end lumina with a thick apical endospore layer, overmature spores

lacking orange oil.

CHEMISTRY —No chemical substances detected by TLC.

SPECIMEN EXAMINED: CHINA. GuIzHOU, Libo, Maolan protection zone, alt. 500 m, on

bark, 2 Nov. 2009, Hai-Ying Wang 20102422 (SDNU).

DISTRIBUTION— Temperate America, extending to Japan (Aptroot 2012). New

to China.

ComMMENTS—Pyrenula punctella is similar to P mastophora (Nyl.) Mill. Arg.,

which differs by lacking pseudocyphellae and having central lumina that are

much wider than long (Aptroot 2012). Our material closely fits the protologue

description of the P (= Verrucaria) punctella holotype specimen (Nylander

1858).

Pyrenula subducta (Nyl.) Mill. Arg., Flora 67: 666 (1884) FIG. 11-L

MorpuHo.toGcy— thallus corticate, yellowish brown, UV-. Perithecia simple,

only aggregated by chance when crowded, often largely covered by thallus,

0.4-0.6 mm diam. Ostioles brown, apical. Perithecial wall <150 um thick.

Hamathecium not inspersed with oil droplets. Ascospores 8/ascus, uniseriate,

brown, 3-septate, with rounded or pointed ends, 40-42.5(-45) x 15-17.5 um,

lumina in a straight line and somewhat rounded, end lumina with an apical

endospore layer, over-mature ascospores without orange oil.

CHEMISTRY—No chemical substances detected by TLC.

SPECIMENS EXAMINED: CHINA. GuIzHou, Tongren, Jiangkou, Mt. Fanjing, alt. 2300

m, on bark, 29 Mar. 2011, Xing-Ran Kou 20111283 (SDNU), Yu-Liang Chen 20111082

(SDNU). Libo, Maolan protection zone, alt. 500 m, on bark, 2 Nov. 2009, Yu-Liang Chen

20102362 (SDNU), Hai-Ying Wang 20102367 & 20102425 (SDNU); alt. 350 m, on bark,

2 Nov. 2009, Dai-Feng Jiang 20102432 (SDNU).

DISTRIBUTION—Neotropical (Aptroot 2012). New to China.

158 ... Fu, Aptroot & al.

ComMMENTS—Pyrenula subducta is similar to P caryae R.C. Harris, which

differs by the presence of black granules at the ascospore tips (Aptroot 2012).

Our material fits the description of India collections of P. subducta quite well,

but the Indian material has a brown to reddish-brown thallus (Ingle & al. 2018).

Pyrenula submastophora Ajay Singh & Upreti,

Geophytology 17(1): 85 (1987) FIG. 1M-P

MorpHo.tocy—Thallus corticate, brownish, without pseudocyphellae,

UV-. Perithecia simple, subglobose, erumpent, 0.4-0.6 mm diam. Ostioles

black, apical. Perithecial wall <160 um thick. Hamathecium not inspersed with

oil droplets, filaments unbranched. Ascospores 8/ascus, uniserial, grey-brown,

3-septate, fusiform with rounded ends, 21-23 x 7.5 um, end lumina with a

thick apical endospore layer. Over-mature spores lacking orange oil.

CHEMISTRY—No chemical substances detected by TLC.

SPECIMEN EXAMINED: CHINA. GuIzHOU, Libo, Maolan protection zone, alt. 500 m, on

bark, 2 Nov. 2009, Hai-Ying Wang 20102312 (SDNU).

DIsTRIBUTION—Eastern palaeotropical India (Aptroot 2012). New to China.

ComMMENTS—Pyrenula submastophora is similar to P acutispora Kalb &

Hafellner, which differs by its ascospores having at least one pointed end

(Aptroot 2012). Our specimen fits the description of Indian collections of

P. submastophora (including the holotype), but the Indian material has larger

perithecia (0.8-1.2 mm diam.; Upreti 1992).

Acknowledgements

The authors thank Dr. Harrie J.M. Sipman (Botanischer Garten and Botanisches

Museum Berlin Dahlem, Berlin, Germany) and Dr. Shou- Yu Guo (State Key Laboratory

of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing, China)

for presubmission reviews. This study was supported by the Emergency Management

Project of National Natural Science Foundation of China (31750001), the National

Natural Science Foundation of China (31400015), and the Youth Science Foundation

(31600100).

Fic. 1. Pyrenula brunnea (Ren 20101318, SDNU). A. thallus with ascomata; B. transverse section

through ascoma; C. ascus; D. ascospores. Pyrenula punctella (Wang 20102422, SDNU). E. thallus

with ascomata; F: transverse section through ascoma. G: ascus. H: ascospores. Pyrenula subducta

(Kou 20111283, SDNU). I. thallus with ascomata; J. transverse section through ascoma; K. ascus

with 8 ascospores; L. ascospores. Pyrenula submastophora (Wang 20102312, SDNU). M. thallus

with ascomata; N. transverse section through ascoma; O. ascus with 8 ascospores; P. ascospores.

Scale bars: E= 1 mm; A, I, M = 2 mm; C, D, G, H, L, O, P = 20 um; B, K, N = 50 um; EF, J = 100 um.

159

New Pyrenula records for China ...

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160 ... Fu, Aptroot & al.

Literature cited

Aptroot A. 2003. Pyrenocarpous lichens and related non-lichenized ascomycetes from Taiwan.

Journal of the Hattori Botanical Laboratory 93: 155-173.

Aptroot A. 2012. A world key to the species of Anthracothecium and Pyrenula. Lichenologist 44:

1-54. https://doi.org/10.1017/S0024282911000624

Aptroot A, Seaward MRD. 1999. Annotated checklist of Hongkong Lichens. Tropical Bryology 17:

57-101.

Aptroot A, Sipman HJM. 2001. New Hong Kong lichens, ascomycetes and lichenicolous fungi.

Journal of the Hattori Botanical Laboratory 91: 317-343.

Fu JM, Wang ZL, Wang CX, Zhang LL. 2018. New records of six Pyrenula species from China.

Mycotaxon 133(3): 473-480. https://doi.org/10.5248/133.473

Ingle KK, Uppadhyay V, Nayaka S, Trivedi S, Sahoo D. 2018. New records and an updated

key of Pyrenula from India. Cryptogam Biodiversity and Assessment 1: 37-46.

https://doi.org/10.21756/cab.esp7

Mendonga CO, Aptroot A, Caceres MES. 2016. Six new species of Pyrenula (Pyrenulaceae) from

Northeast Brazil. Phytotaxa 286(3): 169-176. https://doi.org/10.11646/phytotaxa.286.3.4

Nylander, W. 1858. Expositio synoptica Pyrenocarpeorum. Mém. Soc. Acad. Maine-et-Loire

4: 5-88.

Orange A, James PW, White FJ. 2010. Microchemical methods for the identification of lichens.

2nd edition. British Lichen Society, London.

Seaward MRD, Aptroot A. 2005. Hong Kong lichens collected on the United States North

Pacific Exploring Expedition, 1853-1856 Bryologist, 108(2): 282-286.

https://doi.org/10.1639/0007-2745(2005)108[0282: HKLCOT]2.0.CO;2

Upreti DK. 1991. Lichen genus Pyrenula from India: the species with spores of Pyrenula

brunnea type. Bulletin de la Société Botanique de France 138, Lettres Botaniques (3):

241-247. https://doi.org/10.1080/01811797.1991.10824926

Upreti DK. 1992. Lichen genus Pyrenula from India VII. Pyrenula mastophora spore type.

Feddes Repertorium 103(3-4): 279-296. https://doi.org/10.1002/fedr. 19921030315

Wang ZL, Yan SK, Tang R, Sun MJ, Zhang LL. 2018. New records of Lepraria and Pyrenula

from China. Mycotaxon 133(1): 89-96. https://doi.org/10.5248/133.89

Wei JC. 1991. An enumeration of lichens in China, International Academic Publishers, Beijing.

MY COTAXON

January-March 2019—Volume 134, pp. 161-167

https://doi.org/10.5248/134.161

Spadicoides matsushimae sp. nov., and

Anisospadicoides gen. nov. for two

atypical Spadicoides species

MIN Q1A0’", DE-WEI LI*?, ZE-FEN Yu™,

KAI ZHANG‘, RAFAEL EF CASTANEDA-RUIZ>

‘Laboratory for Conservation and Utilization of Bio-resources,

Key Laboratory for Microbial Resources of the Ministry of Education,

Yunnan University, Kunming, Yunnan, 650091, China

? The Connecticut Agricultural Experiment Station,

Valley Laboratory, 153 Cook Hill Road, Windsor, CT 06095, USA

°Co-Innovation Center for Sustainable Forestry in Southern China,

Nanjing Forestry University, Nanjing, Jiangsu 210037, China

‘Department of Landscaping, Shandong Yingcai University, Jinan, 250104, China

° Instituto de Investigaciones Fundamentales en Agricultura Tropical Alejandro de Humboldt

(INIFAT), Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200

*CORRESPONDENCE TO: zfyuqm@hotmail.com

ABSTRACT—Anisospadicoides is proposed as a new genus for two species of Spadicoides,

S. macrocontinua and S. macroobovata, that have polytretic and blastic conidial ontogeny

on the apical conidiogenous cells. A new species, Spadicoides matsushimae, distinguished

by broadly fusiform to navicular, mostly 2-septate, brown to dark brown conidia, is also

described and illustrated.

KEY worDs—asexual Ascomycota, Helminthosphaeriaceae, hyphomycetes, systematics

Introduction

Spadicoides S. Hughes, typified by S. bina (Corda) S. Hughes, is

distinguished by macronematous, mononematous, and _ generally

unbranched erect conidiophores. Conidiogenous cells are polytretic,

integrated, terminal and intercalary, determinate, with conidiogenous loci

162 ... Qiao & al.

at the apex and laterally below. The conidia are solitary, variously shaped

(ellipsoidal, oblong, obovoid, obclavate, globose, clavate, or cuneate),

unicellular or multicellular, euseptate, smooth or verruculose, subhyaline to

brown or black, and dry (Ellis 1971, Hughes 1958, Ma & al. 2016, Xia & al.

2013, Whitton & al. 2012).

Two Spadicoides species, S. macrocontinua and S. macroobovata, have an

unusual combination of tretic and holoblastic conidial development on the

apical conidiogenous cells (Matsushima 1993, 1995). Conidial ontogeny and

conidiogenous events have been considered the fundamental criteria for

asexual fungal taxonomy and generic delimitation (Kendrick 2017, Seifert

& al. 2011). Consequently, we propose the new genus Anisospadicoides to

accommodate two atypical Spadicoides species.

In 1993, Matsushima published a description and illustrations of a

conspicuous fungus found on rotten petiole of a palm, but he did not

provide a species epithet for it, merely calling it “Spadicoides sp?’ This fungus

is formally named here as a new species, S. matsushimae.

Taxonomy

Anisospadicoides R.F. Castafieda, Qiao & Z.F. Yu, gen. nov. Fie. 1

MycoBAnk MB 827953

Differs from Spadicoides species by its holoblastic, mostly monoblastic conidial

ontogeny, and production of conidia on the apical conidiogenous cells beside or near

the tretic loci.

TYPE SPECIES: Spadicoides macrocontinua Matsush. [= Anisospadicoides macrocontinua

(Matsush.) R.E Castafieda & al.]

EryMo ocy: Greek, aniso- meaning unequal, uneven, or dissimilar + Latin, -spadicoides,

referring to the genus Spadicoides.

CoLonies on CMA medium felted or hairy, brown to dark brown.

CONIDIOPHORES macronematous, mononematous, unbranched, erect,

straight, cylindrical, septate, smooth, brown to dark brown. CONIDIOGENOUS

CELLS biontogenous, sometimes on the same cell: i) polytretic, integrated,

mostly intercalary, cylindrical, sometimes terminal with subapical tretic

loci and apical blastic loci; ii) monoblastic, integrated, terminal, sometimes

with several tretic loci near below the blastic locus on the same cell.

Fic. 1. Anisospadicoides macrocontinua (holotype, MFC OP-302): A. Conidiogenous cells with

apical conidia originated by holoblastic mode and subapical conidia borne from tretic loci;

B. Conidiogenous cells with apical conidia originated by holoblastic mode and subapical tretic

loci.

Anisospadicoides gen. & spp. nov. & Spadicoides matsushimae sp. nov. ... 163

10 um

read

Sette

= *e%*

”

10 um

164 ... Qiao & al.

10 um

Fic. 2. Anisospadicoides macroobovata (holotype, MFC 4P-562): A. Conidia produced after

holoblastic mode on the apical loci; B. Conidia produced after enteroblastic mode on the subapical

and intercalary loci; C. Conidiogenous cells with apical blastic loci and subapical and intercalary

tretic loci.

Anisospadicoides gen. & spp. nov. & Spadicoides matsushimae sp. nov. ... 165

ConrpiA solitary, dimorphic: i) obovoid, subpyriform, truncate at the base,

acropleurogenous, mostly pleurogenous, unicellular or septate, smooth,

brown to dark brown or black, dry, borne on tretic conidiogenous loci;

ii) obovoid, broadly subfusiform, to subclavate, attenuate or slightly fimbriate

towards the truncate base, acrogenous, unicellular or septate, euseptate,

smooth, brown to dark brown or black, dry, borne on blastic conidiogenous

loci.

Anisospadicoides macrocontinua (Matsush.) R.F. Castafieda, Qiao &

Z.F. Yu, comb. nov. Fic. 1

MycoBAnkK MB 827954

= Spadicoides macrocontinua Matsush., Matsush. Mycol. Mem.7: 67 (1993).

Anisospadicoides macroobovata (Matsush.) Qiao, Z.F. Yu & R.F. Castafieda,

comb. nov. Fic. 2

MycoBAnk MB 827955

= Spadicoides macroobovata Matsush., Matsush. Mycol. Mem. 8: 36 (1995).

Spadicoides matsushimae R.F. Castafieda & D.W. Li, sp. nov. FIG. 3

MycoBank MB 809530

Differs from Spadicoides curvularioides by its broadly fusiform, smaller, 2-septate,

brown conidia.

Type: Peru, Rio Sinchicuy, 14°36’N 90°38’W, on decaying petiole of an unidentified

palm, June 1991, coll. T. Matsushima (Holotype, MFC 1P-294).

EryMo_oey: Latin, matsushimae, in reference to Takashi Matsushima, collector of the

holotype specimen.

CONIDIOPHORES on the natural substratum macronematous, mononematous,

unbranched, erect, straight, rigid, septate, brown, smooth, 50-160 um

long, 4.5-6 um wide at the base, CONIDIOGENOUS CELLS polytretic,

integrated, terminal and intercalary, determinate, brown. Conipia solitary,

acropleurogenous, broadly fusiform to broadly navicular, conical-truncate at

the base, rounded at the apex, (1—) 2-septate, smooth, 15-19 x 7-9 um, with

central cell brown to dark brown and larger than pale brown end cells.

Notes: There are 58 previously described species of Spadicoides (Index

Fungorum 2018), of which only 42 were accepted by Goh & Hyde (1996),

Kuthubutheen & Nawawi (1991), Ma & al. (2014, 2016), Whitton & al.

(2012), and Xia & al. (2013). Ma & al. (2016) provided a comparative table

of Spadicoides spp. and a key for their identification. Among the recognized

Spadicoides species, only S. curvularioides B. Sutton & Hodges is superficially

similar to S. matsushimae in producing more or less fusiform or navicular

166 ... Qiao & al.

wil OT

B. Conidiogenous cells and conidiogenous loci.

Fic. 3. Spadicoides matsushimae (holotype, MFC 1P-294):

A. Conidia;

conidia, but S. curvularioides has pale brown conidia that are paler at the

and verruculose (Sutton & Hodges

longer (22-33.5 um),

ends, 3-septate,

1978).

Anisospadicoides gen. & spp. nov. & Spadicoides matsushimae sp. nov. ... 167

Acknowledgments

This work was financed by the National Natural Science Foundation Program of

PR China (31760012, 31570023). The authors express their sincere gratitude to Dr.

Xiu-Guo Zhang and Dr. Josiane S. Monteiro for their critical review of the manuscript.

We acknowledge the facilities provided by Dr. P.M. Kirk and Dr. K. Bensch through

the Index Fungorum and MycoBank websites. Dr. Lorelei Norvell’s editorial review

and Dr. Shaun Pennycook’s nomenclature review are greatly appreciated.

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Kendrick B. 2017. The fifth kingdom (4" ed). Hackett publishing company. 416 p,

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https://doi.org/10.1007/s11557-013-0937-z

Ma J, Castafieda-Ruiz RF, Zhang XG. 2016. Three new species of Spadicoides from Lushan

Mountain. Mycological Progress 15: 43 [8 p.]. https://doi.org/10.1007/s11557-016-1185-9

Matsushima T. 1993. Matsushima Mycological Memoirs no. 7. Matsushima Fungus Collection,

Kobe, Japan.

Matsushima T. 1995. Matsushima Mycological Memoirs no. 8. Matsushima Fungus Collection,

Kobe, Japan.

Seifert K, Morgan-Jones G, Gams W, Kendrick B. 2011. The genera of hyphomycetes. CBS

Biodiversity Series 9. 997 p.

Sutton BC, Hodges CS. 1978. Eucalyptus microfungi: Chaetendophragmiopsis gen. nov. and

other hyphomycetes. Nova Hedwigia 29: 593-607.

Whitton SR, McKenzie EHC, Hyde KD. 2012. Anamorphic fungi associated with Pandanaceae.

125-353, in: SR Whitton & al. (eds). Fungi Associated with Pandanaceae. Fungal Diversity

Research Series 21. https://doi.org/10.1007/978-94-007-4447-9_4

Xia JW, Ma LG, Ma J, Zhang XG. 2013. Two new species of Spadicoides from southern China.

Mycotaxon 126: 55-60. https://dx.doi.org/10.5248/126.55

MYCOTAXON

January-March 2019—Volume 134, pp. 169-175

https://doi.org/10.5248/134.169

New records of Hymenoscyphus, Parascutellinia,

and Scutellinia for Turkey

ALI KELES’

Department of Science and Mathematics Education, Yiiziincti Yil University,

Zeve Campus, Van, Turkey

“ CORRESPONDENCE TO: alikeles61@yahoo.com

ABSTRACT—Three ascomycetous macrofungi—Hymenoscyphus kathiae, Parascutellinia

violacea, and Scutellinia crinita—are recorded for the first time from Turkey, with P violacea

serving also the first record of the genus for the country. The species are described briefly and

accompanied by photographs illustrating their macro- and micro-morphologies.

Key worps—biodiversity, Helotiaceae, Pyronemataceae, Siirt, Van

Introduction

Since studies on Turkish macromycota started in the second quarter of the

20th century, almost 2500 species have been reported. Considering the 15,000

macrofungi reported from Europe (Luki¢ 2009), it can easily be said that there

is still much to be done.

This study aims to identify new records for the mycobiota of Turkey.

During routine field studies some ascomycete samples were collected and

identified as Hymenoscyphus kathiae (Helotiaceae), Parascutellinia violacea

(Pyronemataceae), and Scutellinia crinita (Pyronemataceae). Among the 220

ascomycete species from Turkey covered by current checklists (Sesli & Denchev

2014, Solak & al. 2015) and recent publications on Turkish Ascomycota (Kaya

& Uzun 2015, 2018; Acar & Uzun 2016; Kaya & al. 2016, 2018; Akcay & Uzun

2016; Dogan & al. 2016; Keles & Orug 2017; Uzun & al. 2017a,b, 2018a,b; Isik

& Turkekul 2018), these three species are not cited and represent new records

for the country.

170 ... Keles

Materials & methods

Fungal specimens were collected from Baykan (Siirt) and from Gevas and

Giirpinar (Van) districts during 2012-2018. The fungi were photographed in their

natural habitats and their ecological and morphological characteristics noted.

Specimen samples were mounted in Congo red or Melzer’s reagent and examined

microscopically using a Leica DM500 light microscope. The fungi were identified

after consulting Svrcek (1975), Schumacher (1990), Korf (1999), Garcia & Van Vooren

(2005), Medardi (2006), and Van Vooren & Hairaud (2009). The collections are kept

in the fungarium of Yiiziincii Yil University, Van, Turkey (VANF).

Taxonomy

Hymenoscyphus kathiae (Korf) Baral, Fl. Mediterr. 15: 66 (2005) FIG. 1

APOTHECIA 1-3 mm diam., subsessile to shortly stipitate, cupuliform, then

more or less plane, margin smooth and glabrous. HyMENIuM smooth, pale

yellow to golden yellow, outer surface almost concolorous with the hymenium

or lighter. StrPE noticeably short, broad, yellowish white. Asci 90-110 x 7-10

uum, cylindrical, slightly attenuated at base and arising from croziers, generally

8-spored, some with fewer spores. ASCOSPORES 9-14(-16) x 4-5 um, ellipsoid,

hyaline, smooth, some septate when mature, with two large or small guttules,

uniseriate. PARAPHYSES cylindrical, rarely septate, hyaline, generally filled with

yellowish guttules.

SPECIMEN EXAMINED—TURKEY, Sut, Baykan, central district, on wood remnants,

38°09°37”N 41°47°17”E, 769 m, 17.11.2012, A.K. 1877 (VANE).

CoMMENTS—Hymenoscyphus kathiae is macroscopically similar to H. imberbis,

which differs in its ascospores with two large lipid droplets at the poles

(Dimitrova & Baral 2005). In general, the morphological characteristics of our

Turkish material agree with the descriptions by Dimitrova & Baral (2005) and

Van Vooren & Hairaud (2009). This species is a new record for Turkey.

Parascutellinia violacea (Velen.) Svréek, Ceska Mykol. 29: 129 (1975) FiG..2:

APOTHECIA 3-6 mm diam., violet in colour, usually found in small groups;

surface elliptical, smooth. HyMENIUM carmine red, with some violet tones.

Asci 260-320 x 16-17 um, cylindrical, operculate, inamyloid. AscosPoREs

24-29 x 12-14 um, hyaline, elliptical, thick-walled, with one large guttule

and two smaller guttules. PARAPHYSES multiseptate, branched, with expanded

apex, containing a red-orange to slightly rosy pigment that turns greenish

in Meltzer’s. PARAMARGINAL HAIRS yellowish-brown, fusiform, 75-150 um,

tightly crowded, fasciculate, multiseptate, with acute or obtuse apex, some well

differentiated, others only poorly, with thickened walls <3 um.

Ascomycete species new for Turkey... 171

—

- ed A

Fic 1. Hymenoscyphus kathiae (VANF A.K. 1877)

a. Ascomata; b. Ascospores; c. Tips of asci and paraphyses; d. Ascus;

e. Paraphyses; f. Ascus base.

Fic 2. Parascutellinia violacea (VANF A.K. 2961)

a. Ascomata; b. Ascospores; c. Paraphyses; d. Hairs; e. Asci

SPECIMEN EXAMINED—TURKEY, VAN, Giirpinar, Gizelsu village, river side, under

Salix sp., 38°18’59” N-43°48'00”E, 1977 m, 05.10.2016, A.K. 2961 (VANF).

ComMMENTS— Parascutellinia violacea resembles P carneosanguinea, which

differs by carotenoid pigments within its paraphyses (Svrcek 1975). Although

Ascomycete species new for Turkey ... 173

Van Vooren (2007) reported the same maximum sizes for the ascomata (5 mm)

and asci as for P. violacea, some fruit bodies in our Turkish material measured

almost 6 mm with asci up to 320 um. Our specimens represent the first record

of both genus and species for Turkey.

Fic 3. Scutellinia crinita (VANF A.K. 2963)

a. Ascomata; b. Ascospores; c. Ascal bases; d. Hairs; e. Asci;

f. Paraphyses; g. Apices of paraphyses.

174 ... Keles

Scutellinia crinita (Bull.) Lambotte,

Mém. Soc. roy. Sci. Liege, Sér. 2, 14: 301 (1887) FIG. 3

APOTHECIA 3-7(-9) mm diam., disc-shaped, without stalk. HYMENIUM light

orange, edges and outer surface covered with long dark brown hairs. Hairs

880-1550 x 22-40 um, thick-walled, multiseptate, dark brownish, surface

mostly smooth, sometimes cracked. Asc1 207-250 x 12-22 um, and cylindrical.

ASCosPoRES 16-22 x 15-18 um, broadly rounded ellipsoid, slightly verrucose,

containing many small droplets. Parapuysss filiform, clavate at apex, septate,

somewhat branched.

SPECIMENS EXAMINED— TURKEY, VAN, Gevas, at the vicinity of Kuzgunkiran tunnel,

on rotten and damp wood under Populus sp., 38°22’09”°N-42°47°40” E, 2052 m,

13.06.2018, A.K. 2963 (VANF).

CoMMENTS— The maximum size of ascomata for Scutellinia crinita was

reported as 10 mm by Matoéec & Antoni¢ (1995) and 18 mm by Thompson

(2013), but none of our ascomata measured >9 mm. This species represents a

new record for Turkey.

Acknowledgments

The author would like to thank Prof. Dr. Ertugrul Sesli, Prof. Dr. Ibrahim Tiirkekul,

and Dr. Shaun Pennycook for their helpful comments and careful review.

Literature cited

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Yil Universitesi Fen Bilimleri Enstitiisii Dergisi 21 (1):39-42.

Akcay ME, Uzun Y. 2016. Belonidium mollissimum (Lachnaceae): Turkiye Mikotasi i¢in yeni bir

tiir. Mantar Dergisi 7(2): 118-121. https://doi.org/10.15318/Fungus.2016222679

Dimitrova EG, Baral HO. 2005. Checklist of Bulgarian Helotiaceae (Ascomycetes). Flora

Mediterranea 15: 57-72.

Dogan HH, Bozok F, Taskin H, Biytikalaca $ 2016. Tiirkiye icin bes yeni Morchella kaydh,

Alatarim, 15 (1): 1-11.

Garcia G, Van Vooren N. 2005. Un discomycéte inoperculé plutét discret, Pezoloma ciliifera,

et remarques sur le genre Pezoloma. Bulletin Mensuel de la Societe Linnéenne de Lyon

74: 115-129. https://doi.org/10.3406/linly.2005.13567

Isik H, Tirkekul I. 2018. A new record for Turkish mycota from Tokat Province: Arachnopeziza

aurelia (Pers.) Fuckel. Journal of Fungus 9(1): 54-57.

Kaya A, Uzun Y. 2015. Six new genus records for Turkish Pezizales from Gaziantep Province.

Turkish Journal of Botany 39: 506-511. https://doi.org/10.3906/bot- 1409-3

Kaya A, Uzun Y. 2018. New contributions to the Turkish Ascomycota. Turkish Journal of Botany

42(5): 644-652. https://doi.org/10.3906/bot-1712-1

Kaya A, Uzun Y, Karacan Hi, Yakar S. 2016. Contributions to Turkish Pyronemataceae from

Gaziantep Province. Turkish Journal of Botany 40: 298-307.

https://doi.org/10.3906/bot- 1508-4

Ascomycete species new for Turkey... 175

Kaya A, Uzun Y, Karacan IH, Yakar S. 2018. New additions to Turkish Helotiales

and Orbiliales. Kastamonu University Journal of Forestry Faculty, 18(1): 46-52.

https://doi.org/10.17475/kastorman.290359

Keles A, Orug Y. 2017. Leucocoprinus brebissonii (Godey) Locq, a new record for Turkish

mycobiota. Anatolian Journal of Botany 1(2): 49-51. https://doi.org/10.30616/ajb.348736

Korf RP. 1999. Pezoloma kathiae sp. nov. (Ascomycetes: Leotiales, Leotiaceae), and its placement in

a new subgenus, Phaeopezoloma. Mycotaxon, 73: 493-497.

Luki¢ N. 2009. The distribution and diversity of Boletus genus in Central Serbia. Kragujevac

Journal of Science Sci 31: 59-68.

Mato¢ec N, Antoni¢ O. 1995. The genus Scutellinia (Pezizales, Ascomycotina) in Croatia:

preliminary part. Natura Croatica, 4(1): 1-58.

Medardi G. 2006. Atlante fotografico degli ascomiceti d'Italia. Vicenza, Italy: Centro Studi

Micologici.

Schumacher T. 1990. The genus Scutellinia (Pyronemataceae). Opera Botanica 101. 107 p.

Sesli E, Denchev CM 2014. Checklists of the myxomycetes, larger ascomycetes, and larger

basidiomycetes in Turkey. 6th edn. Mycotaxon Checklists Online.

(http://www.mycotaxon.com/resources/checklists/sesli-v106-checklist.pdf)

Solak MH, Isiloglu M, Kalmis E, All H. 2015. Macrofungi of Turkey, checklist, vol. I. Izmir,

Turkey: Universiteliler Ofset (in Turkish).

Svréek M. 1975. New or less known Discomycetes. II. Ceska Mykologie. 29(3):129-134.

Thompson PI. 2013. Ascomycetes in colour: found and photographed in mainland Britain.

Indiana, USA: Xlibris.

Uzun Y, Acar I, Akcay ME, Kaya A 2017a. Contributions to the macrofungi of Bingél, Turkey.

Turkish Journal of Botany 41: 516-534. https://doi.org/10.3906/bot- 1611-14

Uzun Y, Karacan IH, Yakar S, Kaya A. 2017b. Octospora Hedw., a new genus record for Turkish

Pyronemataceae. Anatolian Journal of Botany 1: 18-20. https://doi.org/10.30616/ajb.342114

Uzun Y, Karacan IH, Yakar S, Kaya A. 2018a. New bryophilic Pyronemataceae records

for Turkish Pezizales from Gaziantep province. Anatolian Journal of Botany 2: 28-38.

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Uzun Y, Yakar S, Karacan IH, Kaya A. 2018b. New additions to the Turkish Pezizales. Turkish

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Van Vooren N. 2007. Description et commentaire de quelques ascomycetes récoltés a la session

de Lamoura (Jura). In Annales 2007. Compte rendu de la session mycologique de la FMBDS

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Linnéenne de Lyon.

Van Vooren N, Hairaud M. 2009. Note sur Hymenoscyphus kathiae (Helotiales). Ascomycete.org

1(4): 11-15.

MY COTAXON

January-March 2019—Volume 134, pp. 177-181

https://doi.org/10.5248/134.177

Lemonniera yulongensis sp. nov.

from Yunnan, China

ZE-FEN YU*”*, YI-FAN Lv*?*, BO FENG’, MIN QIAo**

"Laboratory for Conservation and Utilization of Bio-resources, Key Laboratory for

Microbial Resources of the Ministry of Education & ? School of Life Sciences,

Yunnan University, Kunming, Yunnan, 650091, P. R. China

* CORRESPONDENCE TO: Qidoming@ynu.edu.cn

ABSTRACT—Lemonniera yulongensis, isolated from submerged decaying leaves on Yulong

Snow Mountain, is described as a new species. The fungus is characterized by monophialidic

conidiogenous cells and branched conidia with mainly four septate arms. The new species

also extends the distribution of Lemonniera into higher elevations than is typical for the

genus.

KEY woRDS—aquatic hyphomycetes, fungal diversity, Helotiales, taxonomy

Introduction

Lemonniera De Wild. was proposed with L. aquatica De Wild. as type

species (De Wildeman 1894). The genus is characterized by typically

tetraradiate conidia with branches arising more or less simultaneously from

a globose primordium and phialidic conidiogenous cells growing at the tip of

branched or unbranched conidiophores. Seven species are currently accepted

in the genus; Descals & al. (1977) reviewed the morphology and ecology and

provided a key to six species, and Sinclair & Morgan-Jones (1979) described

a new species isolated from an Alabama stream. Although some Lemonniera

strains have been collected in terrestrial habitats, most have been isolated

from streams, rivers, or ponds (Descals & al. 1977, Sinclair & Morgan-Jones

1979). The cosmopolitan genus has been recorded from Africa, Asia, Europe,

* Yu & Lv contributed equally to this research.

178 ... Fu, Lv &al.

North America, and Australasia (De Wildeman 1894; Ingold 1942, 1958,

1968; Ranzoni 1953; Tubaki 1958; Petersen 1963; Descals & al. 1977; Sinclair

& Morgan-Jones 1979; Seifert & al. 2011).

During a study of aquatic hyphomycetes in Yunnan Province, China, we

encountered an unknown fungus that corresponded with the features of

Lemonniera. We describe it here as a new species, L. yulongensis.

Materials & methods

The culture was isolated from decaying leaves submerged in a river on Yulong

Snow Mountain, Lijiang county, Yunnan province, in April 2015. The leaves were cut

into 2-4 x 2-4 cm fragments and then stretched out on the surface of CMA (20 g

cornmeal, 18 g agar, 40 mg streptomycin, 30 mg ampicillin, 1000 ml distilled water)

for ten days. Single conidia were captured using a sterilized toothpick while viewing

with a CX31 microscope and cultivated on CMA in Petri plates. The cultures were

examined after a one-week incubation at 28°C and morphological characters noted.

Pure cultures and a permanent slide have been deposited in the Herbarium of the

Laboratory for Conservation and Utilization of Bio-resources, Yunnan University,

Kunming, P.R. China (YMF; formerly Key Laboratory of Industrial Microbiology and

Fermentation Technology of Yunnan).

Taxonomy

Lemonniera yulongensis Z.F. Yu, sp. nov. PLATE 1

MycoBank MB 827810

Differs from Lemonniera alabamensis by its wider, 1(-2)-septate conidial arms.

Type: China, Yunnan Province, Lijiang county, Yulong Snow Mountain, 27°06’32”N

100°15’12”E, alt. 3033 m, in a river on submerged leaves of an unidentified

dicotyledonous plant, 4 Apr. 2015, Ze-Fen Yu. (Holotype, YMF 1.044181 [metabolically

inactive permanent slide]); ex-type culture, YMF1.04418).

EryMmo .oey: Latin, yulongensis, referring to the site from which the species was

isolated.

Cotoniges 20 mm diam. on CMA after 20 days at 25°C. Mycelium hyaline,

partly superficial and partly immersed, composed of branched, septate,

hyaline hyphae. CONIDIOPHORES macronematous, mononematous, sometimes

unbranched or bifurcate or penicillate branched towards the apex, septate,

50-200 x 3-4 um, straight or flexuous, CONIDIOGENOUS CELLS monophialidic,

terminal, integrated or discrete, hyaline, 17.5-27.5 x 6-6.5 um. CONIDIA

hyaline, radial, with a very distinct spherical or globose central cell of 4-6

um diam, connected by 4 (rarely 3 or 5) arms radially extending from the

globose central cell, branches more or less equilateral, subulate, apex rounded,

1(-2)-septate, (17.5-)22-42 x 4.8-6.1(-8.0) um.

Lemonniera yulongensis sp. nov. (China) ... 179

PLATE 1. Lemonniera yulongensis (holotype, YMF 1.04418): A, B. Conidia; C. Conidia growing

from old conidia. D, E. Conidiogenous cells. F. Conidiophores and conidia. Scale bars: A-E = 10

um; F = 20 um.

Discussion

Lemonniera yulongensis fits well within the genus Lemonniera based on its

typically tetraradiate conidia with branches arising from the central body and

phialidic conidiogenous cell growing at the tip of main apex or branches of

conidiophores.

180 ... Fu, Lv & al.

Lemonniera species have been divided into two groups based on the

presence or absence of a central body in the conidia. Lemonniera alabamensis

R.C. Sinclair & Morgan-Jones, L. centrosphaera Marvanova, L. pseudofloscula

Dyko, and L. yulongensis all possess central bodies in their conidia. However,

the central conidial body is not as distinct in L. yulongensis as those of the other

three species. The conidial arms in L. centrosphaera and L. pseudofloscula are

longer than in L. yulongensis; L. alabamensis, which produces similarly shaped

conidia, is distinguished from L. yulongensis by its 2-septate, narrower (2-3 um

diam.) arms (Sinclair & Morgan-Jones 1979).

Five Lemonniera species have been reported previously from China:

L. alabamensis from Beijing (Liu & al. 1992), L. centrosphaera from Anhui

and L. cornuta Ranzoni from Beijing (Yu 1988), L. filiformis R.H. Petersen ex

Dyko from Anhui (Yu 1988) and Taiwan (Chao 2006), and L. aquatica and

L. terrestris Tubaki from Sichuan (Zhu & Yu 1992). The localities of all five

occur at low (970-1100 m) elevations, so the discovery of L. yulongensis at

higher altitudes area widens the known distribution of the genus.

Acknowledgements

This work was jointly financed by National Natural Science Foundation Program

of PR China (31570023, 31770026). We are grateful to Prof. X.G. Zhang and Dr.

R.E Castafieda-Ruiz for critically reviewing the manuscript and providing helpful

suggestions to improve this paper, and to Dr. Shaun Pennycook and Dr. Lorelei Norvell

for their critical review and suggestions.

Literature cited

Chao TH. 2006. Aquatic hyphomycetes in Taiwan. Taipei: National Taiwan University. 72 p.

Descals E, Webster J, Dyko BS. 1977. Taxonomic studies on aquatic hyphomycetes. I. Lemonniera

De Wildeman. Transactions of the British Mycological Society 69: 89-109.

https://doi.org/10.1016/S0007-1536(77)80120-4

De Wildeman E. 1894. Notes mycologiques. Fascicle 3. Annales de la Société Belge de

Microscopie 18: 135-161.

Ingold CT. 1942. Aquatic hyphomycetes of decaying alder leaves. Transactions of the British

Mycological Society 25: 339-417. https://doi.org/10.1016/S0007-1536(42)80001-7

Ingold CT. 1958. New aquatic hyphomycetes: Lemonniera brachycladia, Anguillospora crassa

and Fluminispora ovalis. Transactions of the British Mycological Society 41: 365-372.

https://doi.org/10.1016/S0007-1536(58)80051-0

Ingold CT. 1968. More spores from rivers and streams. Transactions of the British Mycological

Society 51: 138-143. https://doi.org/10.1016/S0007-1536(68)80130-5

Liu XJ, Zhu XD, Yu YN. 1992. Aquatic hyphomycetes in water foam from Beijing region. Acta

Mycologica Sinica 11: 111-118.

Petersen RH. 1963. Aquatic hyphomycetes from North America HI. Phialosporae and

miscellaneous species. Mycologia 55: 570-581. https://doi.org/10.2307/3756434

Lemonniera yulongensis sp. nov. (China) ... 181

Ranzoni FV. 1953. The aquatic hyphomycetes of California. Farlowia 4: 353-398.

Seifert K, Morgan-Jones G, Gams W, Kendrick B. 2011. The genera of hyphomycetes. CBS

Biodiversity Series 9. 997 p.

Sinclair RC, Morgan-Jones G. 1979. Notes on hyphomycetes. XXXII. Five new aquatic species.

Mycotaxon 9: 469-481.

Tubaki K. 1958. Studies on the Japanese hyphomycetes. V. Leaf and stem group with a discussion

of the classification of hyphomycetes and their perfect stages. Journal of the Hattori Botanical

Club 20: 142-244.

Yu YN, Li JL, Yu JQ. 1988. Notes on some aquatic hyphomycetes from Sichuan province of

China. Acta Mycologica Sinica 7(3): 138-148.

Zhu XD, Yu YN. 1992. Twenty-three species of aquatic hyphomycetes new records to China.

Acta Mycologica Sinica 11: 32-42.

MY COTAXON

January-March 2019—Volume 134, pp. 183-196

https://doi.org/10.5248/134.183

Contributions to species of Xylariales in China—2.

Rosellinia pervariabilis and R. tetrastigmae spp. nov.,

and anew record of R. caudata

XIN XIE’, LiL Liu?4, XU ZHANG?*, QINGDE LONG?*, XIANGCHUN SHEN®",

SARANYAPHAT BOONMEE®, JICHUAN KANG**, QiRur L1*34*

' Engineering and Research Center for Southwest Bio-Pharmaceutical Resources of

National Education Ministry of China &

? Department of Plant Pathology, College of Agriculture

Guizhou University, Guiyang, Guizhou 550025, China

State Key Laboratory of Functions and Applications of Medicinal Plants,

Guizhou Medical University, Guiyang 550004, China

‘The Key Lab of Optimal Utilization of Natural Medicine Resources,

School of Pharmaceutical Sciences, Guizhou Medical University,

University Town, Guian New District, Guizhou 550025, China

° Center of Excellence in Fungal Research, Mae Fah Luang University,

Chiang Rai 57100, Thailand

“ CORRESPONDENCE TO: “ jckang@gzu.edu.cn *lqrnd2008@163.com

ABSTRACT—Three species of Rosellinia collected from China are reported based on

morphological and molecular characteristics. Rosellinia pervariabilis is proposed as a new

species, differentiated by its narrow ascospores (19.5-24.5 x 4-5 um) without a germ slit.

Rosellinia tetrastigmae is also proposed as new and differentiated by its small stromata.

Rosellinia caudata is reported as a new record from the Chinese mainland. Detailed

morphological descriptions and illustrations are provided, and many new DNA sequences

are cited.

KEY worRDS—Ascomycota, pyrenomycetous fungi, Xylariaceae, taxonomy

Introduction

This is the second in a planned series of papers on Xylariales from China

(Long & al. 2019). Rosellinia, introduced by De Notaris (1844), a large genus

184 ... Xie & al.

in terms of species number in the Xylariaceae Tul. & C. Tul. (Wijayawardene

& al. 2017). Despite the recent resurrection of Hypoxylaceae DC, Rosellinia

remains assigned to Xylariaceae (Wendt & al. 2018, Daranagama & al. 2018,

Wijayawardene & al. 2018). Rosellinia is a species rich genus, especially in

tropical and subtropical areas (Petrini 1992, 2003, 2013a). In 2013, 142 Rosellinia

species were accepted based on the morphological studies of numerous

specimens (including many type specimens) from different parts of the world

(Petrini 2013a). Currently, 41 species of Rosellinia have been reported from

China (Teng 1963, Tai 1979, Ju & Rogers 1990, 1999; Yuan & Zhao 1993, Lu &

al. 2000, Liu & al. 2010, Petrini 2013a, b; Li & Guo 2015, 2016, 2018; Li & al.

2015, Su & al. 2016). The taxonomic concept of Rosellinia and the terminology

applied in this paper follow Petrini (2013a).

During our investigation of Xylariales in China, two new species and one

new record of Rosellinia were found from Guizhou province. In addition, newly

obtained DNA sequence data are reported in this paper.

Materials & methods

Collection & isolation

Fresh materials collected in southwestern China were brought to the laboratory

in paper bags. Macroscopic characters were examined under an Olympus SZ61

stereomicroscope and photographed with a fitted Canon 700D digital camera

(Senanayake & al. 2015). Materials were mounted in water and Melzer’s reagent for

anatomical examination; asci, ascospores, and ascus apical rings were photographed

using a Nikon digital camera fitted to a light microscope. More than 30 ascospores,

20 asci, and 20 apical rings were measured for each specimen. Pure colonies were

obtained by single-spore isolation following Chomnunti & al. (2014), and successful

cultures were preserved in 2 ml screw cap centrifuge tubes with 10% glycerol at —-20 °C

and with sterile water at 4 °C. Herbarium materials were deposited at the herbarium

of Guizhou Medical University, Guizhou, China (GMB) and the Herbarium of

Cryptogams, Kunming Institute of Botany, Academia Sinica, Yunnan, China

(KUN-HKAS). Cultures were deposited at Guizhou Medical University Culture

Collection (GMBC).

DNA extraction, PCR amplification, sequencing

Colonies were transferred to 2% potato-dextrose agar (PDA) medium and

incubated at 25 °C until the hyphae filled the plate. Fresh mycelia were scraped off

the medium with a clean scalpel blade. Total genomic DNA was extracted from fresh

mycelia using a Biomiga GD2416 Fungus Genomic DNA Extraction Kit (Su & al.

2016). DNA products were preserved at —20 °C.

ITS rDNA regions were amplified with primer pairs ITS4 and ITS5; a-actin gene

(ACT) sequence fragments were amplified with primer pairs ACT512F and ACT783R

Rosellinia pervariabilis & R. tetrastigmae spp. nov. (China) ... 185

TABLE 1. Taxa of Rosellinia and related genera used in the molecular analyses.

TAXON

Amphilogia gyrosa

Annulohypoxylon nitens

A. squamulosum

Astrocystis bambusae

A. mirabilis

A. sublimbata

Biscogniauxia anceps

B. arima

B. capnodes

B. mediterranea

B. simplicior

Collodiscula leigongshanensis

C. fangjingshanensis

Daldinia caldariorum

D. childiae

Durotheca comedenS

D. depressa

Hypoxylon rubiginosum

H. shearii var. minor

Jackrogersella cohaerens

Kretzschmaria clavus

K. guyanensis

K. lucidula

K. megalospora

Nemania abortiva

N. illita

N. primolutea

N. serpens

Podosordaria muli

Poronia pileiformis

Rosellinia aquila

R. buxi

R. caudata

R. convexa

R. corticium

R. lamprostoma

CULTURE NUMBER

YMJ 91123101

YMJ 91022108

YMJ 90081905

89021904 (HAST)

94070803 (HAST)

89032207 (HAST)

YMJ 123 [T]

YMJ 122

YMJ 138

YMJ 147

YMJ 136

GZUH0107 [T]

GZUHO0109 [T]

YMJ 263

CBS 122881

BCC25152

BCC28073

YMJ 24

YMJ 29

YMJ 310

YMJ 114

89062903 (HAST)

YMJ 112

YMJ 229

467 (BISH) [T]

YMJ 236 [T]

YMJ 91102001[T]

235 (HAST)

167 (WSP) [T]

88113001 (WSP)[T]

MUCL 51703

99 (JDR)

GMBC0145

GMBC0207

GMBC0208

GZUCC13005 [T]

MUCL 51693

YMJ 89112602

GENBANK ACC. NO.

a-actin B-tubulin

EF025600 EF025615

AY951772 AY951663

AY951774 AY951665

GQ449239 GQ495942

GQ449238 GQ495941

GQ449236 GQ495940

AY951783 AY951671

AY951784 AY951672

AY951787 AY951675

AY951796 AY951684

AY951798 AY951686

— KRO002587

— KRO002589

AY951802 AY951690

KU684039 KU684129

GQ160480 GQ160488

GQ160484 GQ160492

AY951862 AY951751

AY951864 AY951753

AY951766 AY951655

EF025596 EF025611

GQ408901 GQ478214

EF025595 EF025610

EF025594 EF025609

GQ374123 GQ470219

EF025593 EF025608

EF025592 EF025607

GQ389695 GQ470223

GQ455450 GQ844839

GQ455449 GQ502720

— KX271253

GQ398228 GQ470228

MH771031 #MH778499

MH771029 MH778497

MH771030 MH778498

KF885726 KF885725

— KX271254

EF025589 EF025604

REFERENCE

Hsieh & al. 2010

Hsieh & al. 2005

Hsieh & al. 2010

Hsieh & al. 2005

Li & al. 2015

Hsieh & al. 2005

URen & al. 2016

Leessoe & al. 2013

Hsieh & al. 2005

Hsieh & al. 2010

Wendt & al.2018

Hsieh & al. 2010

This study

Su & al. 2016

Wendt & al. 2018

Hsieh & al. 2010

186 ... Xie & al.

R. necatrix YMJ 89062904 EF025588 EF025603 Hsieh & al. 2010

R. pervariabilis GMBC0092 [T] MH771027 MH778495 This study

R. sancta-cruciana 90072903 (HAST) GQ389699 GQ470227 Hsieh & al. 2010

R. tetrastigmae GMBC0030 [T] MH771028 MH778496 This study

Stilbohypoxylon elaeidicola YMJ 173 EF025601 EF025616 Hsieh & al. 2010

S. quisquiliarum YMJ 89091608 EF025591 EF025606 Hsieh & al. 2010

Theissenia pyrenocrata TC11480 GQ247716 GQ247717 Leessoe & al. 2013

Whalleya microplaca YMJ 91111215 EF025599 EF025614 Hsieh & al. 2010

Xylaria acuminatilongissima 623 (HAST) [T] GQ853046 GQ502711 Hsieh & al. 2010

X. brunneovinosa 720 (HAST) [T] GQ853041 GQ502706 Hsieh & al. 2010

Note: New sequences in bold; ex-type sequences designated by ‘[T]’

(Carbone & Kohn 1999); the 6-tubulin gene (BT) was amplified using primer pairs

T11 and T1 (Tanaka & al. 2009; Hsieh & al. 2010). Amplification conditions followed

Li & al. (2015) and Su & al. (2016). After DNA was amplified using the polymerase

chain reaction (PCR), the PCR products were sent to Sino Geno Max in Beijing for

DNA sequencing. All nucleotide sequences obtained were uploaded into GenBank,

and the accession numbers of DNA sequences used in this paper are listed in TABLE 1.

Sequence alignment and phylogenetic analyses

Preliminary BLAST searches of ITS rDNA sequences placed our sequences close

to those of Rosellinia spp. Sequences used for constructing phylogenetic tree in this

paper were chosen according to the latest authoritatively published papers and the

results of preliminary BLAST searches (Su & al. 2016, Wendt & al. 2018, Daranagama

& al. 2018).

Sequences were downloaded and aligned using BioEdit (Hall 1999). The

combined dataset was concatenated from individual ACT and BT alignments, with

alignments checked and improved manually where appropriate and uploaded in

TreeBASE (https://treebase.org/treebase-web/home.html). We used an online tool

(http://sing.ei.uvigo.es/ALTER/) to convert files for RAxML and MrBayes (Ronquist

& al. 2012). The NEXUS file was modified with MrModeltest v. 2.2 (Nylander 2004).

Randomized accelerated maximum likelihood (RAxML) was conducted using

RAXML online (https://www.phylo.org/portal2/). Parameters for RAxML followed

Phookamsak & al. (2015). Bayesian analysis was performed with MrBayes v.3.1.2

(Ronquist & al. 2012) via a uniform [GTR+I+G] model (Iset nst = 6, rates = invgamma,

Prsetstatefreqpr = dirichlet, 1, 1, 1, 1) inferred by MrModeltest 2.3. The phylograms

were visualized in Figtree 1.4.3 and polished in Adobe Photoshop version CS6 (Fic. 1).

Results

Three species of Rosellinia, two new species and a new record for the Chinese

mainland, were recorded.

Rosellinia pervariabilis & R. tetrastigmae spp. nov. (China) ... 187

0.89/94, Rosellinia caudata GMBC0207

1100} Rosellinia caudata GMBC0208

0.99/94 L Rosellinia caudata GMBC0145

uss Rosellinia corticium MUCL 51693

ooiee Rosellinia merrillii 89112601 HAST

1/100 Rosellinia aquila MUCL 51703 Rosellinia

Rosellinia convexa GZUCC 13005 [T]

0.97/88 Rosellinia tetrastigmae GMBC 0030 [T]

1/100 Rosellinia pervariabilis GMBC0092 [T]

1/100 Rosellinia lamprostoma YMJ 89112602

Rosellinia sancta-cruciana 90072903 HAST

Rosellinia buxi 99 JDR

Rosellinia necatrix YMJ 89062904

woo -— Kretzschmaria clavus YMJ 114

1/100 Kretzschmaria lucidula YMJ 112

Kretzschmaria megalospora YMJ 229 Kretzschmaria

1/97 "96l__ Kretzschmaria guyanensis 89062903 HAST

0.99/94 Nemania abortiva 467 BISH [T]

1/100 Nemania primolutea YMJ 91102001 [T]

Nemania illita YMJ 236 [T]

idan Astrocystis mirabilis 94070803 HAST Astrocystis

-/67. Astrocystis bambusae 89021904 HAST

-/65 Astrocystis sublimbata 89032207 HAST

inog7 Collodiscula leigongshanensis GZUH0107 [T]

‘| ollodiscula fangjingshanensis GZUHO109 ‘ Collodiscula

16:

0.99/81

-/63 5

Nemania

Stilbohypoxylon quisquiliarum YMJ 89091608

Stilbohypoxylon elaeidicola YMJ 173 | si ohypoxylon

ch wiod Xylaria brunneovinosa 720 HAST [T] i

Xylaria acuminatilongissima 623 HAST | Xylaria

b 99/48 Poronia pileiformis 88113001 WSP [T] Poronia

Podosordaria muli 167 WSP [T] Podosordaria

0.93/7 Annulohypoxylon nitens YMJ 91022108

0.98/72 1/99 Annulohypoxylon squamulosum YMJ 90081905 | Annulohypoxylon

{51 Jackrogersella cohaerens YMJ 310 Jackrogersella

1/100) Daldinia caldariorum YMJ 263 Daldinia

Daldinia childiae CBS 122881

9 [— Durotheca depressa BCC28073

0.99/79 1/100 ‘Durotheca comedens BCC25152

Theissenia pyrenocrataTC11480 Theissenia

| Durotheca

-/65

6 Hypoxylon rubiginosum YMJ 24

; ‘Hypoxylon shearii var. minor YMJ 29 | Hypoxylon

veel Biscogniauxia anceps YMJ 123 [T]

0.96/62 Biscogniauxia capnodes ¥YMJ 138

199 Biscogniauxia mediterranea YMSJ 147 Biscogniauxia

‘Biscogniauxia arima YMJ 122

Biscogniauxia simplicior YMSJ 136

Whalleya microplaca YMSJ 91111215

Amphilogia gyrosa YMJ 91123101 [T]

Fic. 1. RAxML tree for Rosellinia and related genera based on a combined dataset of 6-tubulin

and a-actin gene segment sequences. Bayesian posterior probabilities >0.95 and bootstrap

support values for maximum likelihood (ML) higher than >50% are marked above the nodes;

an en-dash (“—”) indicates a value <0.95 (PP) or <50% (BS). Strain numbers are noted after the

species names. Our new sequences are shown in bold; and ex-type sequences are annotated with

‘[T]. The tree is rooted via the outgroup Amphilogia gyrosa YMJ 91123101.

188 ... Xie & al.

Phylogenetic analyses

Phylogenetic trees were generated with the RAxML and MrBayes

methods by using combined datasets of BT and ACT sequences. Forty-

seven fungal strains representing 15 different genera were analysed. After

manual adjustment, the alignments consisted of 1130 character positions

in the BT alignment and 334 in the ACT alignment, of which 92 characters

were variable, 715 constant, and 657 parsimony informative. Two different

molecular markers were used, and two tree reconstruction methods were

applied.

The result of the phylogenetic analysis is shown in Fic. 1 with the final

ML optimization likelihood value of -21722.880899. Our isolates fell within

genus Rosellinia. Our strains formed separate branches on the phylogenetic

tree obtained via most parsimonious analysis of the ITS rRNA gene dataset

(supplementary material). Rosellinia tetrastigmae and R. pervariabilis showed

a close relationship with R. lamprostoma Syd. & P. Syd., with R. pervariabilis

and R. tetrastigmae supported as independent species. Our R. caudata three

specimens clustered together with R. corticium with high bootstrap values

(0.99/96).

Taxonomy

Rosellinia pervariabilis Q.R. Li & J.C. Kang, sp. nov. Fic. 2

MycoBAnk MB 827512

Differs from Rosellinia culmicola by its smaller ascospores and the absence of a germ slit.

TypE—China, Guizhou Province: Qiannan Buyei & Miao Autonomous Prefecture,

Maolan National Nature Reserve, on the stem of Bambusa pervariabilis McClure, VII

2016, Li 16029 (Holotype, GMB 0092; isotype, HKAS 101452; ex-type living culture,

GMBC 0092).

EryMoLoGy—In reference to the host, Bambusa pervariabilis.

SAPROBIC on stem of B. pervariabilis. SEXUAL MORPH—SUBICULUM

composed of finely interwoven hypha, brown to reddish brown, persistent.

STROMATA scattered to gregarious, solitary, superficial, subglobose to

conical, dark, 0.3-0.5 mm diam., 0.4-0.8 mm high, each containing a single

perithecium. OsTIOLEs finely papillate. EcrostRoma 20-40 um thick, black,

carbonaceous. ENTOSTROMA confined to base, black. PERITHECIA globose

to subglobose. PARAPHYSES hyaline, septate. Asc1 90.5-138 x 9.5-15 um,

8-spored, cylindrical, unitunicate, short-pedicellate, with a J+ (staining blue

in iodine), long barrel-shaped apical ring, 7.5-9 um high, 3-4.5 um diam.

ASCOSPORES 19.5-24.5 x 4-5 um, overlapping uniseriate, botuliform, ends

Rosellinia pervariabilis & R. tetrastigmae spp. nov. (China) ... 189

Fic. 2. Rosellinia pervariabilis (holotype, GMB 0092). A-E. Stromata on the host. F. Section of

stroma. G-I. Asci. J. Long barrel-shaped J+ apical ring. K, L. Ascospores. Scale bars: A, B = 0.2 cm;

C, D = 500 um; E, F = 200 um; G-L = 10 um.

rounded, brown to yellowish brown, unicellular, smooth, lacking sheath and

germ slit. ASEXUAL MORPH: unknown.

190 ... Xie & al.

CoMMENTS: Rosellinia pervariabilis is diagnosed by its uniquely shaped

ascospores and lack of a germ slit. Although there are many Rosellinia species

producing ascospores within the 15-25 um range (e.g., R. abscondita Rehm,

R. helvetica L.E. Petrini & al., R. hyalospora Theiss), their ascospores have a

much lower (<4) length : width ratio (Petrini 1992, 2003, 2013a). Rosellinia

culmicola has larger ascospores (24.2+1.5 x 5.9+0.6 um) and a 6-7 um long

germ slit; moreover, it has a stromatal ring (Petrini 2013a).

Rosellinia tetrastigmae Q.R. Li & J.C. Kang, sp. nov. FIG. 3

MycoBAnk MB 828157

Differs from Rosellinia megalosperma by its smaller stromata; and from R. procera by its

smaller stromata and its ascospores having a germ slit.

TypE—China, Guizhou Province: Qiannan Buyei & Miao Autonomous Prefecture,

Maolan National Nature Reserve, on the stem of Tetrastigma sp. (Vitaceae), VII 2016,

Li 16012 (Holotype, GMB 0030; isotype, HKAS 101457; ex-type living culture, GMBC

0030).

EryMoLoGy—In reference to the host genus, Tetrastigma.

SAPROBIC on the twigs of dicot wood. SEXUAL MORPH—SUBICULUM

composed of finely interwoven hypha, black, persistent. STROMATA scattered

or gregarious, solitary, superficial, cupulate to oblate sphere, 0.4-0.6 mm

diam., 0.3-0.5 mm high, each containing a single perithecium, embedded in

the subiculum slightly. OsTIoLEs finely papillate or integrated. EcrosTROMA

25-50 um thick, black, carbonaceous. ENTrostroma black, confined to

stroma base. PERITHECIA subglobose, not collapsed. PARAPHYSES hyaline,

septate. Ascr 185-240 x 32-45 um, 8-spored, cylindrical, overlapping

biseriate, unitunicate, short-pedicellate, with a J+, barrel-shaped apical ring,

16-22.5 um high, 8.5-11.5 um diam. Ascospores 72.5-111.5 x 12.5-19.5

um, asymmetrically ellipsoidal with broadly round ends, yellowish brown to

dark brown, smooth, surrounded at each end by slimy caps, with a straight

germ slit extending the length of the spore on the flat side. ASEXUAL MORPH:

unknown.

ADDITIONAL MATERIAL EXAMINED—CHINA, GUIZHOU PROVINCE, Qiannan Buyei

& Miao Autonomous Prefecture, Maolan National Nature Reserve, on the twigs of

unknown plant, VIII 2017, Li 17103 (GMB 0212).

CoMMENTS: Rosellinia tetrastigmae resembles R. procera Syd. & P. Syd. and R.

megalosperma Syd. & P. Syd. San Martin & Roger (1995). Although Ju & Roger

(1999) regarded R. procera and R. megalosperma as synonyms, Petrini (2013a)

treated them as independent species based on the presence of ascospore

germ slits in R. megalosperma and their absence in R. procera. Rosellinia

Rosellinia pervariabilis & R. tetrastigmae spp. nov. (China) ... 191

Fic. 3. Rosellinia tetrastigmae (holotype, GMB 0030). A, B. Stromata on the host. C. Section of

stroma. D-F. Asci. G. Barrel-shaped J+ apical ring. H, I. Ascospores. Scale bars: A = 0.5 cm;

B = 200 um; C= 100 um; D-I = 20 um.

tetrastigmae differs from both by its smaller stromata (R. megalosperma

—600-1000 um high, 750-1100 um wide; R. procera—825-975 um high,

550-875 um wide; Petrini 2013a).

Rosellinia caudata Petch, Ann. Roy. Bot. Gard. (Peradeniya) 10: 135 (1926) Fie. 4

SAPROBIC on the twigs of dicot wood, forming on the host surface. SEXUAL

MORPH—SUBICULUM composed of finely interwoven hypha, well developed,

brown to red brown, persistent. STROMATA scattered or densely gregarious,

192 ... Xie & al.

superficial, subglobose to cupulate, dark brown, 0.8-1.2 mm diam., 0.7-1 mm

high, containing single perithecia. OsT1oxEs finely papillate. EcrostromaA

50-110 um thick, black, carbonaceous. ENTOSTROMA confined to base, black.

PERITHECIA subglobose to cupulate. PARAPHysSES hyaline, septate. Ascr

165-245 x 7.5-15.5 um, 8-spored, cylindrical, unitunicate, short-pedicellate,

with a J+ (staining blue in iodine), long barrel-shaped apical ring, 6-13.5

um high, 5.5-7.5 um diam. Ascospores 21.5-30.5 x 7-11 um, slightly

overlapping uniseriate, asymmetrically ellipsoidal with broadly rounded

ends, cellular appendage at one end in mature ascospores, brown to dark

brown, unicellular, smooth, with a slimy sheath, on convex side with straight

germ slit as long as spore. ASEXUAL MORPH: unknown.

MATERIAL EXAMINED—CHINA, GUIZHOU PROVINCE, Qiannan Buyei & Miao

Autonomous Prefecture, Maolan National Nature Reserve, on twigs of unknown

plant, VII 2016, Li 16102 (GMB 0145; living culture, GMBC 0145); Guiyang, Guizhou

Medical University Campus, on twigs of dicotyledonous wood, IV 2017, Li 17115

(GMB 0207; living culture, GMBC 0207); Huaxi Yingbinguan Hotel Park, on twigs of

dicotyledonous wood, VII 2017, Li 17126 (GMB 0208; living culture, GMBC 0208).

COMMENTSs: Our strains were identified as Rosellinia caudata, which are

morphologically similar to R. corticium (Schwein.) Sacc., R. aquila (Fr.) Ces.

& De Not., and R. merrilli Syd. & P. Syd. (Petrini 2013a). Rosellinia caudata

also shows a close affiliation with R. corticium, R. merrilli, and R. aquila

in the phylogenetic tree (Fic. 1); however, R. corticium differs by its larger

stromata, R. aquila differs by its smaller ascospores and larger stromata,

and R. merrilli differs by its larger ascospores with a cellular appendage at

each end (Petrini 2013a). Petrini (2013a) reported a specimen of R. caudata

from Taiwan, but we believe that our specimens represent a new record for

the Chinese mainland. We also report the first BT and ACT sequences for

R. caudata.

Discussion

Morphological characteristics are widely used to distinguish between

similar species of Rosellinia. Stroma shape, presence of subiculum, texture

and color, apical ring, ascospore shape and dimensions are important

diagnostic characters for Rosellinia (Petrini 2005, 2013a). Rosellinia species

with similarly sized ascospores clustered together clearly in the ITS-

based phylogeny by Su & al. (2015). Our analyses show a similar trend

(Fic. 1). However, out of more than 142 taxa very few sequences have been

uploaded to GenBank, suggesting that Rosellinia is still not yet well studied

phylogenetically. Much herbarium material is in poor condition and cannot

Rosellinia pervariabilis & R. tetrastigmae spp. nov. (China) ... 193

Fic. 4. Rosellinia caudata (GMB 0145). A-D. Stromata on the host. E. Section of stroma.

F, G Barrel-shaped J+ apical ring. H-K. Asci. L-O. Ascospores. Scale bars: A = 0.5 cm;

B-E = 200 um; F-O = 10 um.

194 ... Xie & al.

be used for sequencing. This paper provides sequences from new Rosellinia

collections, which should prove beneficial to systematic molecular taxonomy

of the Rosellinia.

Acknowledgements

The authors are grateful to Dra. D.A. Daranagama (University of Kelaniya,

Sri Lanka) and Dra. Liliane Petrini (Breganzona, Switzerland) for their highly

informative and helpful pre-submission expert reviews. This research was supported

by the Fund of High Level Innovation Talents No. 2015-4029; the Fund of Innovation

Team of Guizhou Province No. 2015-4025; the Fund of Innovated Team of the

Education Department of Guizhou Province No. 2014-31 and the Program for New

Century Excellent Talents in University No. NCET-13-0747; the Fund of Guizhou

Provincial Traditional Chinese Medicine Administration Project QZYY-2016-097;

Doctoral Starting Foundation of Guizhou Medical University YT2017-13; Guiyang

Science and Technology Planning Project No. (2017)30-19; the Open Fund Program

of Engineering Research Center of Southwest Bio-Pharmaceutical Resources,

Ministry of Education, Guizhou University No. GZUKEY20160; and Guizhou

University Talent Introduction Project No. 2017-54.

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MY COTAXON

January-March 2019—Volume 134, pp. 197-213

https://doi.org/10.5248/134.197

Catalog of Penicillium spp. causing blue mold

of bulbs, roots, and tubers

FRANK M. DUGAN" & CARL A. STRAUSBAUGH?

"USDA-ARS WRPIS, Washington State University, Pullman, WA 99164 USA

?USDA-ARS NWISRL, Kimberly, ID 83341 USA

“CORRESPONDENCE TO: fdugan@wsu.edu

ABSTRACT—Accuracy in assigning specific epithets to Penicillium isolates documented

as agents of blue mold of edible and ornamental bulb, root, and tuber crops is highly

variable—with methods ranging from appropriate (recent morpho-cultural criteria,

metabolite production, DNA sequences), to plausible (older morpho-cultural criteria

from monographs), to suspect (unspecified methods, identification via inappropriate

literature). We provide a catalogue appropriate for plausibly assigned names accompanied

by authorities, references, host distribution, and identification methodology. Names are

categorized according to (i) segregates of P. corymbiferum (i.e., names in P. subg. Penicillium)

and taxa in P. ser. Corymbifera associated with Liliaceae s.1.; (ii) taxa in P. subg. Penicillium

other than P. ser. Corymbifera associated with Liliaceae s.1.; (iii) taxa other than P. subg.

Penicillium associated with Liliaceae s.1.; (iv) associates of Beta vulgaris (beets and sugar

beets); and (v) associates of mostly tropical or subtropical roots and tubers. Ambiguities or

deficiencies in assignment of certain specific epithets are noted.

Key worps—Allium, Dioscorea, Iris, Manihot, Zingiber

Introduction

Previously attributed to a small number of Penicillium species, blue mold

pathogens on bulb and root crops have been assigned by modern molecular-

genetic methods to an increasing number of species, many of which represent

“cryptic” species segregated from other species described decades ago (Dugan

& Everhart 2016). In comparing sets of investigations, results from artificial

inoculations reveal substantial areas where host ranges agree as well as

198 ... Dugan & Strausbaugh

instances with contradictory results (Dugan & al. 2014, 2017; Overy & al.

2005). As new species are still being described and new host records published

(e.g., Strausbaugh & Dugan 2017, Oh & al. 2015, Sang & al. 2014), there is a

need for a catalog of names for Penicillium species causing blue mold diseases

of bulb and root crops that also cites the literature on hosts and indicates how

isolates were identified. Because numerous publications record Penicillium

species on various bulb, root, or tuber hosts without clearly indicating how

isolates were identified to species level, we emphasize literature with well-

defined identification methods. Species names in Penicillium are provided with

authors, dates, and publication references (Index Fungorum 2018) for species

of greatest importance to this review (TABLES 1-5). We divide our topic into

three main sections: blue mold on bulbs of Liliaceae sensu lato (TABLEs 1-3),

blue mold on beets and sugar beets (TABLE 4), and blue mold on roots and

tubers, many of which are tropical or subtropical (TABLE 5). We also note

pertinent subgeneric classifications for TABLES 1-3.

TABLES 1-5 concisely summarize methods used to assign specific epithets

for each publication cited. In some instances pertinent isolates that may have

been obtained at different times or simultaneously are those characterized

molecularly and publicly available from culture collections. For instance,

Frisvad & Samson (2004), who characterized isolates using morpho-cultural

methods (including extrolites), published in the same issue as Samson & al.

(2004), who characterized the same isolates emphasizing $-tubulin DNA

sequences; we also include isolates used by Overy & al. (2005), also previously

characterized and readily available. In other instances, plausibly correct names

were assigned earlier based on older monographs (e.g., Pitt 1980; Ramirez

1982; Raper & Thom 1968, 1949) whose methods were current at the time but

not yet molecular-genetic. We also note instances where species names have

become established in pertinent literature but for which identification methods

were not specified or are problematic. We accept subgeneric classifications

by Frisvad & Samson (2004), Houbraken & Samson (2011), Pitt (1980), or

Visagie & al. (2014). Authorities and publication references are provided at first

mention in TaBLes 1-5. We do not provide full synonymies for Penicillium

names (particularly where varieties have been elevated to species or where a

fungus has been transferred to Penicillium from another genus), but we do

reference specific instances of disagreement over synonymy.

For more complete lists of accepted species and synonyms, readers are

encouraged to refer to Farr & Rossman (2018), Frisvad & Samson (2004),

Pitt (1980, 2000), MycoBank (2018), Species Fungorum (2018), Visagie & al.

Blue Penicillium mold on bulbs, roots, and tubers: a catalog ... 199

(2014), and other literature mentioned in tables and text. The primary purpose

of this catalogue is to assess the degree of confidence one should place in

assigning a specific epithet by documenting the method(s) by which the name

was assigned.

Blue mold on bulbs of Liliaceae sensu lato

For decades, blue mold on edible or ornamental bulbs was attributed to

Penicillium corymbiferum Westling, Ark. Bot. 11(1): 92 (1911), presently

regarded as a synonym of Penicillium hirsutum (e.g., Frisvad & Samson 2004,

Pitt 1980, Species Fungorum 2018). Frisvad & Samson (2004) treated these and

the following species within P subg. Penicillium, P. sect. Viridicata (= P. sect.

Fasciculata; Houbraken & Samson 2011), P ser. Corymbifera. Penicillium sect.

Corymbifera is not strictly phylogenetic, but a traditional group of convenience

based partially on host (edible or ornamental bulbs) (Frisvad & al. 2000, Frisvad

& Samson 2004). Dugan & al. (2014) provided details on the typification of

Penicillium allii, P. corymbiferum, and P. hirsutum, focusing on repercussions

for identifying and naming the agents that rot garlic (Allium sativum) in the

field and in storage.

Literature is here summarized for results of artificial inoculation of various

edible bulbs (various cultivars or varieties of onion, Allium cepa L., and garlic,

A. sativum L.) and ornamental bulbs (ornamental onions, Allium aflatunense

B. Fedtsch. and A. stipitatum Regel; crocus, Crocus sativus L.; grass lily,

Ornithogalum umbellatum L.; daffodil, Narcissus ‘Ice Follies; Gladiolus ‘Black

Walnut’ and two unnamed cultivars; Iris xhollandica; Tulipa gesneriana L. and

Tulipa ‘Purple Prince’) (Dugan & al. 2011, 2014, 2017; Overy & al. 2005). Not

all plant taxa were challenged by each set of investigators. Species and cultivars

sometimes differed, as did inoculation and incubation details.

Blue mold on beets and sugar beets

The most frequently reported Penicillium species on Beta vulgaris L. subsp.

vulgaris (table beets, cv. group ‘Conditiva and sugar beets, cv. group ‘Altissima’)

is P. vulpinum (most often reported under its synonym, P. claviforme Bainier),

which is readily identifiable by its conspicuous coremia (Bugbee 1975, Fugate

& Campbell 2009). However, several other species have been conclusively

identified, and others are indicated in literature using ambiguous or potentially

obsolete identification methods. Blue mold of beet is not treated in Koike &

al. (2007), Sherf & MacNab (1986), or Snowdon (1992), but P vulpinum has

commonly been listed among the most important storage rot pathogens along

200 ... Dugan & Strausbaugh

with an Athelia-like sp., Botrytis cinerea Pers., and Phoma betae A.B. Frank

(Bugbee 1993, Strausbaugh & al. 2015). Penicillium vulpinum is an antagonist

of B. cinerea (Bugbee 1976). Penicillium spp. have also been acknowledged to

complicate the re-isolation of the slow growing Athelia-like sp. from sugar beet

roots (Strausbaugh & al. 2015).

Rot lesions on sugar beet roots infested by Penicillium spp. in storage are

normally associated with wounds created by harvest operations or other fungi

(Fugate & Campbell 2009, Strausbaugh & al. 2015). Other Penicillium spp.

documented as pathogenic on sugar beet roots in storage include: P. cellarum,

P. cyclopium, P. expansum, P. funiculosum, and P. polonicum (Bugbee 1975,

Bugbee & Nielsen 1978, Fugate & Campbell 2009, Strausbaugh 2018). A direct

comparison of P. cellarum, PB expansum, and P. polonicum established that

P. expansum and P. polonicum are the most virulent on sugar beet roots in long-

term storage in Idaho (Strausbaugh 2018). In historical literature on sugar beet

storage rots from the United States, P. vulpinum has been described as being

the most prevalent Penicillium sp. in most environments (Bugbee 1993, Fugate

& Campbell 2009). Sheikholeslami & al. (1998) also reported P. vulpinum on

stored sugar beet roots in Iran. However, recent reports describe P expansum

as being the most prevalent species in sugar beet storages in the Czech Republic

(Huijbregts 2013, Zahradnicek 1996) and Japan (Uchino 2001), although the

identification methods have not been detailed. In Idaho, both P expansum and

P. cellarum have recently been established as the most prevalent Penicillium

spp. in long-term sugar beet piles (Strausbaugh 2018). Bugbee (1975) described

P. variabile on stored roots in the Red River Valley (North Dakota), a fungus

now regarded as a synonym of P expansum in Farr & Rossman (2018),

Frisvad & Samson (2004), and MycoBank (2018). Penicillium variabile is not

included in the list of species accepted by Visagie & al. (2014) but is listed as an

independent species by Species Fungorum (2018). Bugbee (1975) determined

that P. variabile was not as virulent or as prevalent as P. vulpinum on sugar beet

roots. Penicillium has also been cited from storages in Belgium, the Netherlands,

and Sweden, but the species was not determined (Huijbregts & al. 2013). Some

species described as pathogenic on sugar beet in storage—such as P. cyclopium,

P. funiculosum, and P. polonicum—appear to be of minor importance.

Blue mold on edible roots and tubers

These crops, with the exception of horseradish, are primarily tropical to

subtropical. Given that current criteria for assignment of species names have in-

corporated molecular-genetic methods, and given the technological challenges

Blue Penicillium mold on bulbs, roots, and tubers: a catalog ... 201

faced by scientists in developing countries, it is unsurprising that the quality

of characterizing of Penicillium isolates from these crops varies considerably.

Accordingly, several names are not cited in the tables, but covered in notes be-

low the tables, with caveats indicated. TABLE 5 occasionally directs readers to a

specific note. Details are provided where a trail of citations ultimately support

plausible criteria and literature for assigning specific epithets. Other details

note where we were unable to locate literature with definitive criteria. There are

instances where specific epithets have been putatively (and invalidly) assigned

by citing literature that does not actually provide species descriptions.

TABLE 1: Penicillium subg. Penicillium sensu Frisvad & Samson (2004),

P. ser. Corymbifera; with comments on hosts in Liliaceae sensu lato

Penicillium albocoremium (Frisvad) Frisvad,

Integr. Mod. Taxon. Meth. Penicill. Asperg. Classif: 275 (2000).

Pathogenic on garlic*, onion, iris, and tulip; less virulent on ornamental onion (Allium stipitatum)

(Dugan & al. 2017; morpho-cultural, DNA sequence $-tubulin).

Pathogenic on onion, tulip; non-pathogenic on garlic*, gladiolus (Overy & al. 2005**; morpho-

cultural, DNA sequence $-tubulin).

Additional substrates in Frisvad & Samson (2004).

Penicillium allii Vincent & Pitt, Mycologia 81: 300 (1989).

Pathogenic on garlic, onion; moderately pathogenic on grass lily and tulip*; non-pathogenic on

crocus, daffodil, ornamental onion (Allium stipitatum), gladiolus, and iris (Dugan & al. 2014;

morpho-cultural, DNA sequence B-tubulin).

Pathogenic on garlic; non-pathogenic on ornamental onions (A. aflatunense, A. stipitatum) (Dugan

& al. 2011; morpho-cultural, DNA sequence 6-tubulin).

Pathogenic on onion, garlic; non-pathogenic on tulip’, gladiolus (Overy & al. 2005; morpho-cultural,

DNA sequence $-tubulin).

Additional substrates in Frisvad & Samson (2004).

Penicillium hirsutum Dierckx, Ann. Soc. Sci. Bruxelles 25: 89 (1901).

Pathogenic on crocus, garlic, onion, tulip, moderately pathogenic on gladiolus and iris; non-

pathogenic on daffodil, ornamental onion, and grass lily (Dugan & al. 2014; morpho-cultural,

DNA sequence $-tubulin).

Pathogenic on onion, garlic, tulip, gladiolus (Overy & al. 2005; morpho-cultural, DNA sequence

B-tubulin).

Additional substrates in Frisvad & Samson (2004).

Penicillium hordei Stolk, Antonie van Leeuwenhoek 35: 270 (1969).

Pathogenic on tulip; non-pathogenic on onion, garlic, gladiolus (Overy & al. 2005; morpho-cultural,

DNA sequence $-tubulin). However, typically on barley (Hordeum vulgare L.) or other cereals,

or soil wherein such are cultivated (Frisvad & Samson 2004), but also once isolated from seed

of Lupinus albus L. in Washington state (Alomran & al. 2013).

Penicillium radicicola Overy & Frisvad, Syst. Appl. Microbiol. 26: 633 (2003).

Pathogenic on garlic*, onion, and iris; non-pathogenic on A. stipitatum and tulip* (Dugan & al.

2017; morpho-cultural, DNA sequence B-tubulin).

Pathogenic on onion, tulip*; non-pathogenic on garlic*, gladiolus (Overy & al. 2005; morpho-

cultural, DNA sequence B-tubulin).

Additional substrates in Frisvad & Samson (2004), e.g., carrot, sometimes without comment on

pathogenicity.

202 ... Dugan & Strausbaugh

TABLE 1, CONCLUDED

Penicillium tulipae Overy & Frisvad, Syst. Appl. Microbiol. 26: 634 (2003).

Pathogenic on garlic*, onion, iris, and tulip, moderately pathogenic on crocus, daffodil, and grass

lily; non- pathogenic on ornamental onion and gladiolus* (Dugan & al. 2014; morpho-cultural,

DNA sequence $-tubulin).

Pathogenic on onion (cultivar-dependent), tulip, gladiolus*; non-pathogenic on garlic* (Overy & al.

2005; morpho-cultural, DNA sequence -tubulin).

Additional substrates in Frisvad & Samson (2004): bulbs of lily (Lilium sp.) and (without plant part

specified) beet (Beta vulgaris) and Jerusalem artichoke (Helianthus tuberosus L.).

Penicillium venetum (Frisvad) Frisvad,

Integ. Mod. Taxon. Meth. Penicill. Asper. Classif.: 275 (2000).

Pathogenic on garlic, onion (cultivar-dependent), gladiolus, grass lily, iris and tulip; non-pathogenic

on crocus and daffodil (Dugan & al. 2014; morpho-cultural, DNA sequence B-tubulin).

Pathogenic on onion, garlic tulip, gladiolus (Overy & al. 2005; morpho-cultural, DNA sequence

B-tubulin).

Additional substrates in Frisvad & Samson (2004): bulbs of Hyacinthus spp. (pathogenic), horseradish

(Armoracia rusticana G. Gaertn. & al., pathogenic, see P hirsutum, TABLE 5), and “licorice root”

(Glycyrrhiza glabra L.; pathogenicity not specified).

* Indicates differences in results between investigators,

possibly reflecting variation in isolates, inoculation, or incubation.

** Overy & al. (2005) challenged bulbs in more than one anatomical location.

Results tabulated above regard lesion formation at any location as a positive result.

TABLE 2. Penicillium subg. Penicillium sensu Frisvad & Samson (2004)

(other than P ser. Corymbifera) with comments on hosts in Liliaceae sensu lato.

Penicillium aurantiogriseum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88 (1901)

[P. ser. Viridicata in Frisvad & Samson 2004 = P sec. Fasciculata in Visagie & al. 2014]

Substrates in Frisvad & Samson (2004) include onion, garlic* (Allium spp., with pathogenicity

implied rather than stated). Non-pathogenic on garlic* in Valdez & al. (2009; morpho-cultural

via Ramirez 1982 and others).

Penicillium crustosum Thom, The Penicillia: 399 (1930) [P. ser. Camemberti].

Pathogenic on onion; moderately pathogenic on iris; non-pathogenic on Allium stipitatum and tulip

(Dugan & al. 2017; morpho-cultural, DNA sequence 6-tubulin).

Additional substrates in Frisvad & Samson (2004).

Penicillium expansum Link, Mag. Gesell. Naturf. Freunde, Berlin 3: 17 (1809) [P ser. Expansa].

Pathogenic on iris, tulip, onion (cultivar-dependent); moderately pathogenic on garlic (Dugan & al.

2017; morpho-cultural, DNA sequence $-tubulin). Symptoms illustrated (Duduk & al. 2017;

morpho-cultural, DNA sequence B-tubulin).

Additional substrates in Frisvad & Samson (2004). The species is most well known as inducing rot of

apple (Malus domestica Borkh.) fruits.

Penicillium polonicum K.W. Zaleski, Bull. Int. Acad. Polon. C1 Sci. Math., Sér. B: 445 (1927)

[P. ser. Viridicata]

Pathogenic on garlic, onion; non-pathogenic on Allium stipitatum, iris*, tulip (Dugan & al. 2017;

morpho-cultural, DNA sequence B-tubulin).

Pathogenic on garlic, onion, moderately pathogenic on iris*; non-pathogenic on crocus, daffodil,

ornamental onion, grass lily, and tulip (Dugan & al. 2014; morpho-cultural, DNA sequence

B-tubulin). Multiple cultivars of onion; symptoms illustrated (Duduk & al. 2017; morpho-

cultural, DNA sequence f-tubulin).

Additional substrates in Frisvad & Samson (2004) include Allium sp.

* Indicates differences in results between investigations,

possibly reflecting variation in isolates, inoculation, or incubation.

Blue Penicillium mold on bulbs, roots, and tubers: a catalog ... 203

TABLE 3. Penicillium, other than P subg. Penicillium sensu Frisvad & Samson (2004);

with comments on hosts in Liliaceae sensu lato.

Penicillium brasilianum Bat., Anais Soc. Biol. Pernambuco 15: 160 (1957)

[P. sec. Lanata-Divaricata, Visagie & al. 2014; but see also Pitt 1980]

Pathogenic on onion (but significantly less aggressive than the pathogenic Aspergillus awamori

Nakaz. or Fusarium oxysporum Schltdl.—presumably FE oxysporum f. sp. cepae) (Sang & al.

2014; morpho-cultural, B-tubulin DNA sequences).

Penicillium citrinum Thom, Bull. U.S. Depart. Agric., Bureau Anim. Indust. 118: 61 (1910)

[P. sec. Citrina, Pitt 1980, Visagie & al. 2014]

Pathogenic on Allium sativum (Hernandez-Anguiano 2006, using Pitt 1988).

Penicillium georgiense S.W. Peterson & B.W. Horn, Mycologia 101: 79 (2009)

[P. sec. Charlesii, Visagie & al. 2014]

Pathogenic on onion, moderately aggressive (lesions ~4-5mm at 10 days) (Oh & al. 2015: morpho-

cultural, 6-tubulin DNA sequences).

Penicillium glabrum (Wehmer) Westling, Ark. Bot. 11(1): 131 (1911).

[P. sec. Aspergilloides, Pitt 1980, Visagie & al. 2014]

Pathogenic on garlic, onion; moderately pathogenic on iris; non-pathogenic on Allium stipitatum,

tulip (Dugan & al. 2017; morpho-cultural, DNA sequence $-tubulin). Symptoms illustrated

(Duduk & al. 2017; morpho-cultural, DNA sequence B-tubulin). Typically confined to outer

layers (Varga & al. 2008).

Penicillium paraherquei S. Abe ex G. Sm., Trans. Brit. Mycol. Soc. 46: 335 (1963)

[P sec. Furcatum, Pitt, 2000; P sec. Lanata-Divaricata, Visagie & al. 2014]

Pathogenic on garlic, onion (cultivar-dependent); non-pathogenic on A. stipitatum, iris, tulip

(Dugan & al. 2017; morpho-cultural, 6-tubulin DNA sequences).

Penicillium purpureogenum Stoll [as “purpurogenum”], Beitr. Morph. Biol. Char. Penicillium: 32

(1904) [P. ser. Miniolutea, Pitt 1980; but see Visagie & al. 2014].

= Talaromyces purpureogenus (Sopp) Samson & al., Stud. Mycol. 70: 177 (2011).

Treated either as an accepted species (Pitt 1980, 2000; Visagie & al. 2014; Farr & Rossman (2018);

Species Fungorum 2018) or as a species complex requiring further study (Samson & al. 2011).

Pathogenic on onion, (Vélez-Rodriguez & Rivera- Vargas 2007; identification corroborated by CABI

Bioscience Identification Services).

Comments on Tastes 1—3

Penicillium allii-sativi Frisvad & al. (Houbraken & al. 2012; Persoonia

29: 89): although the epithet refers to the substrate, Allium sativum, from

which the type was isolated, it is not included in the tables because it is NOT

a pathogen on garlic (Houbraken & al 2012).

Penicillium frequentans Westling, identified from morpho-cultural

criteria, was isolated from tulip bulbs in Poland, and the closely related

P. spinulosum Thom has been isolated from onion bulbs in Lithuania

(Dugan & al. 2014, and sources cited there). Penicillium frequentans is

regarded as either an independent species (MycoBank 2018, Visagie & al.

2014) or a synonym of P glabrum (Pitt 1980, Farr & Rossman 2018, Species

Fungorum 2018).

204 ... Dugan & Strausbaugh

Penicillium gladioli L. McCulloch & Thom (not included in tables) “may

be extinct” ...; pathogenic to Gladiolus corms” (Frisvad & Samson 2004).

In spite of “may be extinct,” one still sees this name assigned to penicillia

reported on Gladiolus (e.g., Farr & Rossman 2018). See Pitt (1980) and Index

Fungorum (2018) for its sometimes ambiguous nomenclatural history.

Penicillium paraherquei and P. brasilianum were previously considered

synonyms of P. simplicissimum (Oudem.) Thom (Pitt 1980), and

P. paraherquei (not cited in Farr & Rossman 2018) is still so regarded at

Species Fungorum (2018). Isolates identified as P simplicissimum based

on Pitt (1980) are recorded from onion (cf. the sources and phylogenetic

summary in Dugan & al. 2017). Penicillium paraherquei and P. brasilianum

are accepted as independent species by Visagie & al. (2014) but are treated

as synonyms in MycoBank (2018).

Penicillium purpureogenum is sometimes assigned the protologue

citation La Cellule 33: 235 (1923). The correct protologue is provided in

TABLE 3.

Some names in Sumner & al. (2008) are problematic because that

publication does not explain clearly how a specific epithet was applied

to agents of blue mold of onion. There are the cases of P citrinum on

onion (a species primarily on Citrus but see Hernandez-Anguiano 2006);

P. cyclopium on onion (from a 1978 publication in Sumner & al. 2008)

is a synonym of P aurantiogriseum in Farr & Rossman (2018) and Pitt

(1980, 2000) but independent in MycoBank (2018) and Visagie & al.

(2014); P aurantiogriseum is documented on onion and garlic (see TABLE

2). Penicillium digitatum (Pers.) Sacc. is also mostly on citrus, but host-

fungus indices in Farr & Rossman (2018) cite P digitatum on Iris, in the

same broad family as onion. The sole record in Farr & Rossman (2018) of

P. funiculosum on onion is from a checklist of fungi in Pakistan, and Farr

& Rossman (2018) have no records for P oxalicum on onion. In addition,

P. discolor Frisvad & Samson, a species noted for its presence on cheeses, is

recorded from onion in two cases by Frisvad & Samson (2004), but with no

indication of pathogenicity.

TABLE 4. Penicillium spp. on Beta vulgaris.

Penicillium cellarum C.A. Strausb. & Dugan, Pl. Dis. 101: 1783 (2017).

[B sect. Fasciculata] Phylogenetically allied to: P aurantiogriseum with ITS-5.8S, B-tubulin] (Visagie

& al. 2014); P camemberti with RPB2.

Pathogenic on sugar beet roots held in long-term storage piles (Strausbaugh & Dugan 2017; morpho-

cultural, DNA sequences: B-tubulin, ITS-5.88, RPB2). PB cellarum is recently described, yet to

be evaluated on other crops.

Blue Penicillium mold on bulbs, roots, and tubers: a catalog ... 205

TABLE 4 CONCLUDED

Penicillium cyclopium Westling, Ark. Bot. 11(1): 90 (1911).

Pathogenic on sugar beet roots (Beta vulgaris) (Bugbee & Nielsen 1978; morpho-cultural in Raper

& Thom 1968).

Penicillium expansum

Pathogenic on stored sugar beet roots (Beta vulgaris) (Bugbee 1975; Strausbaugh 2018, morpho-

cultural and DNA sequences: B-tubulin, ITS-5.8S, RPB2). Bugbee (1975; by implication, using

Raper & Thom 1968) applied the name P. variabile, sometimes considered a synonym of

P. expansum as noted above. Farr & Rossman (2018) do not presently list records of P. expansum

on sugar beet.

Penicillium funiculosum Thom,

Bull. U.S. Dep. Agric., Bur. Anim. Indust. 118: 69 (1910) [P. subg. Biverticillium].

= Talaromyces funiculosus (Thom) Samson & al., Stud. Mycol. 70: 176 (2011).

Pathogenic on sugar beet roots (Beta vulgaris) (Bugbee & Nielsen 1978: morpho-cultural via Raper

& Thom 1968).

Penicillium polonicum

Pathogenic on stored sugar beet roots (Beta vulgaris) in Idaho (Strausbaugh 2018; morph-cultural,

DNA sequences: $-tubulin, ITS-5.88, RPB2).

Penicillium tulipae

On Beta vulgaris (Overy & Frisvad 2003; morpho-cultural, extrolites, DNA sequence 6-tubulin).

Penicillium tulipae was twice isolated from B. vulgaris and given IBT accession numbers; no

indication of pathogenicity provided.

Penicillium variabile Sopp, Skr. Vidensk.-Selsk.

Christiania, Math.-Naturvidensk. KJ. 11: 169 (1912) [P subg. Biverticillium].

= Talaromyces variabilis (Sopp) Samson & al., Stud. Mycol. 70: 177 (2011).

Pathogenic on sugar beet (Beta vulgaris) (Bugbee 1975, deposited accession ATCC 28703 but

without stating identification method; possibly via Raper & Thom 1968, as this was how Bugbee

identified P cyclopium and P. funiculosum above). Penicillium variabile was rare in sugar beet,

relative to P vulpinum (Bugbee 1975).

Penicillium vulpinum (Cooke & Massee) Seifert & Samson,

Adv. Penicill. Asper. System.: 144 (1986) [“1985”]

The synonym Penicillium claviforme is common in phytopathological literature. In older U.S.

literature P claviforme is the most prevalent Penicillium rotting sugar beet (Bugbee 1993).

“Clavate coremia, usually produced at 25°C on both CYA and MEA, distinguish Penicillium

claviforme from all other Penicillium species” (Pitt 1980). Bugbee (1975) presented a photo of

coremia and deposited representative accession (ATCC 28702).

Additional substrates in Frisvad & Samson (2004).

Comments on Tas e 4:

Bugbee (1975) also cited Morochkovsky (1936, in Russian) describing

additional Penicillium spp. inducing sugar beet rot: P. stoloniferum Thom

(regarded as a synonym of P. brevicompactum by Farr & Rossman 2018, Pitt

1980, and Species Fungorum 2018), P. bordzilowskii Morochk. (later regarded

as a synonym of P. cyclopium based on Samson & al. 1976), P. expansum,

206 ... Dugan & Strausbaugh

P. duclauxii Delacr., and P rubrum Stoll. We have not read Morochkovsky’s

publication and do not know how the isolates were identified.

Liebe & al. (2016) listed Penicillium paneum Frisvad based on fungal

ITS sequences present in stored beets. The ITS region alone is considered

insufficient for species assignment in Penicillium, and definitive phylogenetic

analyses use alternative or supplemental gene regions (e.g., Samson & al.

2004; Visagie & al. 2014). Although beets were sorted based on their degree

of deterioration, Liebe & al. (2016) did not describe inoculations (i.e., Koch’s

postulates) and thus did not provide a direct measure of pathogenicity.

TABLE 5: Penicillium spp. on edible roots and tubers.

Penicillium albocoremium

On Zingiber officinale Roscoe, ginger (Frisvad & Samson 2004; morpho-cultural, DNA sequence

B-tubulin). Pathogenicity is implied rather than stated.

Penicillium brevicompactum Dierckx, Ann. Soc. Sci. Bruxelles 25: 88 (1901), [P. ser. Olsonii].

Pathogenic on Zingiber officinale, ginger (Overy & Frisvad 2005; Varga & al. 2008, literature review,

citing sources; morpho-cultural, metabolite profiles).

Penicillium crustosum

On Dioscorea alata L., purple yam (Pitt 1980, citing an IMI accession), with comment on

pathogenicity. See comment on Snowdon (1992) below.

Penicillium expansum

On Dioscorea spp., yam (Frisvad & Samson 2004). On Ipomoea spp., see comment on Holmes &

Clark (2013) below.

Penicillium hirsutum

On horseradish roots (Armoracia rusticana; Frisvad & Samson 2004) but without indicating

pathogenicity; morpho-cultural, DNA sequence $-tubulin). “P hirsutum is a major cause of

loss of stored horseradish” (Beuchat 1987). However, Beuchat (1987) may have been referring

to what was subsequently described as P hirsutum var. venetum Frisvad (Frisvad & Filtenborg

1990), the basionym of P venetum, TABLE 1.

Penicillium oxalicum Currie & Thom, J. Biolog. Chem. 22: 289 (1915)

[P. sec. Lanata-Diviricata, Visagei & al. 2014].

On sweet potato (Ipomoea batatas (L.) Lam.; Paul & al. 2018, with illustrations of symptoms; methods:

morpho-cultural, DNA sequences: B-tubulin, ITS). See comment on Snowdon (1992) in notes.

On Dioscorea spp., see comment on Snowdon (1992) below. Pathogenic on cush-cush yam

(Dioscorea trifida L.f.; Ricci & al. 1979; morpho-cultural, implying use of Raper & Thom 1949

for identification).

Pathogenic on Dioscorea alata, D. cayenensis Lam., D. dumetorum (Kunth) Pax, and D. rotundata

Poir. (Adeniji 1970: “Identifications were confirmed by the Commonwealth Mycological Inst.,

Kew, England”)

Penicillium polonicum

On Chinese yam [Dioscorea batatas Decne. (= D. polystachya Turcz.)], Kim & al. (2008; morpho-

cultural, DNA sequence B-tubulin). No pathogenicity tests were performed.

Blue Penicillium mold on bulbs, roots, and tubers: a catalog ... 207

TABLE 5 CONCLUDED

Penicillium purpureogenum

FRR 1977 “from cassava [Manihot esculenta Crantz] ... Representative of species” (Pitt 1980); no

indication of pathogenicity.

Penicillium sclerotigenum T. Yamam., Sci. Rep. Hyogo Univ. Agric., Ser. 2,

Agr. Biol. 2: 69 (1955) [P. ser. Expansa].

On yam (Dioscorea spp.), tubers and products (Frisvad & Samson 2004).

On Dioscorea batatas (Kim & al. 2008), but without pathogenicity tests; methods: morpho-cultural,

DNA sequence $-tubulin). Pitt's (1980) different classification placed P. sclerotigenum in P. subg.

Furcatum but did describe the species as rotting Dioscorea spp. See comment on Snowdon

(1992) in notes.

Comments on TasLe 5:

Holmes & Clark (2013) provided species names for penicillia causing

blue mold on sweet potatoes but did not specify the identification methods.

“Penicillium spp. identified from ...isolates collected in North Carolina

packing houses: P expansum [the most common], P. bilaiae Chalab.,

P. variabile, P. rugulosum Thom, P. solitum Westling [variously treated as a

synonym of P. aurantiogriseum (Pitt 1980), an independent species (Pitt 2000,

Visagie & al. 2014), or as a separate species with P crustosum a junior synonym

(Species Fungorum 2018)], and P. viridicatum Westling [treated as a distinct

species by Pitt (1980, 2000) and Visagie & al. (2014), but as a junior synonym

of P aurantiogriseum by Species Fungorum (2018)] (Holmes & Clark 2013,

authors of species names added). The reference on Penicillium in Holmes &

Clark (2013) is Harter & al. (1918), which states, “The species of Penicillium

which we most frequently isolated ... was given to Dr. Charles Thom ... it

belongs to the expansum group”). Holmes & Clark (2013) contained a good

photo of P. expansum by Holmes, clearly showing synnemata typical of the

species in vivo. Holmes, on the faculty at North Carolina State University

(Department of Plant Pathology), was co-author on Edmunds & al. (2008)

produced by the North Carolina State University Extension and co-author

with John Pitt (Holmes & al. 1994); Holmes identified the isolates based on

Pitt (1980, 2000) (Gerald Holmes, pers. comm.). Holmes & Clark (2013) did

not provide explicit statements regarding pathogenicity of individual species.

Snowdon (1992) listed the following species on yams: P crustosum

(“very common on yams imported into the UK from Nigeria,’ but without

supplying a citation); P cyclopium (referring only to a previous section on

that fungus, not on yam); P. gladioli (“has been found on importations to

the USA from Cuba and Puerto Rico,” but without a citation); P oxalicum

(citing Adeniji 1970—see TaBLE 5—and Ricci & al. (1978), who provided

no identification methods did reference Ricci & al. 1979, see TABLE 5);

208 ... Dugan & Strausbaugh

P. sclerotigenum (citing Yamamoto & al. 1955 as well as Moura 1980, who

in turn cited Moura & al. 1976). Snowdon (1992) also cited Plumbley &

al. (1985), which does not contain information on identification methods,

but refers to Plumbley & al. (1984, similarly devoid of such methods), who

did cite earlier reports, including Ogundana & al. (1970), which stated “...

microorganisms were isolated and identified” but which otherwise provided

no methods or references by which Penicillium isolates could conceivably be

identified to species. Nonetheless, growth rate, conidial size, and teleomorph

absence detailed in Moura & al. (1976) convincingly led to P. sclerotigenum

in the synoptic key to sclerotigenic species in Pitt (1980).

Snowdon (1992) also mentions Penicillium sp. on “cocoyams (taro

[Colocasia esculenta (L.) Schott] and tannias [Xanthosoma sagittifolium (L.)

Schott]);” and Penicillium spp. on sweet potatoes without providing specific

epithets for penicillia on any of these crops. Farr & Rossman (2018) record

the following on Ipomoea: Penicillium chrysogenum Thom, P. citrinum,

P. crustosum, PB. decumbens Thom, P. expansum, P. funiculosum, P. glabrum,

P. islandicum Sopp, P. italicum Wehmer, P. oxalicum, P. pinophilum Hedgc.,

P. purpureogenum, and P. simplicissimum, but in each instance cite only a

regional checklist. It is difficult to confirm reports of Penicillium expansum

on yam (Dioscorea), and they are not cited in Farr & Rossman (2018).

For cassava (Manihot spp.), Farr & Rossman (2018) cite a checklist of

fungi in Papua New Guinea and one research article from India listing three

Penicillium species on Manihot spp. However, the Penicillium species listed

in India were assessed as non-pathogenic in artificial inoculations, and no

identification methods were provided nor can be inferred from the references

listed.

Discussion

DNA sequence analyses, especially of B-tubulin but also of RPB2 and

even ITS (for subgeneric groups), along with advances in understanding

phylogenetic significance of metabolite production, has substantially

reinforced morpho-cultural methods for identifying Penicillium isolates to

species. Subgeneric assignment changes include transferal of P. vulpinum

(TaBLE 4) from P subg. Biverticillium to P. subg. Penicillium (Pitt 2000), but

the distinctive coremia in P vulpinum consistently enabled assignment of

that specific epithet to isolates regardless of higher level classification. Note

that although Houbraken & Samson (2011) assigned Penicillium species

to only two subgenera—P. subg. Aspergilloides and P. subg. Penicillium—

Blue Penicillium mold on bulbs, roots, and tubers: a catalog ... 209

for heuristic reasons we use prior subgeneric assignments in TABLEs 1-3,

given that most identification literature uses these prior classifications. Pitt

(2000) also noted that he (Pitt 1980) “suggested that only 70-80% of isolates

even from common sources are readily identifiable. The remainder can be

identified, but their identification must increasingly rely on the skill and

experience of the taxonomist and/or newer techniques such as metabolite

profiles or molecular data.” This comment explains the difference between

the large strides in identifying agents of blue mold of edible and ornamental

bulbs (primarily grown in economically and _ scientifically advanced

temperate regions) and the somewhat lagging identification of agents of

blue mold on tropical and subtropical roots and tubers (primarily grown in

tropical regions with developing economies).

There are repeated instances in older phytopathological literature

(from both advanced and developing economies) of cavalier treatment

of identification. Much older and even some newer literature substituted

repetitious “citation” recycling instead of explicitly specifying the

monograph or species description on which the name ultimately rested.

Through diligence, some confusions can be resolved by tracing a citation

trail back to an early monograph (e.g., Raper & Thom 1949), but ultimately

modern morpho-cultural observations and DNA sequence analyses must

be provided.

Several online publications from developing regions cite various editions

of H.L. Barnett & B.B. Hunter (ILLUSTRATED GENERA OF IMPERFECT FUNGI),

T. Watanabe (PICTORIAL ATLAS OF SEED AND SOIL FUNGI), and/or EM.

Dugan (THE IDENTIFICATION OF FUNGI) for determining species names for

Penicillium isolates derived from roots, tubers, or bulbs. Such manuals are

useful for recognizing fungal families and genera, but as they are entirely

inappropriate for applying specific epithets to Penicillium isolates, we omit

literature citing those manuals for assigning species names. We expect

inappropriate citations will diminish as comprehensive monographs on

Penicillium—as well as equipment and skills needed for DNA sequence

analyses—become more widely available. We accordingly anticipate

increased knowledge of agents of Penicillium blue mold of crops such as

cassava, sweetpotato, ginger, taro, and tannia.

Acknowledgements

The authors thank Steven Koike and Wojciech Janisiewicz for constructive

comments on the manuscript, and Shaun Pennycook for support on nomenclature.

210 ... Dugan & Strausbaugh

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NAKARIN SUWANNARACH”’, JATURONG KUMLA’’, KANITTA SATIENPERAKUL?,

WITCHAPHART SUNGPALEE‘’, SUTHEERA HERMHUK’‘*, PIYAWAN SUTTIPRAPAN>,

KRIANGSAK SRI- NGERNYUANG*, SAISAMORN LUMYONG?”*

"Department of Biology &

? Center of Excellent in Microbial Diversity and Sustainable Utilization,

Chiang Mai University, Chiang Mai, 50200, Thailand

Faculty of Economics, *Faculty of Agricultural Production, and

° Faculty of Architecture and Environmental Design,

Maejo University, Chiang Mai, 50290, Thailand

° Department of Entomology and Plant Pathology, Faculty of Agriculture,

Chiang Mai University, Chiang Mai, 50200, Thailand

* CORRESPONDENCE TO: saisamorn.l@cmu.ac.th

AxBsTRACT—Specimens of a clavarioid basidiolichen collected in Thailand were identified

as Sulzbacheromyces yunnanensis, based on morphology and similarities of ITS genes.

A full description and illustration are provided. ‘This is the first record of S. yunnanensis

from Thailand

Key worps—Basidiomycota, Lepidostromataceae, Lepidostromatales, Southeast Asia,

taxonomy

Introduction

Sulzbacheromyces (Lepidostromataceae, Lepidostromatales) was described

by Hodkinson & al. (2014) with S. caatingae (Sulzbacher & Lticking) B.P.

Hodk. & Liicking as the type species. This genus is distributed in Africa, Asia

and the neotropics. Eight species of Sulzbacheromyces have been described:

S. bicolor D. Liu & al., S. caatingae, S. chocoensis Coca & al., S. fossicola

(Corner) D. Liu & Li S. Wang, S. miomboensis De Kesel & Ertz, S. sinensis

216 ... Suwannarach & al.

(R.H. Petersen & M. Zang) D. Liu & LiS. Wang, S. tutunendo Coca & al., and

S. yunnanensis. Sulzbacheromyces is characterized by clavarioid basidiomata

combined with a lichenized crustose basal thallus with a chlorococcoid

photobiont (Hodkinson & al. 2014, Sulzbacher & al. 2016, Liu & al. 2017,

Coca & al. 2018).

Currently, only one Sulzbacheromyces species, S. fossicola has been reported

from Thailand (Liu & al. 2017). During a survey of macrofungi collected

in northeastern Thailand, we found specimens that corresponded to the

description of S. yunnanensis, a species previously reported from China (Liu

& al. 2017). Here we describe and illustrate the morphological characteristics

of the Thai material and provide evidence of ITS gene sequence matches in

GenBank.

Materials & methods

Morphology

Basidiomata were collected from the forest, Sisaket Province, Thailand, and kept

in plastic collection boxes until transported back to the laboratory. Macroscopic

descriptions of specimens were based on examination of fresh basidiomata.

Ecological observations were recorded and photographs were taken in the field.

Color names and codes follow Kornerup & Wanscher (1978). The specimens were

dried at 40—45 °C in an electric food dryer. The micromorphological data were

derived from dried specimens mounted in 95% ethanol and rehydrated in distilled

water, 3% KOH, or Melzer’s reagent. Dried specimens were deposited in the

herbarium of Research Laboratory for Excellence in Sustainable Development of

Biological Resources, Faculty of Science, Chiang Mai University, Thailand (SDBR-

CMU).

DNA isolation, amplification, analyses

Genomic DNA was extracted from fresh material using a Favorgen DNA

Extraction Mini Kit following the manufacturer's instructions. The internal

transcribed spacer (ITS) region of ribosomal DNA (rDNA) was amplified by

polymerase chain reaction (PCR) using ITS4 and ITS5 primers under the following

thermal conditions: 94 °C for 2 min; 35 cycles of 95 °C for 30 s, 52 °C for 30 s,

and 72 °C for 1 min; and 72 °C for 10 min. Negative controls lacking fungal

DNA were run for checking any contamination of reagents. PCR products were

checked on 1% agarose gels stained with ethidium bromide under UV light and

purified using a Macherey-Nagel NucleoSpin Gel and PCR Clean-up Kit following

the manufacturer’s protocol. The purified PCR products were directly sequenced.

Sanger sequencing was performed by 1°' Base Company (Kembangan, Malaysia)

using the PCR primers cited above. Sequences were used to query GenBank

database via BLAST (http://blast.ddbj.nig.ac.jp/top-e.html).

Sulzbacheromyces yunnanensis new for Thailand ... 217

ake

Fic. 1. Sulzbacheromyces yunnanensis (SDBR-CMU-NK0329).

A: basidiomata; B: basidiospores; C: basidia. Scale bars: A = 10 mm; B, C = 10 um.

Sequence analysis

The ITS sequences of our Thai specimens were deposited in GenBank

database as MK085985 and MK086016. Our ITS sequences were 100% similar

to the S. yunnanensis KUN 14-44123 holotype sequence NR159026.

Taxonomy

Sulzbacheromyces yunnanensis D. Lui, Li S. Wang & Goffinet,

Mycologia 109: 742 (2017). FIG. 1

THALLUS crustose, covering an area of 0.5-50 mm diam., distinct and

dark green when growing in shade or depressions, but yellow green and

indistinguishable from soil when growing on exposed ground; forming a thin

layer on the substrate, containing clusters of single-celled chlorococcoid algae,

without prothallus. PHotosiont chlorococcoid, 4-10 um diam., globose

to subglobose, smooth, contiguous, surrounded by a single layer of hyaline

hyphae. Basip1oMa solid, clavarioid or cylindrical, fusiform or heliciform,

simple (rarely once branched), with two conspicuous wide longitudinal

218 ... Suwannarach & al.

depressions or grooves, never circled by transverse cracks at maturity, apex

obtuse to narrowly obtuse to truncate, 25-50 x 0.5-2.5 mm, with hymenium

tissue covering the upper part; surface pruinose, without tomentum or mycelial

patch at the base, orange-yellow (4A7) or light orange (6A4) when exposed to

rain and direct sunshine, orange (6A6) in shaded habitat, base dark ochraceous,

turning ochraceous overall upon drying. Trama hyphae 2-7 um diam., parallel

aligned, clamped, slightly thick-walled, densely agglutinated, with a bulbous

apex. Hymenium 70-80 um thick.

Basidiospores 8.5—12 x 3.5—5.8 um (n = 50), Q = 2.0—2.4, ellipsoid to slightly

reniform, thin-walled, hyaline, smooth, sometimes guttulate. Basidia 50—75 x

5—6.5 um, oblong when young, subclavate to clavate when mature, thin-walled,

hyaline, 4-spored, with basal clamp connections and sterigmata <5 um long.

SPECIMENS EXAMINED — THAILAND, SISAKET PROVINCE, Khun Han District,

14°52’20”N 104°38’01”E, elevation 208 m, on soil, 2 June 2018, Suwannarach N. &

Kumla J. (SDBR-CMU-NK0329; GenBank MK085985); 3 June 2018, Suwannarach N.

(SDBR-CMU-NK0342, GenBank MK086016).

Discussion

The clavarioid basidiomata, lichenized crustose basal thallus, and

chlorococcoid photobiont support placement of our collections in

Sulzbacheromyces (Hodkinson & al. 2014). The two specimens collected in

northeastern Thailand were morphologically identified as S. yunnanensis

based on the descriptions and keys in Liu & al. (2017). The orange-yellow

to orange basidiomata of S. yunnanensis are similar to those of S. caatingae,

S. chocoensis, S. miomboensis, S. sinensis, and S. tutunendo. However, the longer

basidia of S. yunnanensis (50-75 um) clearly distinguish this species from

S. caatingae (23-45 um), S. chocoensis (25-40 um), S. miomboensis (21.1—30.3

um), S. sinensis (15-50 um), and S. tutunendo (25-40 um) (Sulzbacher &

al. 2016, Liu & al. 2017, Coca & al. 2018). The basidiospores of S. caatingae

(5—8.5 um), S. chocoensis (4m), and S. tutunendo (4 um) (Sulzbacher & al. 2016,

Coca & al. 2018) are clearly shorter than those of S. yunnanensis (8.5—12 um).

The ITS sequence matches in GenBank confirmed our two Thai specimens as

S. yunnanensis, supporting the morphological identification (Coca & al.

2018).

Including our new record, four species of Sulzbacheromyces have been

reported in Asia (Liu & al. 2017): S. bicolor in China; S. fossicola in China, India,

Singapore, and Thailand; S. sinensis in China and Japan; and S. yunnanensis

in China and Thailand. Of the four other Sulzbacheromyces species, three

(S. caatingae, S. chocoensis, S. tutunendo) are known from the neotropics

Sulzbacheromyces yunnanensis new for Thailand ... 219

(Brazil and Colombia; Sulzbacher & al. 2016, Coca & al. 2018), while

S. miomboensis is known only from Africa (Democratic Republic of Congo;

Liu & al. 2017).

The present study combining morphological characteristics and sequence

data supports the identification of S. yunnanensis and a new record for

Thailand.

Acknowledgements

This work was supported by grants from Chiang Mai University. The field

collection is under the Biodiversity of Khun Han Forest Plantation, Forest Industry

Organization (FI.O), Sisaket Program. We also thank Dr. Samantha C. Karunarathna

and Dr. Eric H.C. McKenzie for their helpful comments and careful review of this

manuscript.

Literature cited

Coca LS, Licking R, Moncada B. 2018. Two new, sympatric and semi-cryptic species of

Sulzbacheromyces (lichenized Basidiomycota, Lepidostromatales) from the Choco biogeographic

region in Colombia. Bryologist 121: 297-50. https://doi.org/10.1639/0007-2745-121.3.297

Hodkinson BP, Moncada B, Lticking R. 2014. Lepidostromatales, a new order of lichenized

fungi (Basidiomycota, Agaricomycetes), with two new genera, Ertzia and Sulzbacheromyces,

and one new species, Lepidostroma winklerianum. Fungal Diversity 64: 165-179.

https://doi.org/10.1007/s13225-013-0267-0

Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. Eyre Methuen, London.

Liu D, Goffinet B, Ertz D, De Kesel A, Wang X, Hur J, Shi H, Zhang Y, Yang M, Wang L.

2017. Circumscription and phylogeny of the Lepidostromatales (lichenized Basidiomycota)

following discovery of new species from China and Africa. Mycologia 109: 730-748.

https://doi.org/10.1080/00275514.2017.1406767

Sulzbacher MA, Wartchow F, Ovrebo CL, Sousa JO, Baseia IG, Moncada B, Liicking R. 2016.

Sulzbacheromyces caatingae: notes on its systematics, morphology and distribution based on ITS

barcoding sequences. Lichenologist 48: 61-70. https://doi.org/10.10107/S00242829 15000420

MYCOTAXON

January-March 2019—Volume 134, p. 221

https://doi.org/10.5248/134.221

Mycobiota (Funga) new to the Mycotaxon website

AsBsTRACT—In February, Mycotaxon added the first annotated species distribution

list from the Indian Subcontinent (India) to our ‘web-list’ page: ‘A checklist of the

non-gilled fleshy fungi (Basidiomycota) of Kerala State, India by T.K. Arun Kumar,

Anjitha Thomas, Krishnapriya Kuniyil, Salna Nanu, and Vinjusha Nellipunath.

This brought to 131 the number of free access Fungae now available on our website:

http://www.mycotaxon.com/mycobiota/index.html

ASIA

India

T.K. ARUN KUMAR, ANJITHA THOMAS, KRISHNAPRIYA KUNIYIL, SALNA

NANU, VINJUSHA NELLIPUNATH. A checklist of the non-gilled fleshy fungi

(Basidiomycota) of Kerala State, India. 8 p.

AxsstTRACT—Kerala is a geographically unique state in India with more

than half of its area encompassed within the Western Ghats hill ranges.

The peculiar physiographic, edaphic and climatic conditions that prevail in

Kerala contribute to a rich biological diversity. A literature-based checklist

of the non-gilled, fleshy basidiomycetes of Kerala is presented herein. The

list includes 81 species (Agaricomycetes, Dacrymycetes, and Tremellomycetes)

that have been documented and published from the region thus far, excluding

polypores. The listed species belong to 39 genera in 18 families placed in 11

orders. Boletus and Lycoperdon are the genera represented by the most species.