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Illinois
Natural History Survey
Bulletin

Volume 34
1987 -— 1992

State of Illinois
Department of Energy and Natural Resources

Natural History Survey Division
Champaign, Illinois

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CONTENTS

ARTICLE 1.—MONOGRAPH OF THE NEOTROPICAL FERN GENUS POLYBOTRYA
(DRYOPTERIDACEAE). By Robbin C. Moran. November 1987. 138 pp., 55 figs., 21
RIAA OPT RRS sane sec ges AV cane eu esas UaSee-eyeNer stig saveinasceahdensuens saeuilt scsssneppetereteraedeee 1-138
Acknowledgments v, Part One: Introduction and Discussion—Materials and methods 1,
Taxonomic history of the genus 3, Ecology 6, Geography 7, Morphology and anatomy
11, Chromosome numbers 27, Cladistic analysis of the species 27, Subdivision of the
genus 31, Relationships with other groups 34. Part Two: Taxonomic Treatment—Notes
on the presentation of data 37, Description of the genus 38, Key to the species of
Polybotrya 39, Species descriptions 43, Names of uncertain application 119, Excluded
taxa 120, Literature cited 122, Taxa and distribution of Polybotrya 124, Distribution
maps 124, Index to collectors’ numbers 132, Index to taxonomic names 137.

ARTICLE 2.—ASTER AND BRACHYACTIS IN ILLINOIS. By Almut G. Jones. May 1989. 55 pp.,
MELA Pisa ce cas sats avers vaxnssnareastuscaccasbasuasspuvesostoqssstesseesesseacostencvshsutautsieensenseucsvivassieteresctaaste 139-194
Introduction 139, Generic description of Aster L. 143, Conspectus of classification of
Illinois Aster species 144, Key to species of Aster and Brachyactis in Illinois 145,
Descriptions of Aster species 149, Description of Brachyactis species 186, Glossary of
descriptive terms 187, Literature cited 190, Index to scientific species names, including
synonyms 193, Index to vernacular (common) names 194.

ARTICLE 3.—A NOMENCLATOR OF LEPTOSPHAERIA V. CESATI & G. de NOTARIS
(MYCOTA-ASCOMYCOTINA-LOCULOASCOMYCETES). By J.L. Crane and C.A.
pana eet CE Dea VURAL EE LDP ATO PIPE ances ose asecesaee sete ce oc cacccvacuacnert sn csaceatcecteneucass<csacseubesstarsse 195-355
Acknowledgments iv, Introduction and historical background 195, List of Leptosphaeria
species, varieties, and forms 198, Host index 291, Host family index 308, Substrate index
318, Geographic index 328, Appendix 1. Taxonomic division of Leptosphaeria 339,
Appendix 2. Genera historically allied to Leptosphaeria 341, Appendix 3. Synonyms of
Leptosphaeria 347, Appendix 4. Anamorphs of Leptosphaeria 348, Literature cited 351.

ARTICLE 4.—OUR LIVING HERITAGE: THE BIOLOGICAL RESOURCES OF ILLINOIS.
(Proceedings of a symposium in celebration of Earth Day 1990). Edited by Lawrence M.
Page and Michael R. Jeffords. April 1991. 120 pp., figs., maps, photographs. .....357—477
Foreword iii, Introduction 357, Session One: Forests 359, Session Two: Prairies and
Barrens 383, Session Three: Wetlands 400, Session Four: Streams and Caves 416,
Session Five: Agro-Urban Ecology 453, Appendices 463.

ARTICLE 5.—SYSTEMATICS OF LEPTOSPHAERIA SPECIES FOUND ON THE ROSACEAE.
By Sabine M. Huhndorf. May 1992. 55 pp., 21 figs. ........:ecsesesesssssesesssssesseesssesseees 479-534
Acknowledgments iv, Introduction 479, Leptosphaeria species referable to the
Pleosporales 482, Leptosphaeria species referable to the Melanommatales 505,
Leptosphaeria species referable to the Dothideales 508, Leptosphaeria species referable
to the Hymenoascomycetes 518, Species incertae sedis 525, Appendix: Leptosphaeria
species described from the Rosaceae 526, Literature cited 532, Index to taxa 534.

ARTICLE 6.—CATALOG OF TYPES OF THE ILLINOIS NATURAL HISTORY SURVEY
MYCOLOGICAL COLLECTIONS (ILLS). By J.L. Crane and Pamela P. Tazik. May
Introduction 535, Catalog of types 536, Literature cited 550, Appendix 550.

Monograph of the
Neotropical Fern Genus
Polybotrya (Dryopteridaceae)

Robbin C. Moran

Volume 34, Article 1
November 1987

Monograph of the Neotropical Fern Genus

Polybotrya (Dryopteridaceae)

Robbin C. Moran

Illinois Natural History Survey
Illinois Department of Energy and Natural Resources

Department of Plant Biology
University of Illinois at Urbana-Champaign

Illinois Natural History Survey Bulletin
Volume 34, Article |
November 1987

This work is dedicated to my parents, Livia Ann Moran and John Howard Moran.

Dr. Robbin C. Moran currently works at the Missouri Botanical Garden in St. Louis.

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

Printed by Authority of the State of Illinois
(62997—1 ,200—1 1-87)

No charge is made for most publications of the Illinois Natural History Survey, and a list of those
publications is available upon request. Single copies of most publications are available to anyone request-
ing them. Requests for multiple copies should be made in writing and should explain the use to be
made of the publications. Address correspondence to the Office of the Chief at the address below.

Illinois Natural History Survey
Natural Resources Building
607 East Peabody Drive
Champaign, Illinois 61820

Citation:
Moran, R.C. 1987. Monograph of the Neotropical Fern Genus Polybotrya (Dryopteridaceae). Illinois
Natural History Survey Bulletin 34 (1): 1—138.

US ISSN 0073-4918

Contents

SMPERELETIOINCHIS IMME EE eee a ee nn OT Oe Ge BO in i's fi se 0 .e v

Part One: Introduction and Discussion

Rete MMIC IVIC NOUS teh sate fe ls eee soos BoC ay are Wan: ievoinems mtioes | Bae) 1
SRTNCHIs(OTy Onthe Genus, 2.5. cade 5) seats secs ee St sat wo ee 3
SPP cts oc tees Se ica ot Jo; Gay top Spee yee Sayan Saas oils) a sei 6
OSS EDSITITRY 5 o Sumidees Biase emetic ts tale etiam gee ee ec em emer 7
RENEE AATICCATIACOMIY is yore are yk Gags Joho al ects oe gg. “sige bas OP sae 11
PRICED A IITMIUCIS Chest ec cee. ae! ccs) se 5) OG: ye) be cua ee ng 27
PanrewAUALYSISOLMNe SPECIES =. F . . ee os ie ee 27
PPI EHIONIEOM IDE GERUST Ass ty scrim Gn. oe cy om ackice Gr ok be gehes > Sheep cee Set A 31
SermtSMipsuwith’ Other Groups: <>... hae cn ee ee ee ee 34

Part Two: Taxonomic Treatment

Mee enarueseresentationt Ol Data: sei is) xs) cue. aieelercsy sal see, sae ee ets 37

SITE SOD OPEN CST ae ee eee eg ce TE (en eee eee ee ene 38

EM oIEIeTSDeCles OL POLYDOUYG ss oc ene 2 Gs a ee Ss Oe 39

Pa eEPAICSCTIPULOUS Meade et = hehe cogs es oe a lied ephefia. Acie” ot fomSe. AGE Le 43
Polybotrya subgenus Soromanes

leypolybotnyaserrajola (ree) Nilotzschi= -.- 295 = 2) 2) 2 sus) ae 43

2. Polybotrya polybotryoides (Baker) Christ ................ 46

SeaLopoirya suberecta, (Baker) G: Chis 5. 22 42 2 Swe he ee 50

eee OLY DOSY A CNC ACs et 12, hae ich teen os Ge eee) Dee eee) Sue 50

Polybotrya subgenus Sorbifolia

Se OLYDOIVOsSOLDiOla: KANN yin =< cere ye ee Se Bi 2 Sok oe APS 53

6G. Lolybomrya fractisertalis (Baker) J; Smith .> =: ss... 2.26: = < 54

iuoLyborya: crassirhizomaivellinger 5... Gl: 2 1s: 3 3 8 eee 2 Sts 58

8. Polybotrya espiritosantensis Brade .........5.5005564% 60
Polybotrya subgenus Polybotrya

DMEAISDOWY A COUdAA KUNZE Hh.) ae Ben A SRD SI. oe Se 60

EOL DoN.Va) govazensisiBrade, "i. ..5. a se a eee ES 66

DieeROly Dol yarpuDensiMattus. 94 «iis ees eK woke SE 68

iaeolyborrya glandulosa Kohn . . . cia. As 28 en SS 71

iSmrolyboiryailechierianaMetienius . ut... 86 2 2. eee. 71

ame colvoryaratenuata Morante... Sd ee ES Ee: 74

Ip eaiyborvarstolzer Moranits, S20: .. 2. eT Oe as. 76

ResLolporyaraipedibprdes Sere... 6.0 Ae I eee 78

iE bolybotrya botryoides (Baker)'C: Chr, 2 ee Ee. 80

iSmbolyborryalourieisianaellinper . i052 00 128) SA ee: 82

ipmbolpoiryaipiierr Mellingeir. |. 6. aah ere een ee ee ke 84

Ue Col boryarcyinarica Kaulfuss’....¢ 00 |e AO es OO 86

BisPolybotrya: mekeyMMoran 3 2)... 222d. LP AOe Lee toe 88

Doo boryopuverulenta Moran: ©... 3, ) 0. ee PS. 90

DStEE LY DOU VO\ AIA IMIOTAN My. Pp 5 veo. jo ck 6 Siw ev 65 18) odie eas 92

24. Polybotrya aequatoriana Moran. ................-...-.-. 94

25. Polybotrya appressa Motan\ 3.20) 4) ne ee ee 94
26» Polyborryaialtescandens © Chi) ene eee 97
DieRolybotrya gomezit: Morale ee) nes enn 99
28" Polybotryaosmundaced Willd) see) es eee 101
29) /Polybotryacyathifolia REG) a] = eee eis ae a ee 106
30: Polybotrya latisquamosa Moran’ = =>. = == = =.=. 2) 4 - ee 108
Sil Polybonryatsessilisora Moran. =). een ene) ee ee 108
32 eLolybotnya canalichlata owschss =a se a ee 111
33, PolybotryaysemipinnatatEeey= | ys) ee) eee 113
34: Polybotrya speciosa Schott = = = 2. 22 28.) =e = 2) eee 115
35% PolybotnyaipilosaiBrade a. 8-1) eae ee eee 117
Names of WncertainyApplication: <= ei) -neeie) ee see 119
ExclidédWaxa™. © a...) das Ble eG Sees was ees onl Sure ee ee 120
Diterature Cited) 3.5. & hc oh se a, He a 3 Ss, ey ee eee 122
axa and Distribution of: Polybotrya = 7) =) ae ee 124
Distribution:Maps: 2) sons es ss Bw Be 2 eee see See, 124
Index. to:Collectors' Numbers... 2s. «+ «2 JGRSTOESS Skee eae eee 132

Index: to Taxonomic\Names\... 28-6. .o8 6 2415 94% ee eee eee 137

Acknowledgments

No one helped me more than the five pteri-
dologists who carefully reviewed every page
of this monograph: Dr. David B. Lellinger,
Smithsonian Institution, Washington, D.C.;
Dr. Alan R. Smith, University of California,
Berkeley; Mr. Robert G. Stolze, Field Mu-
seum of Natural History, Chicago, Illinois;
Dr. Rolla M. Tryon, Harvard University,
Cambridge, Massachusetts; and Dr. Warren
H. Wagner, Jr., University of Michigan, Ann
Arbor. Not only did they spend long hours
reading my rough manuscripts, but they also
patiently answered my numerous queries
about the intricacies of fern taxonomy.

My first field trip—to Costa Rica—was
made on funds awarded by the Tinker Foun-
dation. Doctoral Dissertation Improvement
Grant # 83-06990 from the National Science
Foundation provided most of my financial
support, primarily for three additional col-
lecting trips in Latin America and for a semes-
ter of study at Harvard University.

Two people were particularly helpful dur-
ing my fieldwork in Latin America. First,
Mr. Luis D. Gomez P. of the Museo Nacional
de Costa Rica, San José, gave me lodging in
his house and extended numerous other cour-
tesies while I worked in Costa Rica. Second,
Professor Francisco Ortega, UNELLEZ,
Guanare, Venezuela, gave me the use of his
herbarium and plant-drying facilities and took
me on collecting trips through the Andes of
western Venezuela. I thank them both for
their hospitality and for sharing their know]-
edge of ferns.

During my semester of study at Harvard
University, Dr. Alice F. Tryon was extremely
helpful in showing me how to obtain high
quality scanning electron microscope photo-
graphs of spores. Dr. Rolla M. Tryon helped
with nomenclatural, systematic, and other
matters.

I learned much about pteridophyte biol-
ogy from two summer field courses with Drs.
Warren H. Wagner and Florence S. Wagner,

the first at Flathead Lake Biological Station,
Montana, and the second at Mountain Lake
Biological Field Station, Virginia. I thank
them for these memorable summers.

Dr. Kenneth R. Robertson of the Illinois
Natural History Survey, Champaign, was my
major advisor and provided much help and
encouragement. Others who served on my
doctoral committee were Dr. J. Leland
Crane, Illinois Natural History Survey,
Champaign; Drs. Johannes M. J. de Wet,
Zane B. Carothers, and Malcolm L. Sargent,
University of Illinois at Urbana-Champaign;
Dr. Rolla M. Tryon, Harvard University,
Cambridge; and Dr. Warren H. Wagner, Uni-
versity of Michigan, Ann Arbor. Dr. Tom
L. Phillips, University of Illinois at Urbana-
Champaign, provided numerous helpful com-
ments, especially on the Ecology and Geog-
raphy sections.

I completed most of the work for this
monograph in the herbarium at the Illinois
Natural History Survey, and I am greatly in-
debted to that institution for its support and
for the use of its facilities. Several biologists
at the Survey helped me in various ways. Mr.
John Taft and Ms. Mary Kay Solecki provid-
ed useful discussion and patiently tested my
keys. Dr. David Swofford was helpful with
the cladistic analysis. Mr. Bill N. McKnight
was a constant friend and critic throughout
the study and assisted with the reproduction
of the illustrations. I am also indebted to the
editorial staff at the Survey, especially Patty
Duzan and Eva Steger who typeset the man-
uscript, Audrey Hodgins who edited it, and
Molly Scott who helped with the production.

I am grateful to the directors and curators
of the following herbaria for the large quan-
tities of valuable material they made available
to me, often for a considerable period of time.
The abbreviations used in the text follow the
names of the herbaria: Armmold Arboretum,
Harvard University (A), Cambridge, Mas-
sachusetts; Herbarium Jutlandicum, Univer-

sity of Aarhus (AAU), Risskov, Denmark;
Herbarium Amazonense, Universidad Na-
cional de la Amazonia Peruana (AMAZ),
Iquitos, Peru; Botanischer Garten und
Botanisches Museum (B), Berlin, Federal
Republic of Germany; British Museum of
Natural History (BM), London, England;
Herbier (CAY), Cayenne Cedex, French
Guiana; Herbario Nacional Colombiano,
Museo de Historia Natural, Universidad Na-
cional (COL), Bogota, Colombia; Herbario
Nacional de Costa Rica, Museo Nacional
(CR), San José, Costa Rica; John G. Searle
Herbarium, Field Museum of Natural History
(F), Chicago, Illinois; Conservatoire et Jardin
botaniques de la Ville de Genéve (G), Swit-
zerland; Gray Herbarium, Harvard Univer-
sity (GH), Cambridge, Massachusetts;
Rijksherbarium (L), Leiden, Netherlands; In-
stituto Miguel Lillo de la Fundacion Miguel
Lillo (LIL), Tucuman, Argentina; Facultad
de Ciencias Naturales y Museo, Divisi6n
Plantas Vasculares, Universidad Nacional de
La Plata (LP), La Plata, Argentina; Herbario
Nacional de Bolivia, Universidad Mayor de
San Andres (LPB), La Paz, Bolivia; Her-
barium of the University of Michigan
(MICH), Ann Arbor, Michigan; Missouri
Botanical Garden (MO), St. Louis, Missouri;
New York Botanical Garden (NY), Bronx,

New York; Muséum National d’Histoire
Naturelle (P), Paris, France; Academy of
Natural Sciences of Philadelphia (PH),
Philadelphia, Pennsylvania; Herbario Uni-
versitario, Universidad Nacional Experimen-
tal de los Llanos Occidentales “Ezequiel
Zamora” (PORT), Portuguesa, Venezuela;
Herbario del Instituto de Ciencias Naturales,
Universidad Central (Q), Quito, Ecuador; In-
stituto de Ciencias, Pontificia Universidad
Catolica del Ecuador (QCA), Quito, Ecua-
dor; Jardim Botanico do Rio de Janeiro (RB),
Rio de Janeiro, Brazil; Department of Bot-
any, University of California (UC), Berke-
ley, California; United States National Her-
barium, Smithsonian Institution (US), Wash-
ington, D.C.; Herbario San Marcos, Museo
de Historia Natural, Universidad Nacional
Mayor de San Marcos de Lima (USM), Lima,
Peru; Instituto Botanico (VEN), Caracas,
Venezuela; Pringle Herbarium, University of
Vermont (VT), Burlington, Vermont; Institut
fiir systematische Botanik der Universitat
Zirich (Z), Ziirich, Switzerland.

This work was originally part of a disser-
tation submitted to the Graduate College of
the University of Illinois at Urbana~Cham-
paign in partial fulfillment of the require-
ments for the degree of Doctor of Philosophy
in plant biology.

Monograph of the Neotropical Fern Genus Polybotrya

(Dryopteridaceae)

Part One:
Introduction and Discussion

The need still is for more monographic work
on tropical species, which represent the great
majority of all ferns.

R.E. Holttum (1982)

The genus Polybotrya (Dryopteridaceae) in-
cludes 35 species of neotropical ferns. It is
distinguished by 1) strongly dimorphic leaves
with fertile leaves that resemble skeletons of
the sterile, photosynthetic ones; 2) usually
high-climbing stems that are covered with
scales; and 3) a unique stem anatomy with 5
to 12 circularly arranged meristeles, each sur-
rounded by a black sclerenchymatous sheath,
with numerous tiny leaf traces arching be-
tween adjacent meristeles. The center of di-
versity of the genus is the Andes, where 23
species occur, 12 of which are endemic. The
coastal mountains of southeastern Brazil are
notable because they contain 5 species, all
endemic. The range of Polybotrya is from
Chiapas, Mexico, southward through Central
America; the West Indies; northern South
America southward along the Andes to
Bolivia and Paraguay and eastward to the
Guiana Highlands; the Amazon River basin
and the Matto Grosso; and southeastern
Brazil. Species of the genus typically inhabit
wet, shaded, primary tropical forests from
sea level to 2500 m, most often occurring at
middle altitudes between 500 and 2000 m.
Polybotrya is divided into three subgen-
era: 1) Soromanes, leaves simply pinnate and
veins anastomosing; 2) Sorbifolia, leaves
simply or twice pinnate and veins free, close,
and parallel; and 3) Polybotrya, leaves de-
compound and veins free. Polybotrya cer-

Robbin C. Moran

vina, a species usually included in Polybot-
rya, is removed to the monotypic genus Ol-
fersia (Moran 1986). Polybotrya is related to
dryopteroid genera such as Arachniodes,
Cyclodium, Maxonia, Olfersia, and Poly-
stichopsis. Carl Christensen, the father of
modern fern taxonomy, observed (1916) that
Polybotrya may have arisen from Maxonia
because both have high-climbing stems and
strongly dimorphic leaves. The morphologi-
cal and anatomical evidence presented here
suggests that Polybotrya may have evolved
instead from a Cyclodium-like ancestor.

I chose Polybotrya for study because two
aspects of the genus immediately intrigued
me: its strongly differentiated sterile and fer-
tile leaves and its long, creeping hemiepiphy-
tic stem (Fig. 1). Since these features evolved
separately in unrelated fern genera, studying
Polybotrya might well provide insight into
broader questions of fern evolution. Polybot-
rya was suited to monographic study because
the number of species (35), all of which are
neotropical, was manageable. Finally, no
previous monographic work had been done
on Polybotrya and many problems of
nomenclature and identification remained to
be solved.

Materials and Methods

This monograph is based on the study of
about 2500 herbarium sheets, which repre-
sent approximately 800 individual collections
from 30 herbaria (see acknowledgments).
Unfortunately, Polybotrya at Kew could not
be examined because their policy precludes
loaning specimens for use by graduate stu-
dents (G.L. Lucas, in litt. 1983; pers.
comm.). Kew, however, did send color slides
of several critical type specimens.

in)

ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

FiGurE |. Habit sketches for two species of Polybotrya. Above, P. crassirhizoma, a climbing species,
below, P. sorbifolia, a terrestrial species. Note that the fertile leaves are skeletonlike and that their
orientation is more erect than that of the spreading sterile leaves.

November 1987

I spent seven months observing and col-
lecting 18 of the 35 species of Polybotrya in
Costa Rica, Ecuador, Peru, and Venezuela
and was able to make such important obser-
vations about the biology of the species as
presence of aerophores and mucilage on
stems, duration and habit of sterile versus
fertile leaves, and variation within and be-
tween populations. During field work, her-
barium, cytological, and anatomical mater-
ials were also collected.

Anatomical cross sections were prepared
by freehand sectioning, and staining was
rarely needed to determine cell types or tissue
layers. Leaf segments were cleared by soak-
ing them in 10 percent NaOH in a warm oven
for 5 to 6 days. Several changes of clearing
solution were usually needed to replace so-
lution that had become darkly stained by
phenolic substances. Since this procedure did
not remove all the dark color from the leaf,
the segments were placed in full-strength
Clorox bleach from 10 to 60 minutes. This
treatment usually rendered even the most
stubborn leaves translucent for microscopic
study.

Taxonomic History of the Genus

Polybotrya was first described by Willdenow
in his fourth edition of Linnaeus’s Species
Plantarum (1810), as distinguished by “Cap-
sules sessile, globose, aggregated in naked
paniculate spikes. Nonindusiate” [my transla-
tion]. Polybotrya osmundacea, the sole
species described in the new genus, received
its specific epithet because of the fancied re-
semblance of its fertile leaf with the fertile
leaf apex of Osmunda regalis. Since strongly
contracted, nonindusiate fertile leaves were
considered the principal feature of the new
genus, other species with similar leaves were
eventually placed in Polybotrya. Since fully
differentiated fertile leaves have arisen sepa-
rately along many phyletic lines, this proce-
dure resulted in a highly unnatural, poly-
phyletic genus. Species placed in Polybotrya
at one time and then excluded are represented
today in as many as 12 genera (see Excluded
Taxa).

MONOGRAPH of POLYBOTRA 3

Past Generic Concepts

Each of the nineteenth-century pteridologists
who wrote about Polybotrya had a slightly
different concept of the genus (Table 1).
Blume (1828) placed many unrelated acros-
tichoid ferns, including species of Loma-
gramma, Stenosemia, and Bolbitis, in Poly-
botrya. Schott (1834-1836) later restricted
Polybotrya to the American species allied to
P. osmundacea. The subsequent recognition
of the Asian acrostichoids as distinct from
the American genus Polybotrya testifies to
Schott’s keen perception. Although Schott’s
classification was not immediately adopted
by his colleagues, it was eventually revived
by Smith (1875) and Christensen (1934).

Pres] (1836) relied heavily on venation
patterns and anatomical characters in defining
his genera and did not accept Schott’s circum-
scription of Polybotrya. He merged species of
Bolbitis sect. Egenolfianae with Polybotrya
because both had free veins, but he removed
the anastomosing-veined species placed in
Polybotrya by Blume (1828).Smith (1841)
largely followed this classification in his ar-
rangement of fern genra.

Fée (1845) was the first pteridologist to
try to make sense out of the increasing
number and diversity of acrostichoid ferns.
He included the species related to P. osmun-
dacea in his subgenus Eupolybotrya and
created two subgenera of Polybotrya (see
Excluded Taxa) in which he put many species
now placed in Bolbitis, Blechnum, Lomariop-
sis, and Teratophyllum. Fée also erected the
segregate genus Soromanes for species of
Polybotrya with anastomosing veins and 1-
pinnate leaves. In my treatment, Soromanes
is a subgenus of Polybotrya.

The German pteridologist Mettenius had
a wide circumscription of Polybotrya. He de-
scribed several new species in the genus,
some of which are here retained. The major-
ity, however, have been placed in Bolbitis,
Elaphoglossum, and Leptochilus.

Hooker (1864: 194-195) and Baker in
Hooker and Baker (1874: 399ff.) subsumed
both Polybotrya and Soromanes as subgenera
in their large, eclectic genus Acrostichum,

4 ILLINOIS NATURAL HISTORY SURVEY

which consisted of any fern with acrostichoid
sori. This decision was essentially a reversion
to Swartz’s (1806) concept of Acrostichum,
and although both genera were recognized as
subgenera, Polybotrya continued to include
unrelated species that have since been placed
in Atalopteris, Elaphoglossum, Psomio-
carpa, and Teratophyllum. Hooker and
Baker’s classification was accepted by sev-
eral pteridologists, especially by flora writ-
ers, until the hegemony of Hooker’s concepts
was supplanted by the more natural views of
Smith (1875). Smith accorded generic status
to Polybotrya, restricting it to P. osmundacea
and its immediate allies; however, he kept
Soromanes as a distinct genus.
Unfortunately, Smith’s views were not
immediately accepted by pteridologists.
Christ (1897) and Diels (1899) classified

Vol. 34, Art. 1

many of the acrostichoids with free veins in
Polybotrya, a decision that again resulted in
an amalgamation of unrelated species. Chris-
tensen (1905) in Index Filicum used Polybot-
rya at the rank of genus, but it had essentially
the same circumscription as it had had under
Hooker. Unlike the writers cited above, he
placed Olfersia cervina in Polybotrya. His
classification was used by Schumann (1915)
in her important work on the vascular supply
in fertile leaves of acrostichoid ferns. Chris-
tensen (1934) later removed the discordant
species from Polybotrya and placed most of
them in the genera where they are found today
(Atalopteris, Egenolfia, Lomagramma, and
Psomiocarpa).

Copeland’s Genera Filicum (1947)
adopted Christensen’s earlier (1905) concept.
Pichi-Sermolli (1977) treated Polybotrya,

TABLE |. Comparative treatment of Polybotrya.
Polybotrya Polybotrya Polybotrya Genera formerly
subgenus subgenus subgenus included in
Reference Soromanes* _ Sorbifolia* Polybotrya* Polybotrya
Willdenow (1810) Polybotrya
Kaulfuss (1824) Polybotrya Olfersia
Blume (1828) Polybotrya Bolbitis, Lomma-
gramma, Stenosemia
Schott (1834-1836) Polybotrya
Presl (1836) Polybotrya Bolbitis
Smith (1841) Polybotrya Bolbitis
Fée (1845) Soromanes Polybotrya Blechnum, Bolbitis,
subgenus Lomariopsis,
Eupolybotrya Teratophyllum
Hooker (1864) Acrostichum Acrostichum Acrostichum Atalopteris,
subgenus subgenus subgenus Psomiocarpa,
Soromanes Polybotrya Polybotrya Teratophyllum
Smith (1875) Soromanes Polybotrya Polybotrya
Christensen (1905) Polybotrya Polybotrya Polybotrya Atalopteris, Bolbitis,
Olfersia, Psomio-
carpa, Teratophyllum
Christensen (1934) Polybotrya Polybotrya Polybotrya
Copeland (1947) Polybotrya Polybotrya Polybotrya Olfersia
Pichi-Sermolli (1977) Soromanes Polybotrya Polybotrya
Tryon & Tryon (1982) Polybotrya Polybotrya Polybotrya Olfersia

* refers to the treatment of the genus in this work.

November 1987

Soromanes, and Olfersia as distinct genera
‘but placed Soromanes and Polybotrya on
separate branches of his phylogenetic dia-
gram. Tryon and Tryon (1982) are the most
recent authors to discuss Polybotrya and re-
lated genera; their concept is like that used
in the present work, except that they include
Olfersia cervina in Polybotrya. In the present
monograph, I restrict Polybotrya to the
species allied closely to P. osmundacea,
make Soromanes a subgenus of Polybotrya,
and keep Olfersia cervina, which has been
placed in Polybotrya by many recent
pteridologists, in its own monotypic genus.
I recognize 35 species of Polybotrya, all of
which are neotropical and 10 of which are
new.

Work at the Species Level

The species of Polybotrya have received less
study over the years than those of most other
fern genera. Fée (1845), Hooker (1864), and
Hooker and Baker (1874) have been the only
monographers of the genus. Although they
attempted to identify all of the then-known
species of Polybotrya, they did not always
see types, overlooked several published
names, put some species in synonymy with-
out adequate study, and did not make detailed
observations on the genus because they were
studying hundreds of other ferns at the same
time. Despite these shortcomings, the works
of these three men have been the best source
for identifying specimens of Polybotrya in
many regions of tropical America.

The difficulty of identifying specimens
of Polybotyra has been partially alleviated
by local or regional floras, but these cover
only a limited portion of the geographic range
of the genus and are often incomplete. In
many of these floras, types were rarely
examined and names, therefore, were often
applied incorrectly. Only Sodiro’s (1897)
treatment of the ferns of Ecuador is available
for the Andean region; yet he was aware of
only 6 of the 23 species that occur in that
region. Vareschi’s (1969) treatment for Ven-
ezuela and Brade’s (1971) for Brazil have
helpful illustrations, but the names are often

MONOGRAPH of POLYBOTRA 5

misapplied. The best treatment of Polybotrya
for Mesoamerica is Stolze’s (1981) excellent
work for the Flora of Guatemala, but
Guatemala lacks several species found in
Mesoamerica. In brief, adequate keys and
descriptions are lacking for identifying
Polybotrya throughout most of its range and,
as a result, many specimens have been mis-
identified.

In this century, most of the research on
Polybotrya has focused on the specific level.
Brade (1935, 1948) described two new
species of Polybotrya in papers that included
other ferns as well. Later, in a series of three
papers published in the Brazilian journal
Bradea (1969a,b,c), he described ten addi-
tional new species from Brazil, Costa Rica,
and Venezuela. These papers were based on
field studies in the American tropics, and
most of the types were plants he had collected
by himself or with his brother Alfred. Brade
(1971) published a synopsis providing keys
and illustrations of the Brazilian species of
Polybotrya. Unfortunately, he died before
completing the descriptions, habitats, and
distributional information. Only five of
Brade’s names are accepted in this mono-
graph as representing valid species; the re-
mainder are relegated to synonymy.

More recently, Lellinger (1972, 1977)
published two important papers on Polybot-
rya. In the first, he described five new species
from South America and offered interesting
ecological notes on the genus. In the second,
which deals with other ferns as well, he de-
scribed two new species that are endemic to
Colombia. Only three of these seven species
are accepted here. About half of the species
described by Brade and by Lellinger have,
therefore, been relegated to synonymy in the
present work, largely because I found older,
obscure names during my research. In some
cases, these names had not been used since
their original publication. The proliferation
of names is one of the pitfalls faced by
taxonomists when synthetic or revisionary
works are unavailable for a group and in this
case indicates how much Polybotrya has
needed a monograph.

6 ILLINOIS NATURAL HISTORY SURVEY

Ecology

Polybotrya inhabits wet, shaded, tropical
forests. These may be hot, humid, lowland
rain forests or cool, cloud forests at high ele-
vations. The genus rarely occurs in disturbed
forests, and when it does, it is represented
by only a few isolated plants. However, small
scale disturbances within mature forest, such
as light gaps created when a huge tree dies,
appear to be important in establishing young
plants (Lellinger 1972). During fieldwork in
tropical America, I never observed young
sporophytes or their associated gametophytes
establishing themselves on roadbanks or on
shaded, disturbed slopes adjacent to roads.

Vol. 34, Art. 1

Furthermore, all herbarium specimens that
contained habitat information listed only pri-
mary forest as the habitat. The restriction of
Polybotrya to wet, shaded, primary, tropical
forest is reflected by its geographic distribu-
tion (Fig. 2a). The genus is absent from such
arid regions as central Mexico and the west-
ern coast of Peru and from such nonforested
regions as the Llanos of Venezuela and Co-
lombia (Figs. 2a & 3). The altitudinal range
of Polybotrya is from sea level to 2500 m,
with most plants collected from 500—2000 m
(Fig. 4). This range certainly corresponds to
my field observations that Polybotrya is most
diverse, frequent, and abundant at middle
elevations.

Figure 2. Distribution of Polybotrya. a. composite of Maps 1-21; b. number of species (left bar),
number of endemics (right bar). Countries or areas are outlined; those with only one bar have no
endemics. See Table 3 for numbers of taxa in each country.

November 1987

Geography

In the following discussion, the distribution
of Polybotrya is given according to the re-
gional centers defined by Tryon (1972).
These regional centers, all mountainous,
were determined by their high endemism and
species richness.

The most important regional center in the
diversification of Polybotrya is the Andean,
from Venezuela and Colombia south to
Bolivia (Fig. 2b, Table 2). The Andean Re-
gion contains 23 species, 12 of which are
endemic, and I suspect that new species of
Polybotrya will be found there. Within this
region, Colombia contains the richest

MONOGRAPH Of POLYBOTRA 7

Polybotrya flora: 17 species, 6 of which are
endemic (Table 3). The coastal mountains of
northern Venezuela contain P. canaliculata
and P. serratifolia, both occurring in cloud
forests from 1200-2400 m. Polybotrya
canaliculata has a very limited distribution
and is a true endemic to the coastal mountains
(Map 19); it occurs outside of the Andean
Region as defined by Tryon (1972). Polybot-
rya_ serratifolia is considered a_near-
endemic, extending southwest into the Andes
around Mérida and eastward to Trinidad
(Map 1).

The Serra do Mar Mountains of the South-
eastern Brazilian Region contain five species

© 100 900 Be «

Prepared by Menaik B Rypkeme

FiGurE 3. Distribution of Polybotrya species within regional centers of diversity and endemism for
tropical American ferns defined by Tryon (1972). Primary centers are indicated with dark shading;
secondary centers, with light shading. The number on the left is the total number of species; the number
in parentheses indicates how many of those species are endemic. See Table 4 for comparisons.

8 ILLINOIS NATURAL HISTORY SURVEY

of Polybotrya, all of which are endemic (Fig.
3, Tables 2 & 3). I find it remarkable that
this region has no species of Polybotrya in
common with those of the other regional cen-
ters (Table 4) or with intervening areas. This
distinctiveness of the Polybotrya flora is ac-
centuated by its lack of the simply pinnate
species, that is, subgenera Soromanes and
Sorbifolia. The biogeography of Polybotrya
strongly supports data from many other
sources that suggest that the isolated Serra
do Mar Mountains are a distinctive biogeo-
graphic center (Tryon 1972).

Only one species, P. goyazensis, occurs
in the Matto Grosso and intervening area be-
tween the Andean and Brazilian regional cen-
ters (Map 6). This species is closely related
to the widespread P. caudata and is probably
an example of the peripheral “budding” of a
new species from the southern part of the
range of P. caudata.

The Amazon basin contains one endemic,
P. glandulosa, which has been collected only
three times (Map 7). Most species of Polybot-
rya in the Amazon basin also grow in the

2500

2000

=
a
[2)
{o)

METERS

1000

500

Vol. 34, Art. 1

Andean Region. In contrast, the coastal
mountains of southeastern Brazil have
contributed no species to the Amazonian low-
lands. My field experience in the Amazon
basin of Ecuador and Peru showed that in
forests on wet, sandy soils P. caudata was
common; forests on lateritic soils, however,
had P. crassirhizoma, P. caudata, P. osmun-
dacea, and P. pubens. All five species were
abundant in the western Amazon basin where
I collected, and I suspect that they extend
farther eastward into the Amazon basin than
is shown on their range maps.

The Guiana Highlands Region does not
contain a distinctive Polybotrya flora. Only
six species occur there and most of these are
widespread elsewhere. The region, therefore,
has a high floristic affinity with the Andean
and Central American regional centers (Table
4). Polybotrya sessilisora, which occurs in
the Guiana Highlands and in the adjacent
Amazon basin (Map 20), is probably more
common in the Highlands and its range may
be centered there. Polybotrya has two notable
examples of Andean-Guianan range disjunc-

FiGure 4. Altitudinal ranges of Polybotrya species. The numbers above the bars correspond to the
species numbers assigned in the taxonomic treatment.

November 1987

tions. The first example, P. lechleriana, is
known in the Guianas from a single collection
at Mount Roraima in Guyana, 1900 km from
its nearest locality in Colombia (Map 7). This
disjunction is probably best explained by
long-distance spore dispersal from Andean
populations. The second example, P. frac-
tiserialis, occurs primarily in the foothills of
the Andes and is disjunct in the southern
Pakaraima Mountains of British Guiana,
2000 km from the nearest Andean population
in Ecuador. Unlike P. lechleriana, P. frac-
tiserialis is common and widespread in the
Guianas (Map 3).

The Central American Region contains
seven species, including one endemic (Fig.
3, Table 2). All seven species occur in Costa
Rica (Table 3). My fieldwork there showed
that P. polybotryoides, P. osmundacea, P.
alfredii and P. gomezii (endemic) were par-
ticularly common in the mountains and that
P. caudata was frequent in the costal low-
lands. Polybotrya alfredii and P. gomezii
were particularly abundant in cloud forests,
often growing together. In the mountains of

MONOGRAPH of POLYBOTRA

central Panama, just outside the Central
American Region as defined by Tryon
(1972), one endemic occurs, P. alata.

The Greater Antilles Region, otherwise
quite rich in fern species, has only one
Polybotrya species, the widespread P. os-
mundacea (Figs. 2 & 3, Table 2). Puerto
Rico lacks Polybotrya but certainly has suit-
able forest habitat. In the Lesser Antilles, P.
osmundacea is found on Grenada. Martinique
and Guadeloupe have the closely related en-
demic P. cyathifolia. The Caribbean islands
have played a minor role in the diversification
of Polybotrya.

Polybotrya caudata and P. osmundacea
clearly have the largest geographical and al-
titudinal ranges of any species in the genus
(Maps 5 & 18). With the exception of south-
eastern Brazil, these two species extend
nearly throughout the entire area covered by
all of the other species in the genus.

Polybotrya occurs on only one oceanic
island—Cocos Island, about 500 km (310
mi) southwest of Costa Rica in the Pacific
Ocean. Three species occur there: P.

TABLE 2. The geographic regions shown in Figure 3 and the species of Polybotrya that occur in them.
Species (numbers corre- No. in Region
Geographic Region spond to those in the (% of totalspecies No. Endemic
(after Tryon 1972) taxonomic treatment) in genus) (% for region)
Greater Antilles 28 1 (3) 0
Lesser Antilles 28; 29 2 (6) 1 (50)
Mexican 0 0
Mexican—Central American 2,9 2 (6) 0
Central American 2535599, 16; 275,28 7 (20) 1 (14)
Central American— Andean 253, 9;16, 23, 28 6 (17) 1 (17)
Andean D223, 45, One oe Mill la, 23 (66) 12 (52)
ISA ISS 167175 18519; 21,
22, 24, 25, 26, 28, 30
Andean—Guianan Lee Oyia le, 20, 9 (26) 1 (11)
BB S2
Guianan 5,6, 95 13,285.31 6 (17) 0
Guianan—Southeast Brazilian 5,7,9, 10,11, 12,31 7 (20) 0
Southeast Brazilian 10, 20, 33, 34, 35 5 (14) 5 (100)
Southeast Brazilian— Andean 10 1 (3) 0

10 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

TABLE 3. Distribution of Polybotrya by country. Endemics are in boldface. Numbers in parentheses
refer to the species numbers assigned in the taxonomic treatment.

Mexico: 2 species, no endemics. polybotryoides (2), caudata (9).

BELIZE: 2 species, no endemics. polybotryoides (2), caudata (9).

GUATEMALA: 3 species, no endemics. polybotryoides (2), caudata (9), osmundacea (28).
HONDURAS: 3 species, no endemics. polybotryoides (2), caudata (9), osmundacea (28).
NICARAGUA: 3 species, no endemics. caudata (9), alfredii (16), osmundacea (28).

Costa RICA: 6 species, 1 endemic. polybotryoides (2), sorbifolia (5), caudata (9), alfredii (16),
gomezii (27), osmundacea (28).

Cocos ISLAND: 3 species, no endemics. polybotryoides (2), caudata (9), osmundacea (28).

PANAMA: 5 species, 1 endemic. polybotryoides (2), caudata (9), alfredii (16), alata (23),
osmundacea (28).

CuBA: | species, not endemic. osmundacea (28).

JAMAICA: | species, not endemic. osmundacea (28).

Hartt: | species, not endemic. osmundacea (28).

GUADELOUPE & MARTINIQUE: | endemic. cyathifolia (29).

GRENADA: | species, not endemic. osmundacea (28).

TRINIDAD: 3 species, no endemics. serratifolia (1), caudata (9), osmundacea (28).

FRENCH GUIANA: 3 species, no endemics. fractiserialis (6), caudata (9), osmundacea (28).
SURINAM: 2 species, no endemics. fractiserialis (6), caudata (9).

GUYANA: 4 species, no endemics. fractiserialis (6), caudata (9), lechleriana (13), osmundacea (28).

VENEZUELA: 6 species, 2 endemics. serratifolia (1), sorbifolia (5), caudata (9), glandulosa (12),
osmundacea (28), canaliculata (32).

CoLomBIA: 17 species, 6 endemics. polybotryoides (2), suberecta (3), sorbifolia (5), crassirhizoma (7),
caudata (9), pubens (11), lechleriana (13), attenuata (14), stolzei (15), botryoides (17),
lourteigiana (18), pittieri (19), hickeyi (21), altescandens (26), osmundacea (28),

latisquamosa (30), sessilisora (31).

Ecuapor: 14 species, 3 endemics. polybotryoides (2), suberecta (3), andina (4), fractiserialis (6),
crassirhizoma (7), caudata (9), pubens (11), lechleriana (13), alfredii (16), puberulenta (22),
aequatoriana (24), appressa (25), altescandens (26), osmundacea (28).

PeERu: 11 species, no endemics. polybotryoides (2), suberecta (3), fractiserialis (6),
crassirhizoma (7), caudata (9), pubens (11), glandulosa (12), lechleriana (13), alfredii (16),
altescandens (26), osmundacea (28).

BoLiviA: 10 species, no endemics. fractiserialis (6), crassirhizoma (7), caudata (9),
pubens (11), lechleriana (13), alfredii (16), hickeyi (21), puberulenta (22), aequatoriana (24),
osmundacea (28).

PARAGUAY: | species, not endemic. goyazensis (10).

BRAZIL: 13 species, 5 endemics. sorbifolia (5), crassirhizoma (7), espiritosantensis (8), caudata (9),
goyazensis (10), pubens (11), glandulosa (12), cylindrica (20), osmundacea (28), sessilisora (31),
semipinnata (33), speciosa (34), pilosa (35).

November 1987

caudata, P. osmundacea, and P. polybot-
ryoides. As noted in the previous paragraph,
P. caudata and P.. osmundacea have the most
extensive range of any species in the genus.
Although not as wide ranging, P. polybot-
ryoides has a long north-south distribution
from Chiapas, Mexico, to Pasco, Peru, and
is the most far-ranging species of the sub-
genus Soromanes. This distribution supports
Tryon’s (1970) findings that the ferns of
oceanic islands tend to be those that are wide-
spread beyond the source area for the island
(the source area for Cocos Island is defined
by Tryon as Central America and Colombia).
According to Tryon (1970), widely distrib-
uted species have a broader ecological
amplitude and therefore establish themselves
more often on oceanic islands because of their
ability to grow under a wide range of environ-
ments.

Morphology and Anatomy

Information on the morphology and anatomy
of Polybotrya has been extremely useful for
assessing affinities of related genera and for
understanding functional aspects of the biol-
ogy of the genus. Every phylogenetic argu-
ment made here relies heavily on evidence
from stem anatomy (along with other fea-
tures, of course) in determining how Polybot-
rya relates to other fern genera and families.
Anatomical studies have also answered other
biological questions about the genus. For

TABLE 4.

MONOGRAPH of POLYBOTRA 11

example, studies of the diplodesmic venation
system showed how the sori are efficiently
supplied with water and nutrients. Studies of
sterile-fertile transitional leaves established
that the diplodesmic veins are homologous
with the veinlets of the sterile leaves. In short,
anatomical studies of root, stem, and leaf
have been valuable in understanding diverse
questions about the biology of Polybotrya.

Roots

A dense tangle of dark, tough, fibrous,
adventitious roots arises from the ventral side
of climbing stems and from all sides of ter-
restrial stems. No positional relationship
exists between the leaf bases and the roots;
roots are apparently borne at random. Roots
arising directly from the stem measure about
1 mm in diameter, but their terminal branches
may be only 0.3 mm wide. Roots branch at
irregular intervals, increasing the number of
tiny supportive rootlets. On climbing plants,
these tenacious rootlets penetrate the outer
layers of tree bark and firmly anchor the stem.
This firm attachment hinders pulling the stem
away from the tree. When a stem is forcibly
pulled away from a trunk, the large roots of
Polybotrya remain attached to the stem,
clinging to shredded pieces of bark, lichens,
and mosses.

I studied the root anatomy of six spe-
cies—P. alfredii, P. caudata, P. gomezii,
P. osmundacea, P. polybotryoides, and P.
sorbifolia. All were similar. Figure Sa shows

Affinities of Polybotrya floras between regional centers (see Fig. 3). Floristic affinity is the

percent species in common (C) of the total species in the two regions (A+B), i.e., LOOC/A+ B.

Regional Center

Total Species

SpeciesinCommon_ Floristic Affinity

(Tryon 1972) (A +B) (C) (%)
Andean & Guianan 29 5 17.2
Central American & Andean 30 5 16.6
Andean & Southeast Brazilian 28 0 0
Central American & Guianan 13 3 23.0
Guianan & Southeast Brazilian 11 0 0
Central American & Southeast Brazilian 12 0 0

12 ILLINOIS NATURAL HistoRY SURVEY

a root in cross section taken about 0.5 cm
from the ventral surface of a scandent stem
of P. osmundacea. The stele is weakly ellip-
tic, protostelic, and comprises about one-fifth
of the root’s diameter. Two protoxylem poles
occur; each is exarch and consists of four to
seven protoxylem elements. The center of
the stele contains three to five large
metaxylem cells. Surrounding the xylem are

Vol. 34, Art. 1

phloem elements—minute, angular, thin-
walled cells about the same diameter as the
protoxylem elements. They are not shown in
Figure Sa because they were poorly pre-
served.

Surrounding the stele is the innermost
layer of the cortex—the endodermis. It con-
sists of a single, continuous, light-colored
layer composed of thin-walled, rectangular

FiGureE 5. Root anatomy and various epidermal features of Polybotrya. a,b, and c are the same size.
a. root cross section of P. osmundacea; b. close-up of a stem meristele in cross section, showing root
trace diverging about midway between the laterally diverging leaf traces; c. phloroglucinol-containing
gland from the stem epidermis of P. caudata; d. stomata on stem aerophore of P. caudata; e. fertile
segment of P. stolzei, adaxial surface. Note different shapes of epidermal cells and the two kinds of
hairs; the flaccid hair represents the type that intergrades with scales. EN endodermis; EP epidermis;
LT leaf trace; PH phloem; RT root trace; SC sclerenchyma; SH sclerenchyma sheath surrounding a

meristele; T tracheid; X xylem.

November 1987

cells. Although the endodermis is narrow, it
contrasts sharply with the dark, thicker-
walled cells on either side. The anticlinal
walls have casparian strips that are easily vis-
ible even in unstained material.

Outside the endodermis is a ring of dark,
thick-walled fibers (Fig. Sa). This ring is the
most conspicuous feature of Polybotrya root
anatomy. Unstained, the walls of the fibers
appear orangy or reddish and may become
so thick that they nearly occlude the entire
lumen of the cell. The walls thicken evenly
on all sides, unlike many angiosperms and
some ferns where the deposition of cell walls
occurs centripetally (Bierhorst 1971; Wagner
1952).

Outside the ring of dark fibers, the corti-
cal cells take on a different appearance. In
contrast to the cells of the inner ring of fibers,
the outer cortical cells are parenchymatous
with thin, irregular, dark brown cell walls.

Cells of the epidermis are compact and
quadrangular, in contrast to the shape of the
cortical cells immediately below. The most
conspicuously differentiated cells of the
epidermis are the root hairs, generally 0.2—
1.0 mm long and straight, narrow, and unicel-
lular. Their color normally varies from brown
to amber, but brilliant yellow hairs occur in
several species. Unfortunately, the root
anatomy of Polybotrya cannot be compared
with that of other genera of dryopteroid ferns
because of inadequate information for those
genera.

Stem

Habit. The stem is climbing in all species
of Polybotrya except for P. fractiserialis and
P. sorbifolia, where it is terrestrial (Fig. 1).
Climbing stems grow horizontally about 0.5—
2.0 cm beneath the forest floor until they
contact a tree. The stem then turns upward,
anchoring itself to the tree by adventitious
roots produced on the ventral surface. Stems
usually climb to a height of 1 to 5 m, but the
highest scandent stem I saw was just under
15 m. The longest underground stem that I
saw was traced back 8 m from where it en-
tered the soil at the base of its support tree.

MONOGRAPH of POLYBOTRA 13

As a stem begins to climb, it grows wider.
The diameter of a climbing stem may reach
3.5 cm; terrestrial stems seldom grow wider
than | cm. The internode length on climbing
stems is generally 10—15 cm.

Stems of the two terrestrial species of
Polybotrya differ in overall appearance from
stems of the climbing species. Stems of ter-
restrial species are about 15—20 cm long with
internodes 1-3 cm long. These short inter-
nodes impart a basketlike aspect to the ar-
rangement of leaves instead of the linearly
spaced aspect typical of the leaves of climb-
ing species (Fig. 1). The amount of scale
cover is another difference. Stems of the ter-
restrial species are less scaly than stems of
climbing species due to the sloughing off or
rotting away of scales, presumably because
the scales are buried in moist, tropical soils
where decay is rapid. Furthermore, the
petiole bases hide the few scales that remain
in terrestrial species; the best place to find
scales is the stem apex, where there is less
decay and fewer petiole bases. Despite these
differences, both types of stems show affinity
by sharing the distinctive anatomy described
below.

Branching is rare in stems of Polybotrya.
I never saw dichotomy of the stem apex, and
lateral branching was exceedingly rare in
scandent stems, although occasional in ter-
restrial portions. I did not investigate the re-
lationship, if any, of branching to leaf posi-
tion, but that examination might provide
further evidence for comparisons with other
genera of dryopteroid ferns.

Anatomy. The stems of all dryopteroid
fern genera have radially symmetric dictyo-
steles; however, the stems of Polybotrya are
distinctive (Fig. 6). The most noticeable fea-
ture in cross section is the vascular bundles —
terete, circularly arranged, each bundle en-
sheathed by a jet-black ring of sclerenchyma.
The 5 to 12 vascular bundles are positioned
around the central portion of the stem, their
dark outlines contrasting vividly with the
matrix of bright white ground parenchyma.
Numerous tiny leaf traces emanate from the

14 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

FiGurE 6. Stem anatomy of Polybotrya osmundacea, typical of the genus. Lower left is a cross section
of the stem with a meristele enlarged above. Note “xylary arm” in 2. AP aerophore; EN endodermis;
GP ground parenchyma; LT leaf trace; MS meristele; P parenchyma; PC pericycle; PH phloem;
RT root trace; T tracheid; SH sclerenchyma sheath consisting of fibers; X xylem; XP protoxylem.

November 1987

sides of the bundles, collectively forming a
C-shaped line that connects adjacent bundles.
This pattern is unmistakable (Fig. 6), and no
other fern genus yet studied has this kind of
stem anatomy. (For comparison of stem
anatomy with closely related groups, see Re-
lationships with Other Groups.)

The epidermis of the stem is darkly
sclerotized and one cell layer thick; it pro-
duces numerous scales (described separately
below). The cell walls of the epidermis are
relatively straight compared to the sinuous
ones of the leaf epidermis (cf. Figs. Sc & 9).

The stems of Polybotrya have minute
glandular hairs on the epidermis. These short-
stalked hairs have, in relation to their length,
exceptionally large, round heads, 54-56
microns in diameter (Fig. 5c). Because these
glands are tiny and few, they had been over-
looked until found by Widen et al. (1983),
who were also the first to report phloro-
glucinols in Polybotrya. The phloroglucinols
are apparently produced and stored in these
glands. In other dryopteroid ferns, such
glands are known to secrete an oleoresin,
along with various lipids, between the cuticle
and the outer layer of the cell wall. The
phloroglucinols are, presumably, located in
this secretion (Widen et al. 1983). Using ex-
tracts from the stem and petiole bases of P.
caudata, Widen et al. found that aspidin BB
was the principal compound, along with
smaller amounts of albaspidin BB, desaspidin
BB, and flavaspidic acid BB. Internal glandu-
lar hairs, which probably also contain
phloroglucinols, occur in the stems and
petioles of other dryopteroid ferns (Mehra &
Mittal 1961). I did not, however, find internal
glands in Polybotrya.

Aerophores occur on the stems of most
species of Polybotrya but are apparently ab-
sent in species having a mat of thick, woollike
scales, such as in P.»aequatoriana, P. alte-
scandens, and P. crassirhizoma. On living
stems, aerophores appear as yellowish white
lines 1-3 mm wide that connect with
aerophores on the petiole (for description of
petiolar aerophores, see section below on

MONOGRAPH of POLYBOTRA 1S

petioles). Aerophores usually extend 2—6 cm
down the stem from the base of the petiole.
In cross section, they project about 0.5—1
mm above the surrounding surface and con-
sist of thin-walled parenchyma cells that in-
terrupt the otherwise continuous, thickened
epidermis (Figs. 6 & 16g). Stomata abound
and can be seen with a binocular microscope
(Fig. 5d). Presumably, aerophores allow oxy-
gen into the stem that is otherwise surrounded
by compact, oxygen-impermeable, scleren-
chyma fibers. Because aerophores shrink and
darken upon drying, thereby matching the
contour and color of the stem’s scleren-
chyma, they cannot be seen on herbarium
material.

Below the epidermis is a well-developed
hypodermis, usually three to eight cells thick
and darker than the epidermis that forms a
conspicuous ring around the white ground
parenchyma. The cells of the hypodermis are
thick-walled sclerenchyma fibers that thwart
cutting the stem, sometimes even with a hefty
machete, and I had problems cutting thin,
even, cross sections for microscopic study.
A ring of hypodermal sclerenchyma also en-
circles the stem in closely related genera such
as Arachniodes, Cyclodium, Maxonia, Olfer-
sia, and Stigmatopteris (pers. obs.).

Below the hypodermis lies the cortex,
which consists of scattered clusters of
brachysclereids (stone cells) in a matrix of
whitish parenchyma. The parenchyma cells
are generally rounded and contain abundant
amyloplasts. In a freshly cut stem, the white
color of these cells contrasts vividly with the
darkly sclerotized hypodermal and meristelar
sheaths. Sclereids never abound, as in Olfer-
sia (Fig. 16i), but are scattered in clusters
throughout the ground parenchyma. The dep-
osition of secondary walls is extremely un-
even; most of the wall is laid down on the
side of the cells facing the inside of the
sclereid cluster. A small, cuplike lumen can
usually be seen on the side of the cell facing
away from the center of the sclereid cluster.
The thick, dark cell walls contain numerous
simple pits and a lesser number of ramiform

16 ILLINOIS NATURAL HisTORY SURVEY

pits. Many Dryopteridaceae and Lomariop-
sidaceae also have sclereid clusters in the
ground parenchyma of their stems (pers.
obs.).

Inside the sclerenchyma sheath that sur-
rounds each meristele lies a band of paren-
chyma two to four cells wide (Fig. 6). The
cells of this layer are more compact and
smaller than the cortical parenchyma, but like
the cortical parenchyma, they contain amylo-
plasts. This inner parenchymatous layer stops
abruptly at the endodermis, which stands out
as a single row of clear, narrow, rectangular
cells interrupting the darker parenchyma on
either side of it (Fig. 6). The next layer toward
the center is the pericycle, which consists of
compact, orangy cells that form a continuous
band around the xylem and phloem.

The phloem consists mostly of clear, an-
gular, thin-walled sieve cells (Fig. 6.2 &
6.4). Phloem is broadest on the distal side
of the stele but forms a thin band elsewhere.
This band is broken in two locations by arms
of the xylem that extend to the pericycle (Fig.
6.2). Scattered among the clear cells of the
phloem are parenchymatous cells.

Inside the ring of phloem is the xylem,
which has large, conspicuous metaxylary
tracheids (Fig. 6.3). These tracheids occur
singly and in clusters within a matrix of dark
brown (when unstained) parenchyma. The
tracheids are generally rounded or somewhat
angular; when adjacent to another tracheid,
the walls are straight. Numerous pits occur
between all the cells of the xylem. Pro-
toxylem elements occur in the xylary “arms”
(Fig. 6.2). Each meristele is, therefore,
diarch and exarch.

Root traces arise between the xylary arms
of the meristele (Fig. 5b). The roots diverge
through the cortex to the ventral surface of
the stem. In contrast to the root traces, the
leaf traces originate from the xylary arms.
Four to six leaf traces are usually produced
at each leaf gap from adjacent meristeles.
During their passage from the cortex to the
petiole, the leaf traces bifurcate and anas-
tomose, forming a reticulum like that in
Maxonia (Chandra 1975). Each leaf trace is

Vol. 34, Art. 1

surrounded by a single layer of sclerotized
cells (Fig. 5b). The walls of these cells are
brownish and slightly thickened on the inner
tangential surface. The distance from the in-
ception of a leaf gap to the point where the
leaf diverges from the stem is about 10—15
cm.

Stem scales. Many species of Polybotrya
have distinctive stem scales. Some species,
in fact, can be identified solely on the basis
of their stem scales (thus the importance of
always collecting part of the stem). Color is
the easiest feature of the scales to use in iden-
tifying the species of Polybotrya. Botanists
should have no difficulty identifying P. alte-
scandens, with its bright golden or yellow
scales; P. crassirhizoma, with its dull orange
or red scales; and P. latisquamosa, with its
shiny, castaneous scales. Depending on the
species, scales range from concolorous to
slightly darker in the center to bicolorous with
a dark central stripe.

Scale habit is another helpful identifica-
tion feature. Some species, such as P. alte-
scandens and P.. crassirhizoma, have narrow,
densely tangled scales that impart a woolly
appearance to the stem. Other species, such
as P. appressa, P. caudata, and P. cylin-
drica, have thick, dark, massive scales that
are conspicuously appressed to the stem. Be-
cause of their great width and spreading habit,
the scales of P. serratifolia are distinct from
those of other species of the subgenus
Soromanes. Most species of Polybotrya have
ascending and slightly spreading scales—a
characteristic that is not particularly distine-
tive because of its prevalence in the genus.

The shape of stem scales is usually not
helpful in identification because most species
have lanceolate to linear-lanceolate stem
scales. Two species, however, P. alata and
P. altescandens (Figs. 42 & 45), have ex-
tremely long, attenuate scales that differ strik-
ingly from those of the other species. Petiole
scales are generally similar to those of the
stem but tend to be shorter and wider. The
petiolar scales of P. latisquamosa have be-
come, as the specific epithet implies, greatly
widened so that this characteristic distin-

November 1987 MONOGRAPH of POLYBOTRA

guishes the species from all others in the — easily visible cell walls (Fig. 7). Most of the
genus (Fig. 50d). scale is one cell layer thick, although the

Despite variation in color, habit, and center may be thicker and darker. The mar-
shape, stem scales are of two general types, gins vary from denticulate to erose, with the
here termed simply Type | and Type 2. Scales _ teeth formed from the projecting ends of two
of Type | are thin and translucent and have adjacent cells (Fig. 7). Marginal teeth in

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18 ILLINOIS NATURAL HISTORY SURVEY

Polybotrya are never glandular, as in Bolbitis
(Hennipman 1977) and Stigmatopteris (pers.
obs.). Scales of Type | are attached basally
at a single point or, more rarely, across the
full length of the base. This point or line of
attachment is always darkly sclerotized (Fig.
7a). Representative species with scales of
Type | include P. crassirhizoma, P. goyazen-
sis, P. puberulenta, and P. serratifolia (Figs.
26, 29, 41, & 19).

Scales classified as Type 2 are thick,
opaque, and composed of many cell layers.
The margins are entire or subentire. The base
is greatly thickened, sometimes almost
stipelike, and is curved and ascending. When
the scale is removed, a circular or oval scar
remains on the stem. Scales of this type are
usually appressed, but they are squarrose in
P. osmundacea. Representative species with
scales of Type 2 include P. appressa, P.
caudata, and P. cylindrica (Figs. 44, 28, &
39). In contrast to scales of Type 1, which
show tremendous variety in habit, color, and
shape, those of Type 2 are relatively
homogeneous.

Evidence suggests that morphological
transition occurs easily between the two types
of scales. Species such as P. lechleriana, P.
lourteigiana, P. osmundacea, and P.. pubens
have both types of scales but on different
individuals. Even on a single plant, scales
belonging to Type | can occur on the climb-
ing portion, with scales of Type 2 and/or
intermediates on the terrestrial portion. If
morphological transition is easily ac-
complished, then evolutionary change in
scale type could be expected to occur in
closely related species pairs. This hypothesis
is supported by two closely related species
P. caudata and P. goyazensis (Figs. 28 &
29); the former has scales of Type 2 whereas
the latter has scales of Type 1. Clearly, the
stem scales of Polybotrya have been mor-
phologically and evolutionarily plastic.

The evolutionary advantage of scales in
Polybotrya is unknown. Their contribution
to protecting the stem from mechanical dam-
age is probably minimal since the stem is

Vol. 34, Ar. 1

surrounded by a tough, resistant, scleren-
chymatous sheath. Scales seem unnecessary
on the older, hardened parts of the stem. Dur-
ing fieldwork in Latin America, I found sev-
eral plants of Polybotrya that had lost all of
their scales on the lower, older part of the
climbing stem, yet the plants appeared vigor-
ous. Perhaps the scales afford protection to
the stem apex, where young tissues have not
yet become sclerified.

Another hypothesis is that the scales re-
tain water by capillary action for later use by
the roots. Retention would be advantageous
to a climbing plant that has the leaves on the
scandent portion of the stem separated by
several meters from the roots in the soil.
Stems collected in the field, however, never
seemed to hold water between the scales and
plants never appeared water stressed. The
bark of the support tree, which is usually
covered with water-retaining mosses,
lichens, and organic debris, probably could
provide most of the water needed by the plant.
Further, the retention of capillary water be-
tween stem scales might be harmful in a hot,
humid, tropical environment that promotes
decay. No satisfactory hypothesis, therefore,
has been offered to explain the adaptive sig-
nificance of scales in Polybotrya.

Even more difficult to explain is the
adaptive advantages of the numerous minor
modifications of scales. What could be the
function of a dark, central stripe, of reddish
color, of an appressed habit, or of a linear
shape? How these and other minor modifica-
tions of scales are significant in evolution, if
indeed they are, is baffling.

Leaves

Petiole. The petiole base in some ferns
is specialized to accumulate food and to per-
sist as a storage organ long after the rest of
the leaf to which it was attached has withered
and decayed. These specialized petiole bases,
called “trophopods,” are a new source of sys-
tematic data in ferns (Wagner & Johnson
1983). Polybotrya, however, lacks troph-
opods, and the food-storing function of the
trophopod is performed by the stem in the

November 1987

amyloplast-containing cells of its ground
parenchyma.

A cross section of the petiole of Polybot-
rya reveals leaf traces that are arranged in a
mushroomlike outline, with the base of the
mushroom oriented adaxially (Fig. 8a). This
pattern also occurs in Cyclodium trianae, a
species closely related to Polybotrya, and in
some species of the similar family Lomariop-
sidaceae (Hennipman 1977; Holttum 1978).
Other closely related dryopteroid genera,
however, such as Cyrtomium, Dryopteris,
and Polystichum, display a C-shaped arrange-
ment of leaf traces in the petiole (pers. obs.).
These genera also have fewer leaf traces than
Polybotrya, probably as a consequence of
their narrower petioles. Further study of
petiole anatomy in dryopteroid genera may
provide helpful information for assessing
their relationships.

In Polybotrya, the two vascular bundles
closest to the adaxial surface of the petiole
differ from the lower vascular bundles in sev-
eral respects. The most obvious difference is
their shape: elongate and slightly curved in-
stead of round (cf. Fig. 8b,d). Inside each
of the two adaxial bundles, the xylem is hook-
shaped in outline, in contrast to the lenticular
shape seen in the lower traces. Only these
uppermost vascular bundles, not the lower,
yield the lateral traces that supply the pinnae.
As they traverse the rhachis and petiole, all
traces remain separate; they do not anas-
tomose to form a reticulum within the petiole.

Aerophores run down the sides of the
petiole and join below with those on the stem.
On living petioles, they appear narrow,
linear, and light green. The surface of the
aerophores bears stomata, and beneath them
is parenchyma that contains intercellular
spaces. This parenchyma, when seen in cross
section (Fig. 8c), interrupts the otherwise
continuous band of collenchyma that encir-
cles the petiole. Upon drying, the aerophores
darken and collapse, leaving a sulcus on
either side of the central, adaxial sulcus of
the petiole. Drying, therefore, makes the
petiole trisulcate; in living petioles only a
central sulcus is present.

MONOGRAPH of POLYBOTRA 19

Petiole and stem mucilage. Mucilage
has apparently never been reported in
Polybotrya or any other genus of Dryop-
teridaceae. During fieldwork in Costa Rica
and Ecuador, however, | found that stems
and petiole bases of P. alfredii usually had
a thick coat of translucent mucilage similar
to that on the stems and/or petioles of certain
Blechnum and Thelypteris species. Upon dry-
ing, the mucilage disappeared completely,
although sometimes the scales exhibited a
matted or flattened appearance, as if stuck
together. Attempts to rehydrate the mucilage
failed.

Other species of Polybotrya probably
have mucilage. Several dried specimens of
P. lechleriana and P. lourteigiana exhibited
matted, flattened scales that suggested the
earlier presence of mucilage. These two
species are closely related to P. alfredii, an
observation that suggests that mucilage is
probably confined to the P. alfredii group
(species nos. 13—20). I did not see matted,
flattened scales in any other species of
Polybotrya. The mechanism of mucilage se-
cretion and its adaptive significance, if any,
is unknown.

Nectaries. Koptur et al. (1982) were the
first to report the existence of nectaries in
Polybotrya. They found them in P. osmun-
dacea on the rhachis of unfurling leaves, near
the yellowish, lateral, linear aerophores. Sev-
eral times during fieldwork in Costa Rica, I
noticed ants on young leaves, but I never saw
them feeding on nectar. The nectar itself
would certainly be nutritious because it con-
tains sucrose, glucose, and fructose in con-
centrations up to 35—75 percent by weight
(Koptur et al. 1982). Amino acids also occur
in the nectar, and Koptur et al. list those
present. Although ants may benefit from the
nectaries, additional fieldwork is needed to
establish whether or not the plants benefit.
Apparently, ants do not defend the plants —at
least I was never attacked by ants while col-
lecting Polybotrya.

Costa-costule architecture. Costa-cos-
tule architecture refers to the morphology of
the juncture of these axes. Several kinds of

20 ILLINOIS NATURAL HistoRY SURVEY Vol. 34, Art. 1

FiGuRE 8. Petiole anatomy of Polybotrya alfredii. a. cross section of petiole 2 cm above the stem
showing the arrangement of vascular bundles in the shape of a mushroom; b. adaxial bundle showing
hooked xylem; c. aerophore; d. abaxial bundle. COL collenchyma; EN endodermis; EPI epidermis;
T tracheid.

November 1987

costa-costule architecture occur; the most
thorough review of them is by Holttum
(1959); more than any other pteridologist, he
has shown the importance of this feature in
the taxonomy of ferns. According to Holttum
(1984), the dryopteroid genera have “midribs
of ultimate leaflets grooved, the groove of
the rachis bearing the leaflets being open to
admit the leaflet-groove, the margin of the
lamina of the leaflet being decurrent (but not
prominent) down the side of the rachis; hairs
of ctenitoid type lacking.” This type of ar-
chitecture is illustrated in Figure 47h.
Holttum (1984) describes the tectarioid gen-
era as having “midribs of ultimate leaflets
more or less prominent (in Tectaria some-
times slightly grooved) and bearing ctenitoid
hairs, usually many.” In Polybotrya, the
basic type of architecture is dryopteroid, a
finding that supports its classification among
the dryopteroid ferns.

Within Polybotrya, however, modifica-
tions are found in the dryopteroid pattern of
costa-costule architecture. Typical dryop-
teroid costa-costule architecture is shown in
the most primitive group in the genus—the
group of species related to P. osmundacea.
In contrast, the most advanced species in the
genus—P. attenuata, P. lechleriana, and P.
stolzei—always have the costular groove
truncated by the ridges of the costa; therefore,
the groove is not decurrent (Figs. 33b, 32b,
& 34f). This truncated costa-costule architec-
ture is considered to have been derived from
the decurrent-grooved condition, as evi-
denced from outgroup comparison with other
dryopteroid ferns. Intermediates, such as P.
aequatoriana and P. alfredii, have costular
grooves that gradually fill up near the costa,
so that the ridges of the costa are only slightly
interrupted by the shallow, weakly decurrent
groove of the costule (Fig. 43d). In P. alfredii
and P. lourteigiana, these three types of
costa-costule architecture can be found on
different parts of the same leaf, although the
intermediate type predominates. In summary,
most species of Polybotrya have dryopteroid
costa-costule architecture, but distinctive
modifications of this basic type occur.

MONOGRAPH of POLYBOTRA 21

Anatomy of the lamina. | studied leaf
clearings and cross sections for 17 species of
Polybotrya to determine differences between
species or subgenera. Leaf clearings revealed
that stomata were absent from the adax-
ial epidermis but abundant on the abaxial sur-
face between the veins. The long axes of the
stomata ran almost parallel to the veins. The
mean length of the stomata (for all species)
was 56 microns, but means for the species
ranged from 48 to 74 microns. Subsidiary
cells surrounded the distal one-half to two-
thirds of the guard cells. The shape of the
subsidiary cells can be like the sinuous epi-
dermal cells or slightly more rounded and
circular (Fig. 9d,f).

In face view, the epidermal cell walls fit
together like the pieces of a jigsaw puzzle
(Fig. 9). This interlocking is best exemplified
by the abaxial epidermal cells, which have a
more highly irregular outline than the adax-
ial ones. The cells above and below the veins,
however, have straight walls. The slightly
elongate axes of the cells are oriented parallel
to the veins and point in the direction of the
leaf margin.

In cross section, the abaxial epidermis
and adaxial epidermis form a single, compact
layer of relatively thick-walled cells (Fig.
9g,h). Palisade parenchyma is absent from
the mesophyll, which is composed mostly of
large intercellular air spaces surrounded by
long, cylindrical, parenchymatous cells. An
arm of collenchyma extends from the abaxial
epidermis to the vein, where it forms a sheath
around the vein but does not continue to the
adaxial side of the leaf. Leaf anatomy was
similar in all species of Polybotrya, probably
because the entire genus occupies a similar
habitat.

Hair types. Two fundamentally different
types of hairs occur in Polybotrya. One type
intergrades with scales, the other does not.
The nonintergrading type is erect or spread-
ing, usually cylindrical, and whitish or
tawny. The size of hairs of this type varies
tremendously, from unicellular hairs that are
less than 0.1 mm long and not visible to the
naked eye to readily visible acicular hairs

ILLINOIS NATURAL HISTORY SURVEY

that have up to 15 cells and are up to 2 mm
long (Fig. 10 a—i,k). The apical cell is point-
ed, rarely rounded. All but five species of
Polybotrya have this kind of hair.

Hair of the second type is fundamentally
different from hair of the first type because

Vol. 34, Art. 1

each hair is a tiny, uniseriate scale and, there-
fore, by definition, a hair. Every intermediate
stage exists between these hairs and the
scales, as evidenced by studying their change
in form from the apex to the base of the costa
(Figs. 10j & 36a,b). At the apex of the costa,

Recall
eet
oe

!

FiGureE 9. Leaf anatomy of Polybotrya. a—f. face views of the abaxial leaf epidermis. The elongated
cells with straight walls lie above the veins. g, h. cross sections of the leaf. a. Polybotrya serratifolia;
b. P. pubens; c. P. fractiserialis, d. P. canaliculata; e, g. P. crassirhizoma; f, h. P. speciosa.

November 1987

only the tiny uniseriate scales, the “hairs,”
occur. These are readily distinguished from
other hairs by their flattened cells, appressed
habit, and darkened transverse walls. To-
wards the base of the costa, the hairs become
longer and are composed of more cells; soon

== SS

a

a
SSS

MONOGRAPH of POLYBOTRA 23

are found “hairs” that are two cells wide near
their base, that is, scales. These small scales
grade imperceptibly into the larger scales on
the rhachis. The scales on the rhachis and
costae are, therefore, developmentally
homologous with this second kind of hair.

0.1 mm

FiGure 10. Hair types in Polybotrya taken from the abaxial surface of the costa. a. P. pubens;,
b. P. semipinnata; c. P. altescandens; d. P. pittieri; e. P. alfredii; f. P. glandulosa, g. P. speciosa;
h. P. osmundacea; i. P. caudata; j. P. lechleriana (reduced scale type); k. P. cylindrica.

24 ILLINOIS NATURAL HISTORY SURVEY

These hairs are called proscales since they
are developmentally, and probably evolu-
tionarily, precursors to scales (Moran 1986).

Proscales occur on the lamina and axes
of all species of Polybotrya and all genera
of dryopteroid ferns. Proscales are, however,
so minute that pteridologists often overlook
them or do not bother to mention them. Pro-
scales are smallest on the distal parts of the
lamina (Fig. 49h) and at least 30 is needed
to see them. Their flattened cells, often hav-
ing reddish cross walls, occasionally twist
upon drying, imparting the appearance of a
Ctenitis-hair. These are not true Ctenitis-
hairs, however, because such hairs never in-
tergrade with scales (Christensen 1913). This
nonintergrading feature is rarely mentioned
in the definition of Ctenitis-hairs.

Fertile Leaves

Moran (1987) has presented a detailed study
of the sterile and fertile leaves (trophophylls
and sporophylls) of Polybotrya. Below is a
synopsis of the subject.

The fertile leaves of Polybotrya resemble
a skeleton of the sterile ones because their
green lamina has been reduced to a narrow
wing above the veins. This extreme dimor-
phy, termed “holodimorphy,” is distin-
guished from other types of dimorphy by the
nearly total loss of photosynthesis in the fer-
tile leaf (Wagner & Wagner 1977). The re-
duction of green tissue is the most conspicu-
ous aspect of dimorphy, but differences in
seasonality, duration, orientation, insertion
on the stem, size, color, texture, and venation
are also involved. Thus, sterile-fertile leaf
dimorphy is an ensemble of characteristics
and not merely a single feature of the leaf.
Pteridologists previously described the sori
of Polybotrya as acrostichoid, but three types
of sori occur in the genus: botryoid,
coenosoric, and a new type here termed “am-
phiacrostichoid.” Botryoid sori, which are
round and discrete, are considered primitive
on the basis of outgroup comparison with
other dryopteroid ferns (Fig. 38b). The
coenosoric type, which are oblong to linear,
probably evolved from the botryoid type by

Vol. 34, Art. 1

basipetal fusion of the sori (Fig. 35h). The
amphiacrostichoid type developed from a
coenosoric ancestor by the expansion of the
margins of the receptacle beyond the original
adaxial surface; as a result, the receptacle
assumed its own adaxial surface and sporan-
gia appeared on both surfaces of the leaf (Fig.
30g). Coenosoric and amphiacrostichoid sori
contain diplodesmic veins that are develop-
mentally homologous with veins of the sterile
leaf.

Sporangia. The sporangial stalk of
Polybotrya is three-celled at its apex, im-
mediately below the capsule. Below the apex,
however, only two rows of cells occur, each
three to four (rarely 5) cells long (Fig. 11).
Where the two rows of the base join the three
cells at the apex, a paraphysis occurs; it is
always uniseriate, unbranched, and multicel-
lular. About three-fourths of the species of
Polybotrya have paraphysate sporangial
stalks, but paraphyses occur on only 40—80
percent of the stalks from a single sample.
In dried or rehydrated material, the apical
cell of the paraphysis is smaller and shriveled
compared to the cells below. In fresh mater-
ial, the apical cell appears swollen and glan-
dular. Sporangial stalks of P. fractiserialis
are unique because they have lightbulb-
shaped, glandular cells at the base of the
paraphyses (Fig. 11g). The functions of these
glandular cells and paraphyses are unknown.

Sporangial capsules of Polybotrya are
typical of those found in related dryopteroid
ferns. The mean number of annular cells for
species of Polybotrya ranges from 12 to 24,
with most species having between 14 and I8.
Both the epistomium and hypostomium con-
sist of three or four thin-walled, transversely
elongated cells. All species of Polybotrya
have glabrous sporangial capsules, except P.
pubens, which has short, subulate hairs at
the top of the capsule near the annulus (Fig.
30e,f). Usually two of these hairs occur on
either side of the annulus, creating the
impression that the capsule has “horns.”

Spores. Spore sizes, measured by the
longest axis, are given (when available) at
the end of each species description. Numbers

November 1987

are for spores measured immediately after
being placed in distilled water because this
medium is readily available to taxonomists.
Fifteen spores were measured per specimen.

Spores of most Polybotrya species range
from 45—65 microns in length, with extremes
of 32 and 80 microns. According to Tryon
and Tryon (1982), Polybotrya has relatively
large spores compared to those of other gen-
era in their tribe Dryopteroideae. The spores
appear dark brown when viewed with trans-
mitted light under a compound microscope
but deep orange when viewed with reflected
light under a dissecting microscope.

Spores of Polybotrya are fairly uniform
within the genus compared to the variation
encountered in other genera of dryopteroid

MONOGRAPH of POLYBOTRA 25

ferns (e.g., Dryopteris and Polystichum).
Polybotrya spores are monolete, with the
aperture linear and one-third to three-fourths
the length of the long axis. The aperture is
often obscured by the broad perispore folds
and smaller spines. The exospore appears
smooth (Fig. 12), as it does in spores of most
genera of dryopteroid ferns. The perispore
consists of two layers: the lower layer is thin
and appressed to the exospore; the upper layer
is thicker, with inflated folds and echinate to
various degrees (Fig. 12). No constant differ-
ences were found between the three subgen-
era or smaller species groups.

The principal differences between species
appear in the prominence and density of peri-
sporal folds and spines. Typically, the in-

e

FiGureE 11. Sporangia of various Polybotrya species. a. P. attenuata; b. P. sorbifolia, showing both
sides of the same sporangium; c. P. espiritosantensis; d. P. speciosa; e. P. osmundacea; f,g. P. fracti-
serialis (note in g the globose, glandular cell at the base of the paraphysis).

26 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

ws :

Se
FicureE 12. Spores of Polybotrya. a. P. semipinnata;b. P. osmundacea; ¢c. P. serratifolia; d. P. pittiert,
e. P. gomezii; f. P. alfredii; g. P. crassirhizoma, h. P. cylindrica; i. P. speciosa. a: Duarte et al

65322 (F). b: C. Schunke 380 (GH). c: Fendler 261 (GH). d: Lellinger & de la Sota 251 (US). e:
Moran 3241 (CR). f: Moran 3168 (CR). g: Plowman et al. 4025 (GH). h: Handro 2228 (GH). i: Webb
26 (GH). c and h are X 5000, all others are X 1000

November 1987

flated folds are well developed and the spines
are so numerous that they impart a “fuzzy”
appearance. However, the folds may be rela-
tively low, as in P. goyazensis and P. sor-
bifolia, or the spines may be reduced in height
and/or density, as in P. speciosa (Fig. 12i).
Species may also differ in spore size. This
range is clearly shown by the two terrestrial
species in the subgenus Sorbifolia: P. sor-
bifolia and P. fractiserialis differ greatly in
spore size (40—47 and 52-56 microns long,
respectively), a difference that probably re-
flects different ploidy levels.

Comparison of the spores shown in Figure
12 with the spores of other dryopteroid ferns
illustrated by Tryon and Tryon (1982) shows
that the spores of Polybotrya are most like
those of Maxonia and Stigmatopteris (Tryon
and Tryon include Cyclodium in Stigmatop-
teris). Cyclodium, which is most closely re-
lated to Polybotrya, has spores similar to
those of Polybotrya but less spiny (A.R.
Smith 1986). This observation supports evi-
dence from external morphology that these
genera form a closely related group.

Chromosome Numbers

The only previous reports of chromosome
numbers in Polybotrya came from two sepa-
rate counts of P. osmundacea, one from
Jamaica and the other from Trinidad; both
gave a chromosome number of n=41
(Walker 1966; Smith & Mickel 1977). To
add to the information about chromosome
numbers in the genus, I collected meiotic
material during fieldwork in Latin America.
Young fertile segments were placed for 3—5
hours in distilled water saturated with
paradichlorobenzene. The segments were
then removed, blotted gently, and placed in
a fixative of 3:1 ethyl alcohol to glacial acetic
acid. The material was stored in a freezer
until it could be examined. Sporangia were
squashed in a drop of aceto-carmine and then
photographed.

This work recorded new chromosome
number counts for four species —P. alfredii,

MONOGRAPH of POLYBOTRA 27

P. altescandens, P. polybotryoides, and P.
serratifolia (Fig. 13). Each of the four counts
had a chromosome number of n= 41, a find-
ing that argues, along with morphological
features, for classifying Polybotrya among
the dryopteroid ferns, all of which have
n=41. Chromosome counts are still needed
from the other species of Polybotrya, espe-
cially where polyploidy may be involved in
the evolution of one species from another,
as in P. fractiserialis and P. sorbifolia.

Cladistic Analysis of the Species

A cladogram of species relationships was
constructed using the PAUP program, ver-
sion 2.3 (Swofford 1985; PAUP is an ac-
ronym for Phylogenetic Analysis Using Par-
simony). Table 5 shows the input data used
in the analysis, and Table 6 gives the charac-
ters and character states used in the input data
matrix. The character state trees (hy-
pothesized evolutionary pathways) for the
characters (Table 6) are shown in Figure 14.
Further information on the characters and jus-
tification of their postulated evolutionary
pathways is given in the Morphology and
Anatomy section.

Because of homoplasy in the data set,
PAUP found a large number of equally par-
simonious cladograms. Therefore, a consen-
sus cladogram (Fig. 15) was printed for the
first 50 trees to determine the branching pat-
terns they all had in common. The 50 trees
agreed on the branching patterns for 12
species— about one-third of the genus. The
groups that had congruent branching patterns
were subgenera Soromanes and Sorbifolia
(Fig. 15, serratifolia—espiritosantensis) and
the group of species related to P. caudata
(Fig. 15, caudata—pubens). Homoplasy in
the remaining species accounted for the
numerous, equally parsimonious clado-
grams. Nevertheless, distinct groups of
species are shown on the consensus clado-
gram, a result that supports the following
subdivision of the genus.

28 ILLINOIS NATURAL HisTORY SURVEY Vol. 34, Art. 1

FiGuRE 13. Chromosome squashes of four species of Polybotrya. All squashes have n=41. My
interpretations are at the right. a. P. altescandens, Ecuador, Pichincha, Moran 3559 (GH); b. P.
polybotryoides, Costa Rica, Cartago, Moran 2178 (MO); c. P. serratifolia, Venezuela, Trujillo, Moran
3709 (MO); d. P. alfredii, Costa Rica, Cartago, Moran 2442 (CR).

November 1987 MONOGRAPH of POLYBOTRA 29

TABLE 5. Data matrix for cladistic analysis of 35 species of Polybotrya. See text for discussion of
character states and polarity. Ancestor = hypothetical ancestor possessing all primitive character states.
?=unknown character state. NA = not applicable.

Character States
Species oA Se ORL Sie om LOM TI IA IS Aw Ise 7 18419520
serratifolia Sea OeINATO es Ola meleOe 28 lee YO) SO 1 30000" 0
polybotryoides Seer ese OMe N Ate OL 0? 2 be 1 iO? OF a 400: 0
suberecta See eo sOmINAG dee Oe 2502 SOU 10, To 10-0 1 i070 0.0
andina Ses OeNA he Or a 2 (O70) te 0 1 1 0 0: OO
sorbifolia Sane OR N AC OMLOs wie Nr Ouere tO. 7O. uO). st .0"0
fractiserialis Sas ORtNAG On Om Nie O ee ae Loe Oe Or e. Oo el .O. I
crassirhizoma POs Om NAG OO be tet One 2a ol 2a! Olt 0) “Oa Oa0
espiritosantensis Oe Ole ee OLOs we lO) 2a lL E210: 0) Sia), 0 sO 0
caudata Fee On sl Ope ee On clam On Oud OF 04,0 0,
goyazensis Sie 4 oe OO) Ol 2 ale OP eee? 0) ORO Ons00
pubens See eee le OOP OOM 10 Sees OP OM a0 O20 «0
glandulosa Slows eal eer ON Oro real” 407S2" TOMOPr 1° “0 s0'0'*.0
lechleriana ieee al OR Ole 2) 10 Oe eee ele OFNOs BON shim
attenuata Pelee: OF MOs i OS On ON is Ome 92" x0) 0) 25.10,
stolzei Pe ee ORO kOe Ot Open a Oa Oe Oh O ne Os 10h 0:
alfredii re OOo le Ol FO le OmeOMOe sO JOC OunO
botryoides OR ee eee O) Slee Or sO le le0) Oe0) 05 -0) 50.0
lourteigiana See See te Ou OM eyelets OnnO oe AO we. 0) 20070) (0's 0
pittieri Cee Oe Os ROR 1s Al O50: [O: Ota Op OnrOs a0
cylindrica es ele O Ol Fee eaO 4 OF Tiel 20 10M Tesh POF 0
hickeyi Del ee sTyLe a ee Opler TORO! Blt OSROMO)e 0,
puberulenta Doane lal ACPO lL 10 1 0) OPsOeO LOMA, YO-0e0r 0
alata Seale eee eee OR Ohl 2 Or Oe el OF ONO) 107.72 0 0
aequatoriana Pal Sele ORO Sha ol OO) “1 0) 20) 10) “1, 70 00
appressa See ales Lee Olea ial Onsale) Oa OieO,
altescandens Ta IS fe) pee Ce Oey Cat Cae | OP OR CORO MiP One OF (Oy °0
gomezii Semi etso ei 0) 0" th a 2 TO (0h) S40) *'0, SOMO:
osmundacea Poets OU nw Geely le Net eOr 0! 1) S080: 0.30
cyathifolia Dee SPO at One eee Olea eet SOMO) CORIO. sO.F,0
latisquamosa ie Uk Oe Olah: 2 SOLO. se 10; 10). 07 JO
sessilisora Deals lnkOe 9 ORS Te ORel OMA O Ol :2y 20'.:0:20%50
canaliculata Leslie 22. OOM Leelee oor ler @L Oe xO NOl HON 0S 0
semipinnata SP Leas) 0 1 0 rl Om Oewinnce OOmOr al! OOF 0
speciosa a Om eee ele Owe a2 a Ont 0 0 0. 0
pilosa Pega OL el OOO, Ol. Onl. anil 0; 105 (000
ancestor Saale agOe Tl Oe Ole a Le Ome Onn DeOLiOuO. 0

30 ILLINoIs NATURAL HISTORY SURVEY Vol. 34, Art. 1

TABLE 6. Characters and character states used in the cladistic analysis of 35 species of Polybotrya.
The numbers given to each character state correspond with those shown on one of the cladograms in
Figure 14.

1. Dissection of sterile leaves (Fig. 14a). 0 = 4-pinnate; 1 = 3-pinnate-pinnatifid; 2 = 3-pinnate; 3 = 2-
pinnate-pinnatifid; 4 = 2-pinnate; 5 = 1-pinnate.

2. Venation (Fig. 14b). 0=close and long-parallel; 1 = obliquely ascending; 2 = anastomosing.

3. Type of fertile leaf (Fig. 14c). 1= botryoid, but with lamina not completely reduced; 2 = fully
botryoid; 3 =coenosoric; 4 = caudate.

4. Pinnule arrangement (Fig. 14d). 0= anadromic; | = catadromic.

n

Symmetry of pinnule base (Fig. 14b). 0 = symmetrical and truncate; 1 = prolonged acroscopically
and truncate; 2 = symmetrical and cuneate.

Submarginal connecting strand (Fig. 14e). 0 = absent; 1 = several connections; 2 = present.
Stem scale base (Fig. 14d). 0= attached by a single point; 1 = attached across the width of the base.

Hair type (Fig. 14b). O=long, acicular; 1 = small, jointed; 2= uncinate.

<9 2) SI oN

Pubescence of laminar surface (Fig. 14b). 0 = both surfaces pubescent; 1 = glabrous; 2 = abaxial
surface pubescent.

10. Pubescence of laminar margin (Fig. 14d). 0= glabrous; | = ciliate.

11. Pubescence of costae (Fig. 14b). 0O= uniform and dense pubescence; 1 = moderately pubescent
or with scattered hairs; 2 = glabrous.

12. Lamina base (Fig. 14b). 0=deltate; 1 = reduced; 2 = cuneate.

13. Scale color (Fig. 14f). 0 = golden or yellow; 1 = brown; 2 = reddish or bright castaneous; 3 = cream
or whitish.

14. Receptacular hairs (Fig. 14d). 0= unbranched; | = branched.
15. Size of lamina (Fig. 14d). O= >1 meter; 1 = <1 meter.

16. Costal scale type (Fig. 14f). O= flaccid and ovate; 1 = linear to narrowly lanceolate; 2 = linear
and tortuous; 3 = caducous.

17. Apex of sterile leaf (Fig. 14d). 0= pinnatifid; 1 = subconform.

18. Stem habit (Fig. 14d). 0=hemiepiphytic; 1 = terrestrial.

19. Shape of the tertiary pinnules (Fig. 14b). 0=ligulate; 1 = oblong or ovate; 2 = obovate.
20. Spore size (Fig. 14d). 0=40—50 microns long; 1 = 52—56 microns long.

0 4
\ 5 { 2
u [ one f 1 : 3
\ vi / / Ale
3 1 1 0 0 1
A B c D E F

FiGure 14. Character state trees used in the cladistic analysis of 35 species of Polybotrya. Table 6
lists characters and character states. For example, tree B represents the hypothesized evolutionary
pathway for character states 2, 11, 12, and 19.

November 1987

attenuata

lechleriana

2
oa
x c
3 =
%
os %
o 20 8
Sj Ses £
a lee S wo
a
iS) 3 c
2 2) 3
a 2g N
= a= &
3 nd 3
a een > 3.2
= os 5
is} =
Log
5h = 2
5 ©
= S

serratifolia

appressa

MONOGRAPH of POLYBOTRA

31

stolzei
fourteigiana

alfredii
* cylindrica

puberulenta

3
2 2 8
o SFG:
a2) © o
Cc a o
% og 2
% Em
o SS 8.0
= as
hai o a
w &
BS5 9.0 £
= Zi w10 &
o “4 —~ ~)
os 1.05 3
35 322 3
BS £ p43 aS
5 S$ 71 +11.0 14:1 2 6
E 432}13.2
fe}

——

ancestor

FicurE 15. Consensus tree for 50 equally parsimonious cladograms of species relationships in Polybot-
rya. The number before the decimal refers to the character; the number after the decimal refers to the

character state (Table 6).

Subdivision of the Genus

I subdivide Polybotrya into three easily rec-
ognized subgenera: 1) Soromanes, sterile
leaves |-pinnate, veins anastomosing; 2) Sor-
bifolia, sterile leaves 1-2-pinnate, veins free,
close, and parallel; 3) Polybotrya, sterile
leaves decompound, veins free.

Subgenus Soromanes ranges throughout
Central America and the Andes, primarily in
mountainous areas, but it is conspicuously
absent from the Amazon basin and from
southeastern Brazil (Maps | & 2). This sub-
genus consists of four well-defined species
(species nos. 1—4) that occur in montane
forests, primarily from 500—2000 m (Fig. 4).

The only contemporary pteridologist who
has maintained Soromanes at the generic
level is Pichi-Sermolli (1977), who also
placed Soromanes and Polybotrya on sep-
arate branches of his phylogenetic diagram,
associated with different generic groups.
Other pteridologists, such as Christensen
(1905), Copeland (1947), and Tryon and
Tryon (1982) have subsumed Soromanes in
Polybotrya. My research on both genera has
shown that Soromanes is the closest genus
to Polybotrya. Both genera share a unique
stem anatomy: a circular grouping of meri-
steles with each meristele surrounded by a
dark, sclerenchymatous sheath (Figs. 6 &
16g). No other fern genus has this unmistak-

32 ILLINOIS NATURAL HisTORY SURVEY Vol. 34, Art. 1

Sasa

Po,
NOSES

Pre

LS III
wn,

Wooo:

\
in

\ =

)

L
é
y

FiGure 16. Polybotrya and closely related genera. a. venation of sterile pinna, Cyclodium meniscioides,
b. venation of sterile pinna, Polybotrya polybotryoides; c. pinna of Cyclodium trianae var. trianae
(compare to P. sessilisora and P. osmundacea); d. sterile pinna of Olfersia cervina, showing venation
entirely unlike any Polybotrya; e—i. stem cross sections (dorsal surface is up) of e. Maxonia apiifolia,
f. Lomariopsis fendleri, g. Polybotrya caudata, h. Bolbitis lindigii, i. Olfersia cervina. AP aerophore;
LT leaf trace; RT root trace; SH sclerenchyma sheath; SC sclereid or stone cell.

November 1987

able stem anatomy, one of the hallmarks of
Polybotrya. Other compelling similarities,
such as holodimorphic leaves, long-creeping
and densely scaly stems, petiole anatomy,
and spore morphology, further demonstrate
that Soromanes and Polybotrya are closer
genealogically to each other than to any other
genus; that is, they are sister groups. In my
opinion, these similarities justify including
Soromanes in Polybotrya.

Subgenus Sorbifolia is primarily South
American (Maps 3 & 4) and consists of four
species (species nos. 5—8) that occur in low-
elevation forests primarily from O-—1200
(1500) m (Fig. 4). Two species, P. sorbifolia
and P. fractiserialis, are very closely related
and contrast sharply with the rest of the genus
because of their terrestrial stems. These two
species also have distinctive venation: the
veins are long, parallel, and sharply ascend-
ing (Figs. 24 & 25). The third species, P.
crassirhizoma, is dissimilar from the others
and has dull orange stem scales and slightly
more spreading veins (Fig. 26). It is an abun-
dant fern in the western Amazon basin, from
Colombia south to Bolivia. Polybotrya es-
piritosantensis is included in this subgenus,
even though its leaves are 2-pinnate because
its venation (Fig. 27b) is exactly like that of
P. fractiserialis (Fig. 25e); this similarity
suggests a close relationship.

The species having decompound leaves,
all free veined (species nos. 9-35), make up
the subgenus Polybotrya, which is further
subdivided into four species groups. Since I
see no purpose in giving these groups formal
taxonomic names, I informally refer to them
with the name of a typical, widespread
species for each group, e.g., the “P. caudata
group.” The first three groups described
below have catadromic pinnules; the fourth
has anadromic pinnules. Pinnule arrange-
ment, however, cannot be used to classify
the species groups at a higher level because
no other characters correlate with it.

The P. caudata group (species nos. 9-12)
is the most distinctive because it has peculiar,
caudate fertile pinnules that are soriferous on
both surfaces, that is, amphiacrostichoid. Ac-
tually, what looks like the adaxial soriferous

MONOGRAPH of POLYBOTRA 33

surface is the expanded, thickened margin of
the receptacle; the true, phylogenetic, adaxial
surface is reduced to a thin green line (see
Morphology and Anatomy section). In addi-
tion to this diagnostic sorus, the group is
further united by the presence of whitish,
septate, acicular hairs (Fig. 10a,f,i). Poly-
botrya glandulosa is tentatively assigned to
this group because of similarities in leaf cut-
ting and pubescence; however, it has botryoid
fertile leaves. This character is constant in
P. pubens, but both P. caudata and P. goya-
zensis can be glabrous. The P. caudata group
is most frequent and abundant at low eleva-
tions from 0—1000 m, although P. caudata
itself occasionally occurs up to 1900 m (Fig.
4).

The second species assemblage, the P.
alfredii group, consists of eight primarily An-
dean species (species nos. 13-20). This
group has a characteristic appearance, al-
though it is difficult to describe because few
features are constant. In general, the pinnae
are short-stalked and crowded near the costa
and rhachis. The pinnae bases are more or
less equilateral and not acroscopically pro-
longed as in the rest of the genus. The tertiary
segments are often oval or rounded, never
narrow or strap-shaped. The grooves of the
costules are either truncated by the ridges of
the costa or very weakly admitted to the
groove of the costa (see Morphology and
Anatomy section). Most species in the P.
alfredii group have botryoid fertile leaves.

The third species group, the P. altescan-
dens group, consists of seven species (species
nos. 21-27). These species, with the excep-
tion of one endemic to the mountains of Costa
Rica, grow in the Andes at slightly higher
altitudes than the rest of the genus (Fig. 4).
This group has pinnule bases slightly to
strongly asymmetric, a characteristic that
helps to distinguish it from the P. alfredii
group.

The fourth species group centers on the
type of the genus, P. osmundacea, and differs
from the rest of the genus by having anadrom-
ically arranged pinnules. As shown in the
illustrations, the species (species nos. 28-35)
are very similar in dissection of the leaf. The

34 ILLINOIS NATURAL HISTORY SURVEY

pinnule base ascends obliquely on the basi-
scopic side and prolongates conspicuously on
the acroscopic side. This asymmetry imparts
a distinctive appearance to the pinnule base
(Fig. 48).

Relationships with Other Groups

Pteridologists have always classified Poly-
botrya with Dryopteris and such closely re-
lated genera as Arachniodes, Cyclodium,
Cyrtomium, Maxonia, Olfersia, Polystichop-
sis, Polystichum, and Stigmatopteris. These
genera share the following features with Poly-
botrya: base chromosome number x= 41,
dryopteroid costa-costule architecture, spores
monolete with prominently inflated perispo-
rial folds, more than three vascular bundles
in the petiole, and petioles and petiolules not
articulate at their bases.

Familial Relationships

The dryopteroid genera are most closely re-
lated to the tectarioid ferns, such as Ctenitis,
Lastreopsis, and Tectaria. These genera con-
stitute a natural group on the basis of their
costa-costule architecture and _— special
“Ctenitis-hairs.” As noted in the Morphology
and Anatomy section of this monograph,
Polybotrya lacks Ctenitis-hairs, and their ab-
sence argues strongly against a close relation-
ship with the tectarioid genera. Furthermore,
Polybotrya has costa-costule architecture of
the dryopterioid type rather than the tectarioid
type.

Polybotrya might possibly be construed
as a member of the Lomariopsidaceae. Three
genera of Lomariopsidaceae—Lomagram-
Lomariopsis, and Teratophyllum—
closely resemble Polybotrya in overall habit
because they have densely scaly, high-
climbing stems, strongly differentiated sterile
and fertile leaves, and nonindusiate, acros-
tichoid sori. Furthermore, at the anatomical
level, the three genera have diplodesmic ve-
nation and dark, sclerenchymatous sheaths
surrounding each meristele in the stem. Al-
though no pteridologist has ever classified
Polybotrya with the Lomariopsidaceae, these

ma,

Vol. 34, Art. 1

similarities prompt me to consider the possi-
bility.

Stem anatomy is tremendously important
in the taxonomy of Polybotrya and the
Lomariopsidaceae. The stem anatomy of
Polybotrya is unique and has been thoroughly
described in the Morphology and Anatomy
section. The stem anatomy of the Lomariop-
sidaceae is also unique among ferns because
it has a broad, strap-shaped, ventral meristele
that differs from the remaining circular or
oblong meristeles (Fig. 16f,h). Like Polybot-
rya, each meristele is surrounded by a dark,
sclerenchymatous sheath. This elongated
ventral meristele creates a dorsiventral dic-
tyostele. Presumably, this ventral meristele
was formed during phylogeny by the fusion
of two, once-distinct meristeles like the upper
ones (Holttum 1978). In fact, the two ventral
meristeles have not completely fused in
Lomariopsis, as evidenced by the shallow
indentation in the ventral band (Fig. 16f).
The ancestor of the Lomariopsidaceae, there-
fore, had a radially symmetrical dictyostele
of several to many, circularly arranged meri-
steles with each meristele surrounded by a
dark, sclerenchymatous sheath. In other
words, the ancestral stem anatomy was like
that of Polybotrya. If one looks at Figure
16g, a cross section of a Polybotrya stem,
and imagines what it would look like if the
two or three ventral meristeles were fused
into a single meristele, the result would be a
replica of the stem anatomy of the Lomariop-
sidaceae. This compelling similarity in stem
anatomy suggests that Polybotrya and the
Lomariopsidaceae arose from a similar an-
cestral stock among the dryopteroid ferns.
The possibility also exists, however, that
parallel evolution accounts for these
similarities in stems adapted for climbing.

The leaf architecture of the two differs,
however, and most species of Lomariop-
sidaceae have simply pinnate leaves. The
leaves of Polybotrya, like those of most
dryopteroid ferns, are primitively decom-
pound. All Lomariopsidaceae, except Bol-
bitis, have articulate leaves and/or pinnae. In

November 1987

contrast, Polybotrya and other dryopteroid
genera have nonarticulate leaves and/or pin-
nae. In the climbing genera Lomariopsis and
Teratophyllum, leaves on the terrestrial por-
tion of the stem (bathyphylls) differ greatly
from those on the scandent portion (ac-
rophylls). In Lomariopsis, the bathyphylls
are less divided (usually simple and entire)
than the acrophylls, and in Teratophyllum
they are more dissected than the acrophylls.
In Polybotrya, however, the bathyphylls and
acrophylls are about the same size and shape.

Venation is yet another difference. Sev-
eral kinds of venation occur in the three gen-
era of Lomariopsidaceae most similar to
Polybotrya. Teratophyllum has simple or
forked veins, as does Lomariopsis, except
that the veins of the former unite with the
cartilaginous margin (Holttum 1978).
Polybotrya also has free veins, but the pattern
is entirely different from that of these two
genera, as can be seen by comparing the il-
lustrations in this monograph with those
shown by Holttum (1978, Figs. 1-8). The
veins of Lomagramma differ completely from
those of these three genera; its veins form a
network of three or more rows of oblique
areoles without main veins. Venation, there-
fore, does not support a relationship between
Polybotrya and the Lomariopsidaceae.

In summary, Polybotrya and some genera
of Lomariopsidaceae share the following
characteristics: climbing stems, dimorphic
sterile and fertile leaves, diplodesmic veins,
and dark sclerenchymatous sheaths surround-
ing each meristele. The two groups, however,
contrast sharply in such features of leaf ar-
chitecture as the amount of dissection, pinnae
articulation, acrophylls versus bathyphyll dif-
ferences, venation patterns, and stem
anatomy. This conflicting evidence is dif-
ficult to assess. Certainly, the climbing habit,
sterile-fertile leaf dimorphy, and diplodesmic
veins have arisen many times in ferns, but
the similarity in stem anatomy is less easily
explained. I suspect, however, that the
Lomariopsidaceae may have had a separate
origin among the dryopteroid ferns, apart

MONOGRAPH Of POLYBOTRA 35

from Polybotrya. Strong evidence exists that
Polybotrya was derived from a Cyclodium-
like ancestor (see below).

Relation to Similar Dryopteroid Genera

Maxonia. This monotypic genus was first
described by Christensen (1916), who ob-
served that his new genus “must stand be-
tween Polybotrya and certain species of
Dryopteris grouped with D. amplissima
[Arachniodes].” Copeland (1947) also held
that Maxonia apiifolia (Swartz) C. Chr. rep-
resented a phylogenetic intermediate between
Arachniodes and Polybotrya (Fig. 17), and
most later pteridologists have agreed that
Maxonia is intimately related to Polybotrya.

Maxonia and Polybotrya appear very
similar because both have highly differen-
tiated sterile and fertile leaves and densely
scaly, climbing stems. Nevertheless, a com-
parison of stem anatomy shows some impor-
tant differences (Fig. 16e,g). The most evi-
dent dissimilarity is that each meristele of
Maxonia is not surrounded by a dark scleren-
chymatous sheath as in Polybotrya. Also, the
dictyostele of Maxonia is dorsiventral
(Chandra 1975; Walker 1972), unlike that of
Polybotrya, which is radially symmetrical.
Clearly, Maxonia lacks the stem anatomy that
characterizes Polybotrya, thus weakening the
hypothesis that these genera are closely
related.

Strong evidence exists that Maxonia is
most closely related to the American species
of Arachniodes, in particular A. macrostegia
(Hooker) Tryon and Conant. Indeed, if
Maxonia apiifolia had monomorphic leaves
it would without doubt be placed in
Arachniodes. The similarities between the
two genera are striking and are best seen in
the sterile leaves. Both genera have pinnules
arranged anadromically, basal pinnae elon-
gated basiscopically, and laminae broadened
notably at the base and of similar thickish
texture. In addition, the pinnules and smaller
segments have cuneate bases and acute
apices, thereby imparting a characteristic
“streamlined” appearance to the lamina that

36 ILLINOIS NATURAL HISTORY SURVEY

contrasts sharply with the truncate or acro-
scopically prolonged bases and rounded
apices in Polybotrya. Both Maxonia and
Arachniodes have costa-costule architecture
of the dryopteroid type (grooves decurrent
into each other; Holttum 1984). Finally, both
genera have remarkably similar brown, thick-
ish, round-reniform indusia.

The evolution of the climbing stem of
Maxonia poses no problem if Arachniodes is
accepted as the ancestor. Since the American
species of Arachniodes have creeping stems,
the potential to evolve a fully hemiepiphytic
stem, as in Maxonia, was initially present
and easily achieved.

In short, Maxonia is similar to Polybotrya
because it arrived at a comparable evolution-
ary grade or level, but it came from a different
source (Fig. 17). Polybotrya was probably
derived from Cyclodium (see below), and
Maxonia from Arachniodes, probably from
an ancestor close to A. macrostegia. Maxonia
should not, however, be subsumed with
Arachniodes because it is derived from it. It
is convenient, and certainly in harmony with
past taxonomic practice, to distinguish
Maxonia generically on the basis of its leaf
dimorphism and climbing stem.

Olfersia. This genus consists of a single
species, O. cervina (L.) Kunze, the place-
ment of which has been controversial. Most
recently, pteridologists have placed Olfersia
in Polybotrya because both have strongly di-
morphic sterile and fertile leaves, nonin-
dusiate sori, similar perispore morphology,
and densely scaly, creeping stems. Yet Olfer-
sia differs from Polybotrya by its venation,
conform apical pinna, and stem anatomy.

Polybotrya
jp
Maxonia

T

Arachniodes

Vol. 34, Art. 1

Moran (1986) studied Olfersia and concluded
that it and Polybotrya are sister taxa best
maintained in separate genera.

Cyclodium (sensu Smith 1986). Polybot-
rya was probably derived from an ancestor
which, if it were alive today, would be placed
in Cyclodium. This genus has two important
prerequisites for the evolution of Polybotrya.
First, the fertile leaves of Cyclodium are
slightly to strongly dimorphic. Second, the
stem is creeping in all species of Cyclodium,
and in several species it becomes fully
hemiepiphytic. Cyclodium, therefore, has the
genetic capacity to evolve two cardinal fea-
tures of Polybotrya: holodimorphic sterile
and fertile leaves and a climbing stem.

Moreover, Cyclodium contains species
that look remarkably like certain species of
Polybotrya. The 2-pinnate-pinnatifid leaf of
C. trianae (Mett.) A.R. Smith var. trianae
is strikingly like some species of Polybotrya
with respect to cutting and venation (cf. Fig.
l6c to P. caudata, P. osmundacea, and P.
sessilisora, Figs. 28,47, & 51, respectively).
In fact, several times during fieldwork in
Ecuador, I mistook sterile leaves of C.
trianae for terrestrial leaves of P. caudata.
Similarity is also seen in the simply pinnate
lamina of C. meniscioides (Willd.) Presl,
which has anastomosing venation notably
like that found in Polybotrya subgenus
Soromanes and is also simply pinnate (Fig.
16a,b). This similarity of venation was noted
long ago by Hooker and Baker (1874). Iam
not suggesting that these two species of Cy-
clodium gave rise to Polybotrya, but within
both genera, species have evolved with simi-
lar characteristics, and this homologous vari-

Maxonia Polybotrya
t
Arachniodes Cyclodium
T T
Ancestor (Dryopteris?)

FiGurE 17. Two phylogenetic hypotheses concerning the relationships of Maxonia and Polybotrya:
left, the hypothesis proposed by Christensen (1916); right, the hypothesis proposed here. See text for

explanation of characters involved.

November 1987

ation implies a close genetic relationship. In
conclusion, the close relationship between
Cyclodium and Polybotrya is evidenced by
their mutual possession of dimorphic leaves,
creeping stems, similar evolutionary tenden-
cies, and, at least in some species, remarka-
bly similar leaf dissection and venation.

Part Two:
Taxonomic Treatment

In evolutionary biology almost all phenom-
ena and processes are explained through in-
ferences based on comparative studies.
These, in turn, are made possible by very
careful and detailed descriptive studies. It is
sometimes overlooked how essential a com-
ponent in the methodology of evolutionary
biology the underlying descriptive work is.
Ernst Mayr (1982)

Notes on the Presentation of Data

In most cases, the key can be used with speci-
mens lacking fertile leaves. Fertile leaves,
however, are often helpful in identification
and are given as additional characters in many
couplets. The key will generally not work

basiscopic

MONOGRAPH of POLYBOTRA 37

when leaves are less than 45 cm long. To
facilitate the comparison of similar species,
I have arranged the species phylogenetically
rather than alphabetically. The synonymy
given for each species is complete. I have
kept the descriptions short by not repeating
characteristics that are constant for the genus.
For example, all species of Polybotrya have
a pinnatifid leaf apex; this characteristic is
mentioned only in the generic description and
is not repeated in each species description.
The distribution maps (see appendix) were
compiled from the specimens listed in the
Specimens Examined section in each species
treatment. In some cases, I could not find
localities despite searching various atlases,
indices, and gazetteers.

The terms basiscopic and acroscopic are
used frequently in the key and in descriptions
to refer to the sides of pinnules (Fig. 18).
The acroscopic side of a pinnule is the side
directed toward the apex of the pinna bearing
the pinnule. Conversely, the basiscopic side
of a pinnule is that side directed toward the
pinna base and rhachis. Two other frequently
used terms, catadromic and anadromic, refer
to pinnule arrangement. The pinnule arrange-
ment is catadromic when the pinnule closest
to the rhachis is basiscopic; when the pinnule
closest to the rhachis is acroscopic, the ar-
rangement is anadromic (Fig. 18). I have de-

acroscopic

side side
costule catadromic

tertiary

segment
rhachis —— costa

anadromic
PINNULE
PINNAE

FiGuRE 18. Terms frequently used to describe the leaves of Polybotrya. See text for further explantions.

38 ILLINOIS NATURAL HISTORY SURVEY

fined the specialized terms that refer to the
fertile leaves, such terms as botryoid,
coenosoric, and amphiacrostichoid, in the
Morphology and Anatomy section and in the
Description of the Genus section.

Description of the Genus

POLYBOTRYA WILLD.

Polybotrya Willd., Species Plantarum, ed. 4.
5:99. 1810. Type species: Polybotrya osmun-
dacea Willd.

Soromanes Fée, Mém. Fam. Foug. 2 (Hist.
Acrost.) 16. 1845.Type species: Soromanes
serratifolium Fée = Polybotrya serratifolia
(Fée) Klotzsch.

Botryothallus Klotzsch, Bot. Zeit. 4:104. 1846.
nom. nudum. Type species: Botryothallus
kunzei Klotzsch nom. nudum = Polybotrya
serratifolia (Fée) Klotzsch.

Stem hemiepiphytic, long-creeping, or (two
species) terrestrial and short-creeping, 1.0—3.5cm
wide (excluding scales), the scandent portions un-
branched, the terrestrial portions occasionally
branched; dictyostelic, in cross section with a
characteristic pattern of 4-10 circularly arranged
meristeles, each meristele surrounded by a dark
sclerenchyma sheath; aerophores present in most
(all?) species, continuous and decurrent from the
lateral aerophores of the petiole, darkening upon
drying and not visible in herbarium specimens, in
fresh material appearing yellowish, linear, 1-3
mm wide, slightly raised and bearing stomata,
rootlets produced only on the ventral surface;
scales numerous, densely covering the stem,
spreading or appressed, primarily of two general
types: 1) thick, dark brown, opaque, margins en-
tire, attached across the length of the curved, thick-
ened base, and 2) thin, variously colored (ranging
from yellow, orange, red to brown), translucent,
margins denticulate or highly erose, attached at a
central, basal point. Sterile leaves up to 2.0 m
long, reclining when mature, internodes generally
10-15 cm apart, bulbils lacking; petiole with 8-16
vascular bundles arranged in a mushroomlike out-
line, the base decurrent for a short distance on the
stem, scaly, with scales shorter and wider than
those on the stem, the aerophores present laterally
as thin yellowish green lines, the adaxial surface
slightly flattened, trisulcate in dried material; color
stramineous, light green or darkened abaxially;

Vol. 34, Art. 1

rhachis trisulcate below becoming unisulcate
above, the groove pubescent within; lamina
papyraceous to subcoriaceous, 1-pinnate (sub-
genera Soromanes and Sorbifolia) or 1-pinnate-
pinnatifid to 4-pinnate (subgenus Polybotrya).
usually lanceolate, the base rarely deltate or
cuneate, the apex pinnatifid; pinnae not articulate
to the rhachis, linear, lanceolate or deltate, sym-
metrical or slightly prolonged acroscopically, a
few species having the basal pair slightly pro-
longed basiscopically; pinnules arranged cata-
dromically or anadromically, symmetrical or
asymmetrical at the base, if asymmetrical then
with the basiscopic side reduced and oblique and
the acroscopic side prolonged; grooves of costules
decurrent into the the grooves of the costa or trun-
cated by the ridges of the costa and therefore not
decurrent (intermediates exist); veins free or (in
subgenus Soromanes) anastomosing, ending close
to the margin, hydathodes absent; hairs of two
types: 1) all species have tiny, reddish, appressed,
jointed, flattened hairs on the lamina that grade
into the scales (these actually represent reduced
scales), and 2) most species have hairs that do not
intergrade with the scales, these are whitish and
cylindrical, ranging from unicellular and less than
0.1 mm long to 15 cells and 2.0 mm long, or
uncinate. Fertile leaves appearing as a skel-
etonized version of the sterile, more ephemeral
than the sterile, produced only on the scandent
portion of the stem in hemiepiphytic species: sori
nonindusiate and of three types: 1) round, discrete,
usually at the tips of pinnately arranged stalks
(botryoid type), 2) oblong or linear formed by the
fusion of several different sori (coenosoric type),
and 3) sori apparently occupying both surfaces of
a caudate pinnule (amphiacrostichoid type); di-
plodesmic veins present except in botryoid sori;
sporangia with 64 spores per capsule, the capsule
glabrous except in P. pubens which has setose
hairs at the apex near the annulus; sporangial stalks
with three rows of cells at the apex, two-rowed
below, paraphysate in most species by lateral
hairs, these multicellular and unbranched
(branched only in P. speciosa), annulus of 15—22
indurated cells; spores (32)45—65(80) microns
long, dark brown when viewed with transmitted
light under a compound microscope, deep orange
when viewed with reflected light under a dissect-
ing microscope, monolete, aperature linear and
Vs—¥s the length of the long axis, exospore
smooth, perispore bilayered with inflated folds
and echinate to various degrees. x= 41.

November 1987 MONOGRAPH of POLYBOTRA 39

Key to the Species of Polybotrya

1. Sterile leaf 1-pinnate.
2. Veins of sterile leaf anastomosing (subgenus Soromanes).
3. Plants of Mesoamerica; vein tips joined into a faint, continuous, submarginal connecting

ESA MEE foe oc act ce Su aaciansifyaciuatecai ni aces cose sueae tea neenn tule 2. P. polybotryoides, p. 80.

3. Plants of South America and Trinidad; vein tips usually, but not always, free.

4. Lamina pubescent abaxially with uncinate hairs. Western cordillera of Ecuador.

5. Hairs on abaxial surface 0.1—0.3 mm long, colorless, erect, 1—3 celled; free pinnae pairs
6-13 below the pinnatifid apex, mostly 15—21 X 4—S cm. ........ see ee eee eee ee ee eee eens
Ree cialasairca ei Atecratas Os diate eeaiaghs eels eohiea eee uineh aes lewe ere aee anette 3. P. suberecta, p. 50.

5. Hairs on abaxial surface 0.5—1.2 mm long, tawny, spreading, 5—12 celled; free pinnae
pairs 4—7 below the pinnatifid apex, mostly 17—33 * 5.7-10 cm. oo... eee ee eee
areas teats intncie Piateiaibaieleie Ages no a SeePale pista Wintsaya dois laiecioboine sec awralnstdcs a caarsadrere 4. P. andina, p. 50.

4. Lamina glabrous abaxially.

6. Apex evenly pinnatifid, not similar to the lateral pinnae (Fig. 20a); vein tips free or in-
completely anastomosing.

7. Stem scales linear, stiffish, mostly 0.3—1.0 mm wide, usually opaque and concol-
orous; fertile pinnules more than 2.0 cm long. Colombia, Ecuador, and Peru.
S or creeisn erie lepine somal wigtoticoanaia noteubibiosiosSiamaane vapeur aecacnmesteng Soe IS UDEnecta? Ds (90;
7. Stem scales lanceolate, flaccid, mostly 1-2 mm wide, often with a dark central stripe
and lighter borders; fertile pinnules generally less than 2.0 cm long. Andes of Vene-
vAv (el Field rote V(c (cee rt See RRR A Se, RR Me Pe aoe 1. P. serratifolia, p. 43.

6. Apex subconform, similar to lateral pinnae, or with one or a few small basal lobes
(Fig. 20b,c); vein tips joined into a faint continuous submarginal connecting vein (see
text for observation of this character). Southern Mexico to Peru. ...................0.06+
Meese ois os keene aa RM clreMeta oe asics Taueeonlecmeoaombierte cee 2. P. polybotryoides, p. 80.

2. Veins of sterile leaf free (subgenus Sorbifolia).
8. Stem terrestrial and short-creeping; scales brown, essentially concolorous; fertile pinnules com-
monly round or oblong, usually less than 1 cm long.

9. Largest pinnae 7—10 times longer than broad, abaxial surface often with white or light
brown, sessile, globose glands; apex of 3—7 pinnalike lobes (Fig. 24a), these with
long decurrent bases; stem scales shiny brown, mostly transparent and denticulate; spores
ANAT s microns MOM epee wc scutes cote emotes ee eer eeselsaer gic 5. P. sorbifolia, p. 53.

9. Largest pinnae 5—7 times longer than broad, abaxial surface without glands; apex of 2 or 3
lobes, their bases not long decurrent (Fig. 25b,c); stem scales dull brown, mostly opaque
with entire or subentire margins; spores mostly 52-56 microns long. ................0.0645
Beate eer aha ead ca SE a rs esis Ais alata an aiapeleepincioaelsiestes celal 6. P. fractiserialis, p. 54.

8. Stem hemiepiphytic and long-creeping; scales reddish brown or orange, with a prominent cen-
tral stripe and lighter borders; fertile pinnules linear and usually more than | cm long. .....

ret ay aNe 3 eM Ta fear PAS atc te coos a ticbom atte Sea ESS eS 7. P. crassirhizoma, p. 58.

1. Sterile leaves 1-pinnate-pinnatifid to 4-pinnate (subgenus Polybotrya).
10. Pinnules of medial pinnae catadromic (Fig. 18).
11. Rhachis and costae pubescent, hairs 1.0—2.5 mm long. Amazonian lowland forests.

12. Lamina more than 15 cm wide at the base; petiole more than 15 cm long; largest pinna
lobes with entire margins; stem scales usually denticulate, translucent, cream to casta-
neous; fertile pinnules caudate, 3—15 mm wide, sori acrostichoid, apparently covering
both surfaces; sporangial capsules setose (Fig. 30e,f). ........ 11. P. pubens, p. 68.

12. Lamina up to 10 cm wide at the base; petiole up to 4 cm long; largest pinna lobes with ser-
rate margins; stem scales entire, dark, and opaque; fertile pinnules botryoid, sori round,
discrete, 1-3 mm wide; sporangial capsules glabrous. ..............ccccceceeeeeneeeeuens
TCE CULL DO RCC E CC COTE Te tle: POUCICT Ge ECE SOC ce ROA EERE 12. P. glandulosa, p. 71.

11. Rhachis and costae glabrous, or if pubescent, hairs shorter than | mm long. Amazonia and
elsewhere.

40

13.

13%

ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

Ultimate segments or lobes of sterile leaf 0.5—1.5 mm wide and only one-nerved; lamina often pubes-

cent on both surfaces. Andes of Colombia to Bolivia, Guyana. ....... 13. P. lechleriana, p. 71.

Ultimate segments or lobes of sterile leaf more than 1 mm wide and with several nerves; lamina rarely

pubescent on both surfaces.

14. Stem scales bright golden or yellowish and the pinnatifid portions of the pinnae or pinnules with
a single veinlet running towards the sinus arising directly from the costa or costule between the
main lobes or vein groups (Fig. 45d). Andes of Colombia to Peru. .....................2.222242-
nae, Fee Rae reece ala aoen cada vate cous cuidate wenn cntcers 26. P. altescandens, p. 97.

14. Plants without the above combination of characters.

15. Laminar margins sparsely ciliate, hairs minute, less than 0.1 mm long (Fig. 28a) and fertile
pinnules caudate, sori amphiacrostichoid, covering both surfaces of the leaf.
16. Stem scales dull brown, opaque, appressed-ascending, margins subentire, base curved

and thickened (Fig. 28h). Widespread. ..................2.220++ 9. P. caudata, p. 60.

16. Stem scales shiny reddish to castaneous, membranous, spreading, margins dentic-
ulate, base usually cordate at point of attachment (Fig. 29f). Paraguay and Brazil.

Pat cae Rat ceeeee sot cacene tet acnessvscic sete ee ace e eat eeee eee see 10. P. goyazensis, p. 66.

15. Laminar margins glabrous, or if sparsely ciliate, fertile pinnules not caudate and sori not am-
phiacrostichoid.
17. Abaxial surface of sterile leaf pubescent, hairs fine, erect, whitish, less than 0.1 mm
long and costae scaly with numerous, golden brown, tortuous scales. Panama.
ite chate aides chide sianuseas salealamadetlodsiecieconsSesed ve senessaesesasane suesedss 23. P. alata, p. 92.
17. Plants without the above combination of characters. Panama and elsewhere.

18. Plants nearly glabrous throughout, even within the grooves and pinnule mar-
gins cut less than % of the way to the costule. Costa Rica. ...........-.........+-
sniadetcaucedsoes recs deeesscansecesceaqeenesaqueewsorsticcnscceseases 20a is POMOC SEieaeae

18. Plants usually pubescent on the major axes and within the grooves; pinnules
cut more than % of the way tohe costule.

19. Lamina margins sparsely ciliate, hairs 0.1 mm long and stem scales
opaque, appressed, the base curved and thickened. Southeastern Brazil.
iddpac vocecnaesece cacaieet sds cuateanseenevececddtaessmeces = cunlU 5 ki VIEICCC Ey meas

19. Lamina margins glabrous; stem scales as above or thin, spreading, and trans-
lucent. Plants not from southeastern Brazil.

20. Pinnule bases more or less symmetrical (Fig. 35); tertiary segments often
inserted at about right angles to the costule, ovate to oblong, usually
less than 2.5 (3.0) times longer than broad.

21. Pinnules up to 2.5 X 1.2 cm, with only 5—7 segments and/or lobes;
lamina up to 55 X 26 cm, apex long-attenuate; major axes with nar-
row, dark, tortuous scales. Colombia. ..................-2.22seeeee00
CER se vase temecn shee e neers, aaroenenrehcrees 14. P. attenuata, p. 74.

21. Pinnules larger than 2.5 x 1.2 cm, with more than 5 segments
and/or lobes; lamina often larger than 55 x 26 cm, apex rarely long-
attenuate; major axes rarely with narrow, dark, tortuous scales.
22. Sterile lamina slightly reduced at the base; basal pinnae less than

3.5 times longer than broad, usually elongated basiscopically;
fertile leaves coenosoric; sori not stipitate but placed close to
the midrib.

23. Sterile lamina pubescent adaxially. Colombia. ...........
Geren ctustttoasn ac conetne sc ttm Gree et acee 15. P. stolzei, p. 76.

23. Sterile lamina glabrous adaxially.
24. Margins of tertiary segments entire to crenate, serrate
or lobed; pinnules of medial pinnae cut more than half
way to the costule for most of their length in pinnae 25
cm or longer. Mesoamerica, Ecuador, and Bolivia.
patuoune vcvcecweveeterese Cooesete 16. P. alfredii, p. 78.

November 1987 MONOGRAPH of POLYBOTRA 41

20.

24. Margins of tertiary segments entire, never crenate, serrate, or lobed; pinnules of medial

pinnae cut less than half way to the costule for most of their length. Colombia. ...

eee ee ener a ata cae orale io oaivels sac Waee de cenaaa de meeceacd 18. P. lourteigiana, p. 82.

22. Sterile lamina broadest at the base; basal pinnae 3.5—4.0 times longer than broad, not

strongly elongated basiscopically; fertile leaves botryoid; sori stipitate, stalks 1-4 mm long.

RONCHI EN a ale ce te ae cata ata ante als Stace taunt sna eepaieesenteannen 19. P. pittieri, p. 84.

Pinnule bases acroscopically prolonged (Fig. 44); tertiary segments and lobes usually inserted
obliquely to the costule, somewhat elongate and falcate, more than 2.5 times longer than broad.

25. Costal scales scattered, dark, ovate to lanceolate, flaccid (Fig. 52b); fertile leaves botryoid

(Fig. 52c). Cloud forests of northern Venezuela. ........... 32. P. canaliculata, p. 111.

25. Costal scales absent, or if present, then linear, tortuous; fertile leaves rarely botryoid.

26. Sterile lamina 3—4-pinnate-pinnatisect, ultimate segments or lobes 1-2 mm wide;

major axes on all sides with numerous, narrow (1—3 cells wide), reddish brown, tortu-

ous, spreading scales; fertile leaves botryoid. Colombia. ....................0.0e0eeeee es

CROCRCO SE PORE C SCRE EICE. CECE RET CRORE Sc On PERE C rR Cn hone eocricocon 17. P. botryoides, p. 80.

26. Sterile lamina to only 3-pinnate-pinnatifid, ultimate segments or lobes more than 1—2
mm wide; major axes lacking numerous scales, or if scales present, then not as above.

27. Lamina puberulent on both surfaces; veins prominulous. Andes of Ecuador and

BOLUVIA ihe cisasisce Jai abidehiva vesetetneanaee Mei eeiene se eetee 22. P. puberulenta, p. 90.
27. Lamina always glabrous adaxially and usually so abaxially; veins not promin-
ulous.

28. Stem scales dull brown, thick and opaque, attached across the length of the
curved and thickened base, margins subentire.
29. Tertiary segments lobed, lamina usually 3-pinnate-pinnatifid.

30. Costae evenly pilosulous, hairs less than 0.1 mm long; costular
grooves truncated by the ridges of the costae and therefore not decur-
rent; grooves glabrous or nearly so within. Andes of Colombia and
BOLIVIA Pi. Saniss oo te eee neous sezeee ears 21. P. hickeyi, p. 88.

30. Costae glabrous or with only scattered hairs; costular grooves decur-
rent into the costal grooves; grooves filled with reddish or brownish
hairs. Widespread species. .......... 28. P. osmundacea, p. 101.

29. Tertiary segments entire or only minutely serrate at the apex; lamina
mostly 2-pinnate-pinnatifid. Andes of Ecuador. ........................05

daa to oeleaiac ac tethine x tesahetagenicacbn ion eSeieg apnea enete 25. P. appressa, p. 94.

28. Stem scales reddish brown or golden, thin and translucent, attached at a cen-
tral point, base not curved or thickened, margins denticulate to erose.

31. Stem scales reddish brown; lamina with punctate, resinous glands; costal
grooves packed with protruding hairs 0.3—0.8 mm long. Guadeloupe

ANGI MartimiQue sy sicasecnescnactacnsacecek oreo 29. P. cyathifolia, p. 106.

31. Stem scales golden; lamina lacking punctate, resinous glands; costal

Prooves glabrous. Andes Of ECUAGON: h<...cpencmch orcelansse sues bnewasiasecer cies

geet es rtd as vapeteplon thee else bac ge zeus 24. P. aequatoriana, p. 94.

10. Pinnules of medial pinnae anadromic (Fig. 18).

32:

32.

Stem scales yellowish to golden; lamina usually with a single veinlet springing directly from the

costa or costule between the main lobes or vein groups (Fig. 45d). Andes of Colombia, Ecuador,

PRIDE I Tene Ree tases een Ay a cists cote acre trac ate eee eee nc cepa ita te ate 26. P. altescandens, p. 97.

Stem scales brown, castaneous, or reddish; lamina lacking a veinlet as described above.

33. Sterile leaves 2-pinnate; pinnules entire, bases symmetrical and cuneate; fertile pinnules cy-
lindrical and entire. Known only from the state of Espirito Santo, southeastern Brazil.
CEPOL STORET ESE Ee PORE ECE ECP E TEPER TE COTE CEE TES 8. P. espiritosantensis, p. 60.

33. Sterile leaves 2—4-pinnate; pinnules divided, lobed, or serrate, the bases usually prolonged
acroscopically; fertile pinnules lobed and/or divided.

34. Scales of the petiole base S—6 mm wide, broadly ovate, castaneous. Colombia. ...
2A Sc ce CNC AE OU CEE POCTE OT MOLLUS OLOP PORE OCC CO TEED 30. P. latisquamosa, p. 108.

42 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

34. Scales of the petiole base less than 4 mm wide, usually narrowly lanceolate, brown, castaneous, or
reddish.
35. Abaxial surface of lamina evenly pubescent; stem scales reddish. Southeastern Brazil.

36: Gostal|hairsiless)than’l'mmilong 7 22:..72-2---22---------eee-o--2=) S40 SPECIOSA, Ba Mase
36;, |Costal hairs: 1—2(25) ammilongy ves.csanae eee oe cee seas se seeeeeeee 35. P. pilosa, p. 117.

35. Abaxial surface of lamina glabrous; stem scales variously colored. Southeastern Brazil and else-
where.

37. Fertile leaves botryoid, i.e., all ultimate soriferous segments round and discrete, not fusing
to form an oblong or linear sorus (Fig. 52c).

38. Sterile lamina to 4-pinnate; stem scales dull to unaided eye, dark brown, with entire or
subentire margins; costal scales broadly ovate to lanceolate; costae glabrous or more
rarely pubescent; hairs short, reddish, less than 0.1 mm long; sori short-stalked (Fig.
52c). Cloud forests, Andes of Venezuela. .............. 32. P. canaliculata, p. 111.

38. Sterile lamina to 3-pinnate; stem scales shiny, dark castaneous, with denticulate-erose
margins; costal scales narrowly lanceolate to linear; costae often pubescent, hairs 0.5—
1.0 mm long, whitish; sori sessile (Fig. 51c). Lowland forests, northern Amazon basin
and Guiana Highlands. ..................:::.:eeeeeeeeeeeeeees 31. P. sessilisora, p. 108.

37. Fertile leaves coenosoric, 1.e., some or most of the ultimate soriferous segments, especially
those near the apex, fusing to form an oblong to linear sorus (Fig. 47g).

39. Abaxial surface of costules evenly hirsute, hairs less than 0.1 mm long, or if glabrous,
stem scales reddish. Southeastern Brazil.

40. Stem scales reddish; margins of tertiary segments or lobes crenate to lobed.

apintelnass elle SORE ROath ac Sate a earache de bmaeee ee abemeee sees at 34. P. speciosa, p. 115.

40. Stem scales bright castaneous; margins of tertiary segments or lobes entire. ...

1 PRO c BSS OEE REM en ee ae SHE Boe 33. P. semipinnata, p. 113.

39. Abaxial surface of costules glabrous or variously pubescent by soft whitish hairs more
than 0.1 mm long; stem scales rarely reddish. Plants not of southeastern Brazil.
41. Costal grooves glabrous within; sterile lamina mostly 2-pinnate-pinnatifid. Costa

RiG ait eS ae o.oo a 2 | ROMER ROS

41. Costal grooves pubescent within, hairs reddish to brownish; sterile lamina mostly
3-pinnate-pinnatifid.

42. Basal acroscopic segment of pinnules usually with a slight basal gibbosity on
both margins (Fig. 49c); lamina membranaceous, almost always with reddish
punctate glands abaxially. Guadeloupe and Martinique. ......................
d2s ciaweleaieleiaitaw Rhea de nev ee ctw cedeceene soeetenteeabn! LOO. CVGInOa: pelo.

42. Basal acroscopic segment of pinnules lacking basal gibbosity, usually slight-
ly reduced or oblique (Figs. 47 & 48); lamina thicker, papyraceous to charta-
ceous, rarely with reddish punctate glands abaxially. Widespread. .........
cseioe lo esbisienuecies seaceloets quevindece MeewOeee reece ceeeeeen LO. Gan ONMENCUCEUS R= aiNaan

November 1987

Species Descriptions
POLYBOTRYA subgenus SOROMANES

(Fée) Moran comb. & stat. nov.

Soromanes Fée, Mém. Fam. Foug. 2 (Hist.
Acrost.) 16. 1845. Type sPEcIES: Soromanes
serratifolium Fée = Polybotrya serratifolia
(Fée) Klotzsch.

Polybotrya section Soromanes (Fée) Klotzsch,
Linnaea 20:430. 1847.

Polybotrya subgenus Soromanes (Fée), attri-
buted incorrectly to Klotzsch by Fée, Genera
Filicum 50, invalid.

Acrostichum subgenus Soromanes Hooker,
Species Filicum 5:256. 1864. Type SPECIES:
Acrostichum caenopteris Hooker = Polybotrya
serratifolia (Fée) Klotzsch.

Sterile leaves \-pinnate; veins from adjacent
costules anastomosing, 4—6 pairs, curved ascend-
ing, the basal pair joining 4—'/ the distance from
the margin to the costa, then with an excurrent
vein to the base of the above anastomosing pair,
the distal veinlets strongly curved towards the
apex, vein tips at the margin free or, in P. polybot-
ryoides, joined by a submarginal vein. Fertile pin-
nae pectinate.

1. Polybotrya serratifolia (Fée) Klotzsch (Fig.
19, Map 1).

Polybotrya serratifolia (Fée) Klotzsch, Lin-
naea 20:430. 1847.

Soromanes serratifolium Fée, Mém. Fam.
Foug. 2 (Hist. Acrost.) 82, tab. 43. 1845.
Type: P! (3 sheets), in Bory Herbarium, cited
by Fée as “Polybotrya serrata, Galeotti, in
herb. Bory, Habitat in Mexicana republica,
Lagunetta (Galeotti). - V.S. in herb. Bory.”
but probably collected by Linden in Ven-
ezuela and later distributed by Galeotti (see
text discussion). I have selected as the lec-
totype that sheet which Fée illustrated for his
tab. 43.

Soromanes dentatum Fée, Mém. Fam. Foug.
2 (Hist. Acrost.) tab. 43. 1845. Fée’s tab. 43
was erroneously labeled as “S. dentatum” in-
stead of S. serratifolium. His tab. 43 is a
precise illustration of the lectotype of P. ser-
ratifolia.

MONOGRAPH of POLYBOTRA 43

Soromanes integrifolium Fée, Mém. Fam.
Foug. 2 (Hist. Acrost.) 82, tab. 42. 1845.
Fertile leaf only, the sterile one is that of
Cyclodium meniscioides. Type: Alexander
Braun Herbarium (B).

Botryothallus kunzei Klotzsch, Bot. Zeit.
104. 1846. nom. nudum, cited by Ettings-
hausen (1864).

Polybotrya kunzei Ettingshausen, Denkschr.
Akad. Wien. 22:66, fig. 2. 1864. Type: Only
“in Colombia” was mentioned.

Acrostichum caenopteris Hooker, Species
Filicum. 5:256. 1864. nom. nov. for Soro-
manes serratifolium Fée, non Kaulf. 1824,
with same type.

Polybotrya coenopteris (Hooker) Christ,

Farnkr. 44. 1897.

Soromanes coenopteris (Hooker) Christ,
Bull. Herb. Boissier, II. 3:613. 1903.

Acrostichum hartii Baker, J. Bot. 371. 1881.
Type: Trinidad, Hart 228 (holotype: K, photo
at US!; isotype: NY!).

Polybotrya hartii (Baker) C. Chr., Index
Filicum. 504. 1906.

Polybotrya crassa Morton, Fieldiana Bot.
28:13. 1951. Type: Venezuela. Monagas:
south-facing forested slopes above lime-
stone bluffs, northeast of Guacharo, alt. 1300
—1400 m, 11 April 1945, Steyermark 6199]
(holotype:US!; isotypes: F!, VEN!).

Stem hemiepiphytic, 1.5—3.0cm thick; scales
0.8—1.5 mm long, 0.5—2 mm wide, narrowly
lanceolate, reddish brown to light orange, with or
without a prominent dark central stripe, membran-
ous, lustrous, spreading, margins highly erose to
merely denticulate at the apex. Sterile leaves up
to 0.8 m long, dull green adaxially, pale green
abaxially, chartaceous to subcoriaceous; petiole
scaly at base, up to 30 cm long, trisulcate and
flattened adaxially; /amina up to SO X 27 cm, 1-
pinnate; pinnae 6-12 pairs, mostly (7)10—
20(23) x (2)3—4.5(5.5) cm wide, lanceolate, base
rounded, cuneate to truncate, margins entire to
serrate, apex acuminate; veins in pinnated groups
3—6 mm apart, the tips arcuate, free; axes with a
few scattered scales, rarely pubescent, the hairs
tiny, less than 0.1 mm long, unicellular, whitish;
grooves nearly glabrous within. Fertile leaves 2-
pinnate, often about the same length as the sterile

Vol. 34, Art. 1

44 ILLINOIS NATURAL HiIsTORY SURVEY

h

|

15cm

; 4

lcm
FicurE 19. Polybotrya serratifolia (Fée) Klotzsch. a. habit; b. stem scales (n
dark central stripes); c. petiole scale; d. sterile pinna; e~g. fertile pinnae showing variation in cutting;
h. sporangium with paraphysis. a—c: van der Werff & Wingfield 3430 (MO). d: Aristeguieta 3963
(VEN). e: Fendler 261 (MO). f,h: Broadway 9947 (GH). g: type, probably Linden (P).

ote erose margins and

November 1987

leaves; pinnae linear to narrowly triangular,
evenly long-tapered to apex, up to 14 2.5 cm;
pinnules entire to occasionally slightly lobate,
linear, oblong or sometimes clavate, 4—15(20) mm
long, 2—3 mm wide, the adaxial margins folding
together at maturity, giving the pinnules a cylindri-
cal appearance; sori coenosoric, continuously cov-
ering the pinnule, occasionally lobed at the base
or botryoid; sporangial stalks paraphysate; spores
(44)50—60(63) microns long. n= 41.

Other illustrations: Fée’s tab. 43 is an excel-
lent, precise illustration of the lectotype; Vareschi,
Fl. Venez., Helechos, vol. 1, tab. 71. 1969 (as
P. crassa).

Polybotrya serratifolia grows in undisturbed,
wet, premontane and cloud forests from 1200—
2400 m elevation. It is known only from the moun-
tains of northern Venezeula and Trinidad (Map
1). Van der Werff and Smith (1980) report this
species (as Polybotrya sp., aff. crassa) from the
state of Falcon, Venezuela, where it grows in wet
premontane forest with another Venezuelan en-
demic, P. canaliculata.

The distinctive, wide, flaccid, erose scales of
P. serratifolia (Fig. 19b,c) distinguish it from the
other species in subgenus Soromanes. Polybotrya
serratifolia differs from the similar P. polybot-
ryoides by its submarginal connecting vein. This
vein, however, is not easily seen because the con-
necting vein of P. polybotryoides is very faint and
the vein tips of P. serratifolia, though free, are
arcuate and simulate a submarginal connecting
vein.

Polybotrya hartii and P.. crassa are placed in
synonymy, although at first sight they look differ-
ent. Both were originally distinguished from P.
serratifolia by their more dissected (botryoid) fer-
tile leaves. But these botryoid leaves represent a
break-up of the coenosorus—a phenomenon that
I interpret as atavistic. These highly divided fertile
leaves are not uncommon; even the type at Paris
contains a sheet with a 3-pinnate fertile leaf (Fig.
19g). As might be expected, intermediates exist
between coenosoric and botryoid fertile leaves
(Fig. 19f). Still, the most common type of fertile
leaf in P. serratifolia is 2-pinnate (coenosoric)
with relatively short pinnules (Fig. 19a,e).

In addition to its finely divided fertile leaf,
Polybotrya crassa was distinguished by having an
acroscopic auricle and several lobes at the pinna
base. This condition represents nothing more than
a part-fertile, part-sterile leaf; such transitional

MONOGRAPH of POLYBOTRA 45

forms are commonly found in other species of
subgenus Soromanes and frequently cause
taxonomic confusion. Accordingly, P. crassa is
here placed in synonymy.

Fée’s citation of the type locality and collector
(“Mexicana republica, Lagunetta (Galeotti). -V.S.
in herb. Bory’’) seems to be the result of confusion.
One of the labels on the type specimen reads (my
translation): “Lagunetta, sent from Galeotti, Oc-
tober 1845.” The word Lagunetta appears without
mention of Mexico. Fée probably thought that
Galeotti had collected the specimen in Mexico,
the principal American country in which Galeotti
collected (Morton 1971:63). Galeotti returned to
Europe in 1840, where he sold many duplicates
of his own and other collections. Morton (1971)
observed: “Galeotti collected only in Mexico and
a few specimens in Cuba but is often cited as the
collector of plants from Brazil, Venezuela, and
Colombia, but these plants were actually collected
by Linden, and Galeotti was merely the distributor
of the Linden plants.” Presumably, Jean Jules Lin-
den collected the type of P. serratifolia, which
was later distributed by Galeotti. Although Linden
collected in Cuba, Mexico, and Guatemala, his
last and most profitable trip was to Venezuela and
Colombia (1841—1844) where, based on other col-
lections, this species is known to occur. Polybot-
rya serratifolia has not been collected in Mexico.
I cannot find a town called “Lagunetta” in Mexico,
but a town with that name is located in the state
of Lara, Venezuela, and I suspect that is where
Linden collected the type.

Specimens examined: TRINIDAD. Prestoe 149]
(MO), /492 (MO); heights of Aripo, Broadway 9947
(GH, US), 9949 (F, NY, US), Fay 859 (BM).

VENEZUELA. Aragua: Cordillera Interior, Cerro
E] Pauji, Topo El Pauji, al sur de El Consejo, Steyermark
& Stoddart 118051 (GH, VEN); cerca Tejerias, Vareschi
7764 (VEN); Parque Nacional “H. Pittier,” bosque de
Rancho Grande, Tschudi 167 (VEN); Colonia Tovar,
Fendler 235 (BM), 26] (GH, K, MO, NY, PH, US);
Colonia Tovar, Moritz 277 (BM). Faleén: Curimagua
(Coro), van Cotthem 1327 (UC); Sierra de San Luis,
Montana de Paraguariba, van der Werff & Wingfield
3430 (MO). Distrito Federal: between El Junquito and
Colonia Tovar, Steyermark 91756 (GH, VEN); El Jun-
quito, Schnee 6/5 (VEN); off road Caracas—Colonia
Tovar, in forest below Club Jundolandia, Berry 948
(VEN). Lara: Lagunetta, Galeotti s.n. (P); Ditto. Moran,
8.7 mi SE of Sanare, Parque Nacional Yacambu, A.R.
Smith 1259 (PORT, UC); Dtto. Moran, Rivero et al.
1608 (PORT); Dito. Iribarren, Parque Terepaina, /riarte
52 (PORT); Dtto. Andres Elroy Blanco, 7 km de Sanare,

46 ILLINOIS NATURAL HISTORY SURVEY

Rivero 513A (PORT); selva arriba de Sanare, Aris-
teguieta 3963 (US, VEN); 2200 m sobre los banos de
Rio Claro, A.R. Smith 4055. Mérida: vertientes del Rio
Capaz, arriba de La Azulita, Steyermark & Rabe 97133
(VEN); rich forest above Hacienda Agua Blanca, above
La Azulita, Steyermark 56114 (F, US). Monoagas:
south-facing forested slopes above limestone bluffs,
northeast of Guacharo, alt. 1300-1400 m, 11 April
1945, Steyermark 6199] (F, US, VEN). Portuguesa:
15 km E of Chabasquen, 67 km NNW of Guanare,
Steyermark et al. 126675 (PORT), 126680 (UC, VEN).
Trujillo: arriba de Escuque, entre Escuque y La Mesa
de San Pedro, Steyermark 104717 (MO, VEN); Dtto.
Bocono, ca. 10 mi SW of Batatal on road to Bocono,
Laguna de Aguas Negras, A.R. Smith et al. 922 (PORT,
UC); 2 km NW of Caserio Cerros de Guaramarcal, 42
km SE of Bocono, Moran 3709 (F, MO, PORT, VEN).
Yaracuy: El Amparo hacia Candelaria, a 7-10 km al
Norte de Salom, Steyermark et al. 106758 (NY, MO,
PORT); Distrito Bruzual, Serrania de Aroa, 11-15 km
NNE of Urachiche, 3 km NE of Caserio Buenos Aires,
Steyermark et al. 124749 (PORT, UC, VEN); Dtto.
Bolivar, entre las Parchitas, Tierra Fria y Ojo de Agua,
Ortega & Smith 2491 (PORT), 25/0 (PORT).

2. Polybotrya polybotryoides (Baker) Christ
(Figs. 20 & 21, Map 2).

Polybotrya polybotryoides (Baker) Christ,
Bull. Herb. Boissier, II. 1:70. 1901.

Acrostichum polybotryoides Baker, J. Bot.
207. 1881. Type: Colombia. Norte de San-
tander: Ocana, on trees in the forest, 7000
ft., Kalbreyer 1254 (holotype: K, color slide
at MO!; photo GH!, MO!).

Acrostichum juglandifolium Baker, J. Bot.
207. 1881. nom. illeg., non Kaulfuss, 1824.
Type: Colombia. Antioquia: Kalbreyer 1798
(holotype: K, color slide at MO!; photo GH!,
MO!).

Polybotrya juglandifolia Christ, Bull. Herb.
Boissier, II. 4:965. 1904. nom. nov. for
Acrostichum juglandifolium Baker, non Kaul-
fuss, with same type.

Polybotrya juglandifolia Christ var. lobata
Christ, Bull. Herb. Boissier, II. 6:168. 1906.
Type: Costa Rica. Cartago: Rio Navarro,
Coll. Inst. Costaricensis no. 16769, Wercklé
sn. (P?).

Polybotrya kalbreyeri C. Chr., Index
Filicum. 504. 1906. nom. superfl. for
Polybotrya juglandifolia Christ; with the
same type.

Vol. 34, Art. 1

Polybotrya aucuparia Christ, Bull. Herb.
Boissier, II. 6:166. 1906. TyPE: Costa Rica.
Cartago: Valley of the Rio Navarro, 1400 m,
Wercklé 16770 (P!; photos F!, NY!, UC!).

Stem 0.5—2.0cm thick, hemiepiphytic; scales
linear, mostly 9-15 mm long, 0.5—1.0 mm wide,
ascending with spreading tips, lustrous, golden
yellow when fresh, turning yellow brown to purple
brown upon drying, margins denticulate. Sterile
leaves up to 1.45 m long; petioles up to 45 cm
long; lamina up to 1.0 m, 1-pinnate, but occasion-
ally becoming pinnate-pinnatifid to 2-pinnate in
transitional sterile-fertile leaves, subcoriaceous,
dark green adaxially, pale green abaxially, nearly
glabrous on both surfaces; pinnae up to 12 pairs,
mostly 12—20(30) x 3—6(8) cm, lanceolate, the
base round, cuneate or subtruncate, with the acro-
scopic side slightly prolonged, the margins entire,
crenate or serrate, the apex acuminate; veins in
pinnate groups 5—12 mm apart, the side branches
ascending and anastomosing at acute angles with
the adjacent ones (rarely, the veins fail to anas-
tomose locally), the vein tips connected by a faint
intramarginal connecting strand; axes nearly gla-
brous or with scattered hairs, these unicellular and
colorless. Fertile leaves 2-pinnate, 12—28 cm
broad; pinnules oblong to linear, 0.9—2(2.5) x I-
2 mm; sori coenosoric, continuously covering the
abaxial surface of the pinnules; sporangial stalks
paraphysate; spores mostly (45)50—65(68) mi-
crons long. n=41.

Other illustrations: Ettingshausen, Farnkr.
fig. 2. 1864 (as P. kunzei, portion of pinnae show-
ing vein); Hooker, Icones Plant. 1877. pl. 1690
(as Acrostichum polybotryoides) and pl. 1691 (as
A. juglandifolium); Rovirosa, Pterid. Sur. Mex.
pl. 60. 1909 (as A. coenopteris); Smith, Flora of
Chiapas, part II, fig. 71. 1981 (as P. aucuparia);
Stolze, Fieldiana Bot. II. fig. 60d, 1981 (portion
of pinna with venation, as P. aucuparia).

Polybotrya_ polybotryoides grows in wet,
shaded, tropical forests from sea level to 1400
(1850)m. It has the largest range of any species
in subgenus Soromanes, even occurring on Cocos
Island (Map 2). Gomez (1976) reported this
species from Nicaragua, but I have not seen a
specimen from that country.

The shape of the leaf apex varies clinally from
southern Mexico to Peru (Fig. 21). From southern
Mexico to Costa Rica, the apex is pinnatifid, hav-

November 1987 MONOGRAPH of POLYBOTRA 47

FicurE 20. Polybotrya polybotryoides (Baker) Christ. a. northern apex form, Guatemala; b. sterile
leaf with intermediate apex form, Panama; c. southern apex form, Ecuador; d. fertile leaf; e. sterile
pinna; f. stem scales. a: Steyermark 37320 (F). b: Croat 12143 (MO). c,d: Moran 3569 (F). e,f: Moran
2178 (CR).

48 ILLINOIS NATURAL HISTORY SURVEY

ing two to four basal lobes; this apex does not
resemble the lateral pinnae. South of Costa Rica,
the apex resembles the lateral pinnae because it
has only one or two basal lobes with the remainder
of the apex relatively narrow and the margins en-
tire to crenate. Since apex shape varies clinally
and no other characters correlate with it, I interpret
the two extreme shapes as geographic variations
of the same species and do not believe that these
should be named. Plants having an intermediate
apex shape do not appear to be hybrids, since they
do not have aborted spores. In South America,
the nearly conform apex of P. polybotryoides is
taxonomically useful because the three other
species of subgenus Soromanes, which might pos-
sibly be confused with this species, have strongly
and evenly pinnatifid apices. The subconform
apex in southern populations of P. polybotryoides
is derived within the genus, as evidenced by com-
parison to all other species of Polybotrya and to
all other genera of dryopteroid ferns.

Polybotrya polybotryoides is most like P.. sub-
erecta. Another morphological character separat-
ing these two, besides apex shape, is the submar-
ginal connecting vein present in P. polybotryoides
(Fig. 20e) but only partially formed in P. suberecta
(Fig. 22d). Unfortunately, the connecting vein is
faint and difficult to see because of the thick lamina
and, in some specimens, margins that have be-
come revolute upon drying and thereby hide the
region just inside the margin. The connecting
strand is best seen with magnification on the abax-
ial surface of the pinna or with the unaided eye
and the pinna held up to strong light.

Specimens examined: Mexico. Chiapas: Tum-
bala, Rovirosa 972 (GH, PH); Finca Mexiquito, Purpus
6761 (BM, F, GH, MO, NY, UC, US), 7245 (BM,
GH, US); 18-20 km N of Ocozocoautla de Espinoza,
along road to Mal Paso, Breedlove & Smith 21886

(MICH, NY, UC).
BELIZE. Cayo: Hwy 28.5 mi S of Belmopan, Croat

24566 (CR, MO); Toledo district, Maya Mts., between
Rio Caraval and Union Camp, Boutin & Schlosser 5902
(MO).

GUATEMALA. Alta Verapaz: 7 mi up the road to
Oxec along road off Hwy. 7E between Tucuru and El
Estor ca. 6 km NE of Panzos, Croat 41637 (CR, MO).
Izabal: along Rio Bonita, Cerro San Gil, Steyermark
41698a (F, US); Cerro San Gil, Steyermark 41870 (F).
Quezaltenango: near Calahuache, Standley 67135 (F);
Finca Pireneos, below Santa Maria de Jest, Standley
68203 (F). San Marcos: above Finca El Porvenir, Vol-
can Tajumulco, up Loma Bandera Shac, Steyermark
37320 (F).

Vol. 34, Art. |

HoNpuRAS. Atlantida: Lancetilla Valley, near
Tela, Standley 53955 (F, US); Montana La Manga, 30
km SE de La Ceiba, Nelson et al. 3296 (MO); Cordillera
Nombre de Dios, Gémez 7027 (CR). Comayagua:
Quebrada Potrero, Cerro Azul de Meambar, Gémez 69/4
(CR). Cortés: mountains on N side of Lake Yojoa,
Morton 7629 (US).

COSTA RICA. Cartago: Valley of the Rio Navarro,
1400 m, Wercklé 16770 (P, photos F, NY, UC); along
Camino Raiz de Hule, SE of Plantanillo, Croat 36800
(CR, MO); Raiz de Hule, Moravia de Chirrip6, Ocampo
727 (CR); Chitaria, forest near old jailhouse, Moran
2170, 2171, 2173, 2176, 2178 (CR, F, MO, NY);
Chitaria, Valerio 329 (US), 33/33 (CR); Finca Navarro,
Maxon 639 (NY). Cocos Island: Wafer Bay, Gomez
3324 (CR, F, US); Wafer Valley, Pittier 16232 (CR,
US); Wafer Bay river valley, Holdridge 5/53 (GH),
Klawe 1545 (US); trail between Chatham and Wafer
bays, Gomez 18064 (CR); Chatham Bay, Jiménez M.
3209, 3210 (CR, F, GH, MICH); Chatham Bay, Four-
nier 357 (NY). Puntarenas: Osa Peninsula, on ndge
9.5 km W of Rincon de Osa, Mickel 2742 (NY); above
San Vito at Finca Wilson, NE of home, Evans & Bowers
3152 (MO); vicinity of biological field station at Finca
Wilson, 5 km S of San Vito de Java, Mickel 3180 (NY).
San José: Carnllo, Brade 372 (UC).

PANAMA. Coclé: Continental divide on road to Coc-
lesito, Hammel 3503 (CR). Darién: El Llano—Carti
road, Churchill & de Nevers 4993 (MO); Serrania de
Pirre, along ascent of Serrania de Pirre above Cana Gold
Mine between Rio Cana and Rio Escucha Ruido, Croat
37757 (MO), 37794 (MO). Panama: El! Llano—Carti
road, 13.7 km N of Pan-American Highway, Folsom
3590 (MO); Cerro Campana, ca. 10 km SW of Capira,
trail to summit, Mori & Kallunki 3574 (MO, NY); 1 mi
upstream from Frizzel’s Finca Indio, on slopes of Cerro
Jefe, Foster & Kennedy 1814 (F, MO); Cerro Jefe, Web-
ster et al. 16467 (UC); along road to Cerro Campana,
Croat 14687 (F, MO, NY); trail to Cerro Campana,
Kirkbride & Hayden 274 (MO, NY); Cerro Campana,
FSU Field Station, Kennedy et al. 2074 (MQ); cloud
forest on Cerro Campana above Su Lin Motel, Croar
14742 (MO); Cerro Campana, above Su Lin Motel,
Croat 4266 (MO); Cerro Campana, near FSU building,
Croat 12114 (F, MO), 12/43 (MO, US), 14786 (MO);
8-12 km N of El Llano, along new El Llano—Carti
toad, Nee et al. 8768 (MO); Cerro Campana, lower
slopes above FSU cabin, Croat 22789 (MO); Cerro Cam-
pana along trail to summit, Croat 17/6] (MO, US),
high point of ridges S of Ipeti, 5—6 hr walk from Choco
village, Serrania de Maje, Knapp et al. 4542 (MQ);
along trail to top of Cerro Campana, SW slope from
road, Nee & Stockwell 11610 (MO); Cerro Campana,
Armond 298 (MO). Veraguas: 5 mi NW of Santa Fé,
slope above Rio Primero Brazo, below Escuela Agricola
Alta Piedra, Croat 23244 (MO), Liesner 820 (MO);
Valley of Rio Dos Bocas along road between Escuela
Agricola Alto Piedra and Calovebora, 15.6 km NW of
Santa Fé, Croat 27648 (MO); 0.6 mi beyond Escuela

November 1987

Agricola Alto Piedra, Croat & Folsom 34051 (MO),
vicinity of Escuela Agricultura Alto Piedra near Santa
Fé along trail to top of Cerro Tute, Antonio 4019 (MO);
ridge of Cordillera de Tute, trail to Cerro Tute, above
Escuela Agricola Alto de Piedra, just W of Santa Fé,
Knapp & Dressler 5461 (MO).

CoLomBiA. Antioquia: Kalbreyer 1798 (K, color
slide at MO; photo GH, MO); vic. Planta Provedencia,
28 km SW of Zaragoza, Alverson et al. 342 (MO, US);
cerca de Porcesito en el valle de Rio Medellin, Hodge
6780 (GH); Barbosa, Henri-Stanislas 1709 (US).
Chocé: Trail to Miniquia E of Puerto Mutis, Lellinger
& de la Sota 39 (US); Rio Mutata ca. 3 km above
junction with Rio El Valle, NW of Alto del Buey, Lel-
linger & de la Sota 190 (US). Norte de Santander:
Ocana, on trees in the forest, alt. 7000 ft., Kalbreyer
1254 (K, color slide at MO; photo GH, MO).

MONOGRAPH of POLYBOTRA 49

Ecuapor. Bolivar: Lim6n, estrivaciones in-
feriores de la Cordillera Occidental, Acosta Solis 6358
(F). Napo: 73 km al noreste de Baeza, Cascada de San
Rafael, Moran 3593 (Q, QCA), Foster 85-162 (UC),
camino Baeza—Tena, 15 km al norte de Tena, Moran
3530 (Q, QCA). Morona-Santiago: Cordillera de
Cutucu, western slopes, trail from Logrono to Yaupi,
Madison et al. 3356 (GH). Pastaza: 6.5 km W of Mera,
Moran 3590 (F, Q, QCA). Pichincha: Hotel Tinalandia,
bosque primario al lado norte del Rio Toachi, Moran
3562 (Q, QCA). Tungurahua: 25 km al este de Banos,
detras el pueblo Rio Negro, Moran 3569 (F, Q, QCA).

PERU. Huanuco: SW slope of Rio Lulla Pichis
watershed, ascent of Cerros del Sira, Dudley 13290D
(GH). Pasco: Prov. Oxapampa, Abra los Mellizos, 4—8
km from Enanas, Skog et al. 5036 (US).

FiGure 21. Clinal variation in apex shape of Polybotrya polybotryoides (Baker) Christ. Plants from
the northern portions of the range have deeply and evenly pinnatifid apices. Southward, this shape
gradually changes into a subconform apex. Intermediates are found primarily in Costa Rica and Panama.

50 ILLiNnois NATURAL HISTORY SURVEY

3. Polybotrya suberecta (Baker) C. Chr. (Fig.
22, Map 1).

Polybotrya suberecta (Baker) C. Chr., Index
Filicum 506. 1906.

Acrostichum suberectum Baker, J. Bot. 207.
1881. Type: Colombia. Antioquia: 4000—
4500 ft., Kalbreyer 1877 (holotype: K, color
slide at MO!; photo GH!, US!).

Acrostichum hackelianum Sodiro, Anal.
Univ. Quito XII (78):21. 1895. (Crypt. Vasc.
Quit. 491. 1893.) Type: Ecuador. Pichincha:
banks of the Rio Pilaton, Sodiro s.n. (holo-
type: Q!).

Stem 1-2 cm thick, hemiepiphytic; scales
dark brown, narrow, linear, 0.2—0.4(1.0) x 8—15
mm, stiff to spreading. Sterile leaves up to 1.6 m
long; petioles about 2 to ¥% the length of the
lamina, scaly at base, becoming less so distally;
lamina 1-pinnate, mostly 0.5—1.0(1.2) m long,
lanceolate; pinnae oblong-lanceolate, (10)15—
21(25) x (3)4—5(6.5) cm, 6—13 free pairs, gla-
brous to densely pubescent beneath, the hairs un-
cinate, multicellular, reddish or clear, 0.1—0.6
mm long, the margins entire, or (more rarely)
shallowly and inconspicuously serrate, the base
cuneate to broadly rounded, the apex acuminate;
veins anastomosing, the tips sporadically uniting
to form a discontinuous, submarginal connecting
strand; axes with a few narrow, scattered scales,
usually glabrous, rarely pubescent by uncinate
hairs, or rarely by short (less than 0.2 mm), sub-
ulate, clear hairs. Fertile leaves 2-pinnate to 2-pin-
nate-pinnatifid, 10—30 cm wide, pectinate; mod-
erately scaly, the scales appressed, linear; pinnules
linear, 25—55 X 1—3 mm; sori coenosoric, con-
tinuously covering the abaxial surface of the pin-
nule; sporangial stalks paraphysate; spores (43)
46—55(58) microns long.

Other illustrations: Hooker, Icones Plant. 17,
pl. 1692. 1877 (as Acrostichum).

Polybotrya’ suberecta grows in montane
forests from 600 to 1800 m elevation. Most of the
specimens are from the western slopes of the
Andes (Map 1). These locations correspond to my
field experience in Ecuador, where this species
was common on the western side of the Andes
but absent from the eastern side.

The range of this species overlaps with P.
polybotryoides, a similar species. The two species
are most easily distinguished by the shape of the
leaf apex; P. polybotryoides has a subconform

Vol. 34, Art. 1

apex in Colombia and Ecuador, but P. suberecta
has an evenly tapering, pinnatifid apex. Other dis-
tinguishing characters of P. suberecta are the un-
cinate hairs that occur in some specimens (Fig.
22a) and the discontinuous, submarginal connect-
ing vein (Fig. 22d). Polybotrya serratifolia also
resembles P. suberecta but differs by its wide,
flaccid stem scales and a distribution that is re-
stricted to high altitude forests in Venezuela and
Trinidad (Map 1). Polybotrya suberecta \ooks
very much like P. andina—a species with which
it grows in the western Andes of Ecuador. See P.
andina for comparison.

Specimens examined: COLOMBIA. Antioquia:
4000-4500 ft., Kalbreyer 1877 (K, color slide at MO;
photo GH, US). Choco: hills above Rio Capa and Rio
Mumba, up river from Lloro, Juncosa 1467 (MO, COL);
NW side of Alto del Buey, Lellinger & de la Sota 248
(COL, LPB, US); 0.3 km E of the Ciudad Bolivar—
Quibd6 road, across the suspension bridge at km 141,
Lellinger & de la Sota 894 (COL, US). Cundinamarca:
Ojo de Agua, S side of Rio Guavio, 32 km E of Gachala,
Grant 10556 (US). Magdalena: below Valparaiso, H.H.
Smith 983 (F, GH, NY); “Cincinnati,” lower slopes of
Mt. San Lorenzo, near Sta. Marta, 1300 m, Seifriz 24
(US). Santander: along highway between Pamplona
and Bucaramanga, Munic. Tona, Corregimento Cor-
cova, Vereda la Marina, Croat 56510 (MO, UC).

Ecuapbor. Los Rios: Patricia Pilar, Dodson 7380
(COL, MO, US), Dodson et al. 8679 (MO, US).
Pichincha: banks of the Rio Pilaton, Sodiro s.n. (Q):
2.5 km E of Comejo Astorga, Moran 3546.5 (F, GH,
Q, QCA); 3 km from El Paraiso, road El Paraiso—
Saguangal, Qllgaard et al. 37823 (AAU, Q, QCA); 3
km from La Armenia, road Gualea—La Armenia,
Qligaard et al. 37859 (AAU); about 40 km WNW of
Quito, 3.5 km N of Mindo, Moran 3565 (F, GH, Q,
QCA); Tinalandia Resort, N side of Rio Toachi, about
25 km E of Sto. Domingo, Moran 356/ (F, GH. Q,
QCA); ca. 2 km N of Mindo, Hacienda San Vicente,
Foster 85-37 (UC).

PERU. Junin: Prov. Tarma, Chanchamayo, Esposto
10928 (USM).

4. Polybotrya andina C. Chr. (Fig. 23, Map 2).

Polybotrya andina C. Chr., Index Filicum 7.
1905. nom. nov. for Acrostichum insigne
Baker, non Fée 1872—73, with same type.
Acrostichum insigne Baker, J. Bot. 167.
1877. nom. illeg., non Fée 1872—73. TYPE:
Ecuador. Pichincha: “Andes of Quito,” So-
diro (holotype: K, photo GH!, US!; isotypes:
AAU!, GH!, Q!, UC!).

November 1987 MONOGRAPH of POLYBOTRA 51

a
reeef

\ rs
a
= =
15 cm
e
f
5 cm

FiGuRE 22. Polybotrya suberecta (Baker) C. Chr. a. uncinate hairs from abaxial leaf surface: b. stem
scales; c. sterile leaf; d. sterile pinna; e,f. fertile pinnae. a: @llgaard et al. 37823 (AAU). b—d: Moran
3546.5 (F). e: Smith 983 (NY). f: Dodson 7380 (US).

ILLINOIS NATURAL HISTORY SURVEY

Vol. 34, Art. 1

5cm

b

cm

1

e

d
FiGuRE 23. Polybotrya andina C. Chr. a. fertile pinna; b. sterile pinna; c. sterile leaf; d. multicellular
uncinate hairs from abaxial surface of the leaf; e. stem scales. a: Sodiro s.n. (AAU). b—e: Moran 3563 (F).

November 1987

Stem 1—1.5 cm thick, hemiepiphytic; scales
mostly 8—13 mm long, dark castaneous, shiny or
dull, opaque, margins denticulate. Sterile leaf up
to 1.0 m long; petiole up to 30 cm long, %4 to as
long as the lamina, scaly at base, becoming less
so upwards; lamina mostly 60—80 x 30—S0 cm,
deltate or broadly ovate-lanceolate, subcoria-
ceous, glabrous above, below densely pubescent
with spreading, tawny, uncinate, 4-to-10-celled
hairs, these 0.5—1.2 mm long; pinnae 17—33 x
5.7-10 cm, mostly less than 3.5 times as long as
broad, 4—7 pairs below the pinnatifid apex, mar-
gins entire, base cuneate-rounded, petiolulate,
gradually becoming sessile distally; veins anas-
tomosing, the tips uniting discontinuously; axes
pubescent, the hairs like those on the lamina. Fer-
tile leaves slightly smaller, 40—80 x 20—32 cm,
pubescent with hairs like those of the sterile
lamina; pinnae pectinate; pinnules proximally
lobulate, becoming entire apically, 3—6 mm wide;
sori coenosoric, covering most of the abaxial sur-
face of the pinnule; spore length unknown.

Sodiro (1897) noted that this species “grows
in the tropical and subtropical region up to 1200
m in the forests around Sto. Domingo de los Co-
lorados and in the Nanegal and Mindo valleys”
(translation mine). This species is very rare and
restricted in range and has been collected only
twice from the western cordillera of Ecuador west
of Quito (Map 2). I saw only three plants at the
Tinalandia site. Polybotrya suberecta and P.. poly-
botryoides also occurred at this locality and were
more abundant.

This species is most closely related to P. sub-
erecta, from which it differs by its longer and
wider pinnae, fewer (4—7) free pinnae pairs below
the apex, and spreading, multicellular, tawny hairs
on the abaxial surface (Fig. 23d). Pubescent plants
of P. suberecta occur in the region of Ecuador
where P. andina grows. Within this region, the
hairs of both species are uncinate, but in P. sub-
erecta they are much shorter (0.1—0.3 mm), 1-
or 2-celled, erect, and easily overlooked by the
naked eye (Fig. 22a). Only these two species of
Polybotrya have uncinate hairs. Because the fertile
leaf of P. andina, known only from Sodiro’s type
collection, had very few spores, I could not make
an adequate measurement of spore length.

Specimens examined: ECuApor. Pichincha:
Hotel Tinalandia, casi 25 km al este de Sto. Domingo
de los Colorados, bosque primario arriba la montana al
lado norte del Rio Toachi, 1000 m, Moran 3563 (F, Q,
QCA); “Andes of Quito,” Sodiro s.n. (K, photo GH,
US; AAU, GH, Q, UC).

MONOGRAPH of POLYBOTRA 53

POLYBOTRYA subgenus SORBIFOLIA

Moran, subgen. nov.

Type species: P. sorbifolia Kuhn, Linnaea
34:64. 1869.

Folia 1-pinnata, pinnae lineares vel lanceo-
latae, 6-10 sexies vel decies longiores quam
latiores, apice longiacuminato; venae sibi paral-
lelae pinnatae, 3—5 in quoque turmae, liberae.

Stem terrestrial and short-creeping (2 spp.) or
hemiepiphytic and long-creeping (1 sp.). Sterile
leaves \-pinnate; pinnae linear to lanceolate, 6-10
times as long as broad, apices long-acuminate;
veins in pinnate groups of 3—5, strongly ascend-
ing, free all the way to the margin. Fertile leaves
pectinate or more rarely, moniliform.

5. Polybotrya sorbifolia Kuhn (Fig. 24, Map 3).

Polybotrya sorbifolia Kuhn, Linnaea 36:64.
1869. Based on var. salicifolium Hooker and
with same type.

Acrostichum caenopteris Hooker var. salici-
folium Hooker, Species Filicum 5:257. 1864.
Type: Brazil. Pernambuco: Serra do Araripe,
Gardner 1901 (holotype: K, photo GH!; frag-
ment NY!).

Polybotrya salicifolia Lellinger, Amer. Fern
J. 62:54. 1972. Type: Colombia. Santander:
vicinity of Puerto Berrio, between Carare and
Magdalena River, 100-700 m, 8 June 1935,
Haught 1757 (holotype: US!; isotype: BM!).

Stem terrestrial, 1—2.5 cm thick; stem scales
shiny brown, linear, 0.4—1 x 10—20 mm, mem-
branaceous, spreading, the margins denticulate.
Sterile leaves up to 1.3 m; petiole 15—50 cm long;
lamina lanceolate to oblanceolate, 0.3-1.5 x
0.15—0.50 m, chartaceous, with 12-18 pinnae
pairs, the apex abruptly acuminate, its lowermost
lobes decurrent; pinnae lanceolate-acuminate,
(7.3) 10—22(25.5) x (0.7)1—3(3.5) cm, the base
truncate on its acroscopic side, cuneate on the
basiscopic side, the margins entire, crenate, usu-
ally serrate at the apex, the abaxial surface gla-
brous or glandular, the glands scattered, round,
whitish to reddish, sessile glands; veins slender,
sharply ascending, long parallel, usually oblique
to the costa, the tips free; axes with a few scattered,
appressed scales. Fertile leaves lanceolate, 0.3-
1.3 x 15—40 m, pinnate-pinnatifid; pinnae linear,
4-15 x 0.7-1.0 cm, moniliform because of the
round to oblong pinnules, these 1-3 mm wide;

54 ILLINOIS NATURAL HISTORY SURVEY

sori coenosoric, completely covering the abaxial
surface of the pinnules; sporangial stalks paraphy-
sate; spores mostly 40—47 microns long.

Other illustrations: Murillo, Cat. Illustrado
de las Plantas de Cundinamarca, 2:103. 1966 (as
P. serratifolia), Vareschi, Flora Venez.,
Helechos, vol. I, tab. 73. 1968 (as P. serratifolia),
Brade, Bradea 1(9):62, fig. 3. 1971 (as P. ser-
ratifolia), Lellinger, Amer. Fern J. 62, figs. 3, 4,
10. 1972 (as P. salicifolia).

The distribution of P. sorbifolia is spotty but
extensive (Map 3). This species looks like P. frac-
tiserialis, see that species for comparison. Unlike
all other species in the genus, both P. sorbifolia
and P. fractiserialis are terrestrial—never climb-
ing—and it is doubtful that they could climb be-
cause their stems are only short-creeping. Both
species grow in shaded, rocky habitats along
streambanks, from 0—1200 m. I collected P. sor-
bifolia in Costa Rica at the El Rodeo site. It is
common at this locality, growing on talus along
a stream in the bottom of a humid, limestone ravine
that is surrounded by dry uplands.

Unfortunately, the specific epithet “salici-
folia” cannot be used for this species even though
its pinnae bear a strong resemblance to leaves of
certain Salix species, most notably S$. nigra and
S. amygdaloides. This resemblance was noted by
Hooker (1864) and Lellinger (1972).

Specimens examined: Costa Rica. San José:
Alajuelita, Alfaro 8073 (US); El Rodeo, Moran 3145
(CR, F, GH, MO, PORT), Knight s.n. (US), Gomez
7122 (CR), Hunnewell 16514 (MICH).

VENEZUELA. Aragua: Parque Nacional “H. Pit-
tier,” Rancho Grande, Tschudi 162 (VEN), Steyermark
et al. 95827a (US). Portuguesa: Dtto. Araure, Fila San
José, al oeste de Sta. Lucia, Ortega & Grimann 2707
(PORT). Yaracuy: “La Enjalma” al sur de Chivacoa,
Vareschi & Pannier 2636 (US, VEN).

CoLomBia. Boyaca: valle del Rio Cusiana, entre
Pajarito y Guazul, Murillo 1457 (COL), 1491 (COL).
Cundinamarca: entre Nilo y la quebrada de Agua de
Diosito, Murillo et al. 289 (COL, US). Magdalena:
Santa Marta, near Jiracasaca, H.H. Smith 1052 (F, GH,
L, MICH, MO, NY, PH, US). Meta: along Cono Rosa
Blanca, a small stream outside of Villavicencio,
Kirkbride 404 (MO, NY). Santander: vicinity of Puerto
Berrio, between Carare and Magdalena River, Haught
1757 (BM, US).

BraziL. Goias: Serra Dourada, 17 km (straight
line) S of Goids Velho, 6 km NE of Mossamedes, An-
derson 10152 (NY). Para: Serra dos Carajas, Serra

Vol. 34, Art. 1

Norte, near waterfall near AMAZ Exploration Camp,
Silva et al. BG 526 (AAU, F, GH, MICH, NY, UC).
Pernambuco: Serra de Araripe, Gardner 190] (NY,
photo of K specimen at GH). Roraima: Indian trail
from Surucucu to Uaica, Maita Mts. Prance et al. 10466
(NY).

6. Polybotrya fractiserialis (Baker) J. Smith
(Fig. 25, Map 3).

Polybotrya fractiserialis (Baker) J. Smith,
Hist. Filicum 133. 1875.

Acrostichum fractiseriale Baker, Synopsis
Filicum 414. 1868. Type: Peru. San Martin:
“in sylvis montis campana, terrestre,” Dec.
1855, Spruce 4337 (holotype: K, color slide
at MO!; photo GH!, US!).

Acrostichum plumbicaule Baker, Synopsis
Filicum 413. 1868. Type: Peru. San Martin:
Terapoto, Ad rupes secus rivularum, Aug.
1855, Spruce 4090 (lectotype: K, photo GH!,
photo and fragment P!, US!).

Polybotrya plumbicaulis (Baker) J. Smith,
Hist. Filicum 133. 1875.

Stem terrestrial, 1—-1.5 cm thick, short-creeping
with internodes 1—3 cm long, apex scaly, behind
the apex usually naked or only sparsely scaly;
scales dull brown, opaque, 0.3—0.9 X 8-17 mm,
ascending, margins entire to more rarely denticu-
late. Sterile leaves to 1.4 m tall; petiole about
equaling the lamina, stramineous to lead-gray:
lamina slightly reduced or broadest at the base,
chartaceous to subcoriaceous, apex with one or
two major basal lobes, merely crenate-lobulate
above; pinnae mostly 9-15 pairs, linear to oblong,
(11)15—25(28) X (2.1)2.5—4.0(4.5) cm, margins
serrate, especially towards the acuminate apex,
base rounded to cuneate, with the acroscopic mar-
gin usually more oblique; veins numerous, fine,
and in closely parallel pinnate groups, occasion-
ally with a simple vein springing directly from the
costa; axes usually glabrous or with fine, subulate,
0.1—0.2 mm hairs. Fertile leaves erect, 2-pinnate,
commonly taller than the sterile, to 1.5 m long:
pinnae linear, mostly 7—12(17) X 5—10(15) cm;
pinnules round, oblong, or linear, |—3 mm wide;
sori coenosoric, completely covering the abaxial
surface of the pinnule; sporangial stalks paraphy-
sate, often with a bulbous glandular cell at base
of the paraphysis; spores (48)52—56(60) microns
long.

MONOGRAPH of POLYBOTRA 55

November 1987

Ficure 24. Polybotrya sorbifolia Kuhn. a. sterile leaf; b. stem; c. fertile leaf; d. sterile pinna; e. stem

scales. a—e: Moran 3145 (F).

56 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

FiGure 25. Polybotrya fractiserialis (Baker) J. Smith. a. fertile leaf; b. apex of sterile leaf; c. sterile
leaf; d. stem scales; e. sterile pinna; f. stem. a: C. Schunke 666 (F). b,d,e: Moran 3536 (F). c: Gonggrijp
& Stahel 3300 (MO). f: Tryon & Tryon 5221 (GH).

November 1987

Other illustrations: Brade, Bradea 1(9):62,
fig. 4. 1971 (as P. polybotryoides).

This species, like P. sorbifolia, is terrestrial
and grows in wooded talus slopes, usually along
rocky streamsides. The altitudinal range of P. frac-
tiserialis is from 200-1500 m, but about 80 per-
cent of the collections come from the 400—800 m
range. Although principally a forest species, I have
seen vigorous fertile plants growing along sunny
streamsides in Ecuador, an observation which
suggests that this species has more tolerance to
higher light conditions than its cogenerics.

The range of P. fractiserialis occupies two
discontinuous regions: 1) the Andes from Bolivia
to Ecuador, and 2) the Guianas (Map 3). A dis-
tance of 900 km separates the northernmost popu-
lation in Ecuador from the westernmost population
in British Guiana. All populations of this species
occur on the eastern side of the Andes. Their ab-
sence from the western side is probably due to the
extreme dryness that characterizes that side of the
Andes south of the equator.

Polybotrya fractiserialis and P. sorbifolia
look very much alike and can be easily confused.
Several characters, however, distinguish the two.
The easiest feature to recognize is the shape of
the medial pinnae: those of P. fractiserialis are
relatively shorter and broader than those of P.
sorbifolia (see key and descriptions); this stoutness
is accentuated just below the apex (Fig. 25). The
apex of P. sorbifolia is another difference; it has
more numerous, decurrent, narrower pinnae (Fig.
24). Usually, the veins of P. sorbifolia are at a
more oblique angle to the costa than those of P.
fractiserialis. Another difference, albeit more rec-
ondite, is found in the sporangial stalks of P.
fractiserialis, which have a lightbulb-shaped gland
at the base of the paraphysis (Fig. 11f,g); P. sor-
bifolia lacks such a gland (Fig. 11b). In fact, no
other species of Polybotrya has a glandular cell
on the paraphysis or elsewhere on the sporangium.
Polybotrya sorbifolia, however, has round, sessile
glands on the lamina; such glands are lacking on
P. fractiserialis.

Spore size is yet another diffence: P. frac-
tiserialis has much larger spores than P. sorbifolia.
This difference suggests that P. fractiserialis may
be a polyploid, but chromosome counts are still
wanting for the two species. Polyploidy has not
yet been documented cytologically in Polybotrya.

Polybotrya fractiserialis and P. plumbicaulis
were described at the same time, in the same pub-
lication, and were distinguished on the basis of

MONOGRAPH of POLYBOTRA 57

leaf size, number of pinnae pairs, and two versus
three or four veinlets in each pinnate group. These
characters, however, intergrade completely even
on leaves from a single plant. I chose the name
P. fractiserialis, rather than the simultaneously
published name P. plumbicaulis, because it was
used by Posthumus (1928) in his treatment of
Surinam ferns.

Specimens examined: FRENCH GUIANA. Guy-
ane: Cayenne, 1847, Leprieur s.n. (P). Inini: Cocao,
La Comte Riv., 1847, Leprieur s.n. (P); Tumuc Aumac,
Haut Itany, Hoock s.n. (P); ca. 45 km SE of Saul,
Granville 3674 (CAY, Z); region de Paul Isnard, Mon-
tagne Lucifer, Granville 5219 (CAY, Z); Saul, Monts
du Fumée, Granville 512] (CAY, Z).

SURINAM. 3 km S of Juliana Top, 12 km N of
Lucie Riv., Maguire et al. 54384 (MO, NY), Irwin et
al. 54669 (NY), 54784 (F, MO, NY); Brownsberg,
Gonggrijp & Stahel 3300 (MO, US).

GUYANA. Essequibo: Southern Pakaraima Mts.,
Kopinang Falls, Maguire et al. 46068-A (NY).

ECuADoR. Napo: casi 10 km SE de Tena, 3 km
E de Puerto Nuevo por camino a Puerto Misahualli,
Moran 3536 (F, GH, MO, Q, QCA, US). Pastaza: Rio
Capihuan, tributary of Rio Pastaza, @llgaard et al.
35124 (AAU).

PERU. Amazonas: Prov. de Bagua, Valley of Rio
Maranon above Cascadas de Mayasi near Campamento
Sta. Montenegro (kms. 280-284 of Maranon rd.), Wur-
dack 1854 (US). Cuzco: Prov. Paucartambo, Vargas
11280 (GH). Huanuco: Prov. Hudnuco, Tingo Maria,
Tryon & Tryon 5221 (GH); Tingo Maria, Allard 20661
(US), 22305 (GH, US), 22526 (US); Previsto, antes de
Aguaitia, Aguilar 947 (USM); Prov. Leonicio Prado,
Distrito Rupa Rupa, al este de Tingo Marfa, cerca al
Cerro Quemado, J. Schunke 10173 (MO); hills E of
Tingo Maria, Croat 2/152 (MO). Junin: Prov. Satipo,
Pichanaki, Leén 226 (USM); Schunke Hacienda, above
Ramon, C. Schunke A214 (US); Chanchamayo Valley,
C. Schunke 165 (F, US), 666 (F), 1396 (F). Loreto:
above Pongo de Manseriche, Mexia 6246a (UC, US);
Aguaytia, Croat 2/0/4 (MO). Madre de Dios: Prov.
Manu, town of Atalaya, 2-3 km W of village, Foster
7455 (F, MO, USM); Pantiacolla, serranfa across Rio
Alto Madre de Dios from Shintuya, Gentry et al. 27365
(MO); prov. Manu, km 139 on road between Cabo de
hornas and Shintuya, Foster 4020 (F, USM); Parque
Nacional de Manu, Cocha Cashu Biological Station,
Foster P-84-9] (F); valle de Marcapata, Herrera 1/200
(US). Oxapampa: Prov. Pasco, Iscoazin, Foster 7937
(F, USM). San Martin: Mt. Campana, Spruce 4337
(color slide of K specimen at MO; photo GH, US);
Tarapoto, Ad rupes secus rivularum, Aug. 1855, Spruce
4090 (photo of K specimen at GH; photo and fragment
P, US); Prov. de Lamas, beween Tarapoto and
Moyobamba, ca. 10 km NW of Tabalosas, Croat 5//54
(MO).

58 ILLINOIS NATURAL HISTORY SURVEY

Bo.LiviA. La Paz: Cordillera Real, Rio Chimate,
Tate 502 (NY), 513 (NY); Mapiri, Williams 1188 (NY,
US); Mapiri, Buchtien 1015 (US), 2138 (US); Mapiri
region, San Carlos, Buchtien 13 (US); Mapiri,
Charopampa, Buchtien |] (MICH, P). Proy. unknown:
Challana Riv., Cardenas 1249 (LIL, US).

7. Polybotrya crassirhizoma Lellinger (Fig.
26, Map 4).

Polybotrya crassirhizoma Lellinger, Amer.
Fern J. 62:49, figs. 1, 8. 1972. Type: Peru.
Loreto: Gramitanacocha, Rio Mazan, alt.
100-125 m, J. Schunke 268 (holotype: US!;
isotypes: F!, GH!, NY!, UC!).

Polybotrya macbridei Lellinger, Amer. Fern
J. 62:51, figs. 2, 9. 1972. Type: Peru. Junin:
Hacienda Schunke, La Merced, Macbride
5602 (holotype: US!; isotype: F!).

Stem 1-2 cm thick, hemiepiphytic; scales 10—
25 x 0.2-1.1 mm, dull orange or reddish brown,
rarely shiny and dark brown, with a dark central
stripe and lighter borders, rarely concolorous,
margins denticulate to erose. Sterile leaves up to
1.35 m; petiole 2 to V4 the length of the lamina,
scaly at base; /amina lanceolate, to 1.0 x 0.44 m;
pinnae 6-20 pairs, (11)15—20(24) x 2—4 cm, lan-
ceolate to oblong-lanceolate, the base unequal,
broadly rounded or truncate on the acroscopic side,
slightly excavate or tapering on the basiscopic
side, the margins crenate-dentate or more rarely
lobate-serrate , apex acuminate, often serrate; veins
free, often with a single veinlet springing from
the costa between the main pinnate groups; axes
with a few, scattered, narrow, appressed scales,
occasionally pubescent, the hairs short, less than
0.1 mm, erect, colorless, subulate, unicellular.
Fertile leaves 2-pinnate, occasionally 2-pinnate-
pinnatifid in large individuals, 2 to “3 smaller
than the sterile leaves; axes similar to those of the
sterile leaves but with larger more numerous
scales; sori coenosoric, completely covering the
abaxial surface of the pinnules, occasionally dis-
crete and round at the pinnule base; sporangia
nonparaphysate; spores (48)52—56(61) microns
long. n=41.

Other illustrations: See original descriptions
of P. crassirhizoma and P.
above.

macbridei, cited

Polybotrya crassirhizoma grows primarily in
lowland forests of the western Amazon basin and
in premontane forests of the eastern Andes (Map

Vol. 34, Art. 1

4) from 100 to 1500 m. The cluster of dots in
eastern Ecuador reflects recent collecting activity
in that area and this species’ true abundance there.
I saw this species in nearly every forest that I
visited in the Amazon basin of Ecuador, and it
was one of the most frequent and abundant ferns.
This abundance suggests that P. crassirhizoma
occurs farther east in the Amazon basin than is
shown in Map 4. Apparently, P. crassirhizoma
flourishes on clayey soils and does not grow in
sterile, sandy soils.

I chose the name P. crassirhizoma over P.
macbridei because of the more widely distributed
type collection. Polybotrya crassirhizoma and P.
macbridei were said to differ by stem thickness,
scale luster and color, and the presence versus the
absence of lobing at the base of the sterile pinnae
(Lellinger 1972). These characters, however, in-
tergrade completely as shown by a series of speci-
mens collected near Coca in the Amazon basin of
Ecuador (Moran 3616). The most obvious charac-
ter that supposedly distinguishes P. macbridei is
the basally lobed pinnae, but this character repre-
sents the lobing that typically occurs in transitional
sterile-fertile leaves. This example illustrates how
part-fertile leaves can cause taxonomic confusion.
Accordingly, I placed P. macbridei in synonymy.

Specimens examined: COLOMBIA. Amazonas:
Loreto-Yaco, Schultes & Black 46-266 (GH).

Ecuapor. Napo: 27 km SE of Coca, petroleum
well Auca 4, Moran 3616 (F, GH, NY, Q, QCA, US);
20 km NE of Coca, 5 km N of Joya de las Sachas,
Moran 3615 (F, GH, NY, PORT, Q, QCA, US); about
10 km SE of Tena, 3 km E of Puerto Nuevo on road to
Puerto Mishahualli, Moran 3535 (F, GH, NY, Q, QCA,
US); Bimbino, on the Rio Pacuno, 10 hr W of confluence
with Rio Napo, Whitmore 752 (BM); Rio Cuyabeno,
about 2 km upstream from Puerto Bolivar, Brandbyge
et al. 33684 (AAU); San Pablo de los Secoyas,
Brandbyge & Asanza C. 3288] (AAU); San Pablo de
los Secoyas, Rio Wai si aya, Brandbyge et al. 32617
(AAU); San Pablo de los Secoyas, on path to
Shushufindi, Brandbyge et al. 32544 (AAU); 50 km NE
of Coca, Lugo S. 3293 (GH); Canton Putumayo, Rio
Aguarico, town of Dureno, Plowman et al. 4025 (GH);
Anagua, Parque Nacional Yasuni, SEF forest project
area, Ollgaard et al. 38836 (AAU), 39084 (AAU). Pas-
taza: Curaray, Valle de la Muerte, Holm-Nielsen et al.
22484 (AAU, Q, QCA); basin of Rio Pastaza, Pacayacu-
Sarayacu region, Gill 47 (NY); village of Rio Chico, 8
km from Puyo, Shemluck 280 (F); Rio Bobonaza, be-
tween oil exploration camp Chichirota and Destacamento
Cabo Pozo, Qlilgaard et al. 34855 (AAU, Q, QCA);
Rio Bobonaza, between Cachitama and the outlet of Rio
Bufeo, @llgaard et al. 34748 (AAU), Rio Pastaza, near
the Peruvian border, around Destacamento Ishpingo,

November 1987 MONOGRAPH of POLYBOTRA 59

15 cm

5 cm
11cm

FIGURE 26. Polybotrya crassirhizoma Lellinger. a. fertile leaf; b. sterile leaf; c. fertile pinna; d. sterile
pinna; e. stem scales. a: Ollgaard et al. 34855 (AAU). b—e: Moran 3615 (F).

60 ILLINOIS NATURAL HISTORY SURVEY

Allgaard et al. 34970 (AAU); 2 km W of Yuralpa, S
border of Rio Napo, Holm-Nielsen & Jeppsen 987
(AAU).

Peru. Huanuco: Prov. Pachitae, Rio Pozouso,
Foster 9284 (F, MO, USM). Junin: Chanchamayo Val-
ley, C. Schunke 157 (F), 158 (F), 661 (F), 8/2 (F);
Schunke Hacienda, above San Ramon, Killip & Smith
24605 (NY, US); E of Quimiri Bridge, near La Merced,
Killip & Smith 23876 (NY, US); Hacienda Schunke, La
Merced, Macbride 5602 (F, US); La Merced, Chan-
chamayo, Soukup 1065 (F). Loreto: Prov. Maynas,
Quebrada Yanomono, Explorama tourist camp, Rio
Amazonas above mouth of Rio Napo, Transect 6, Gentry
et al. 27880 (MO), Moran 3640, 3641 (F, Q, QCA);
Mishuyacu, near Iquitos, Klug 1386 (NY, US);
Gamitanacocha, Rio Mazan, J. Schunke 268 (F, GH,
NY, UC, US), /4285 (US); Veradera de Mazan, Croat
20763 (MO); Rio Napo near Entrada de Isla Inayuga,
Croat 20551 (MO).

Bo.iviA. Cochabamba: Prov. Carrasco, conflu-
ence of Rio Leche with Rio Isarsama, Beck 1635 (LPB).

Brazit. Acre: Cruzeiro do Sul, vicinity of Serra
da Moa, Prance et al. 12180 (NY).

8. Polybotrya espiritosantensis Brade (Fig. 27,
Map 21).

Polybotrya espiritosantensis Brade, Rodri-
guesia 10:28, tab. 3. 1948. Type: Brazil. Es-
pirito Santo: Municipality of Itaguagu, Jati-
boca, virgin forest, 700-800 m, A. C. Brade
18224 (holotype: RB!).

Stem 1-2 cm wide, hemiepiphytic; scales
mostly 10-15 x 1.0—2.5 cm, bright reddish brown,
spreading, membranous, the center often darkened
to varying degrees, margins highly erose-denticu-
late, occasionally fimbriate. Sterile leaves up to
1.2 m long; petiole to 30 cm long, about half as
long as the lamina, scaly at base; lamina ovate,
2-pinnate proximally, becoming 1-pinnate dis-
tally, 60—90 x 50-70 cm, subcoriaceous, pale
green, nearly glabrous on both surfaces, the apex
subconform and pinnalike but with one or two
basal lobes; pinnae alternate, 6—8 pairs, the apex
conform, like the lateral pinnules, the upper pinnae
simple and resembling the pinnules of the lower;
pinnules anadromic throughout, 3—4 pairs, ellip-
tic, 10-13 x 2-3 cm, the margins entire, the apex
acuminate, the base cuneate, the proximal pin-
nules with stalks 2-4 mm long, the distal pinnules
sessile; veins slender, none conspicuously thicker
than the others, long, parallel, 1-2 branched, the
tips free; axes with a few scattered, appressed,
narrow scales, otherwise glabrous. Fertile leaves

Vol. 34, Art. 1

2-pinnate, amphiacrostichoid; pinnules caudate,
entire, appearing cylindrical when mature; sporan-
gial stalks paraphysate; spores (52)54—60(62)
microns long.

Other illustrations: Brade’s original descrip-
tion has an excellent photograph of the type;
Brade, Bradea 1:67, tab. 6, fig. 5. 1971 (stem
scales only).

Polybotrya espiritosantensis is endemic to the
state of Espirito Santo in southeastern Brazil (Map
21). I expect this species will eventually be found
in other parts of mountainous coastal Brazil.
Polybotrya espiritosantensis has the most distinc-
tive laminar cutting of any species in the genus.
No other species has the simple, entire pinnules
that evenly taper at the base and apex (Fig. 27b).
Unlike all other species of Polybotrya that have
pinnatifid apices, the leaf and pinnae apices of
this species are conform or nearly so (Fig. 27a).
The numerous, close, long, parallel veins (Fig.
27b) are like those of the 1-pinnate species P.
fractiserialis and P. sorbifolia and probably indi-
cate a close relationship.

Specimens examined: BRazit. Espirito San-
to: Santa Thereza, 900 m, Foster & Foster 854 (GH,
US); Municipality of Itaguagu, Jatiboca, virgin forest,
700-800 m, A. C. Brade 18224 (RB).

POLYBOTRYA subgenus POLYBOTRYA

TYPE SPECIES: P. osmundacea Willd., Sp. Pl.
ed. 4, 5:99. 1810.

Polybotrya subgenus Eupolybotrya Fée, Mém.
Fam. Foug. (Hist. Acrost.) 2:16. 1845.

Acrostichum section Polybotrya Hooker,
Species Filicum 5:244. 1864.

Stem hemiepiphytic; sterile leaves 1-pinnate-
pinnatifid to 4-pinnate; veins free.

9. Polybotrya caudata Kunze (Fig. 28, Map 5).

Polybotrya caudata Kunze, Linnaea 9:23.
1834. Type: Peru. Hudanuco: Pampayaco,
July 1829, Poeppig s.n. (B!).

Polypodium adiantoides Aublet, Hist. Pl.
Guiane Francoise 2:962. 1775, nomen illeg.,
non Burm. (1768). Type: Guyana. Aublet
s.n. (holotype: BM!, Morton photo 6626 at
B!, F!, GH!).

=
a
<
x
8
Z

3
=
5

62 ILLINOIS NATURAL HISTORY SURVEY

Olfersia caudata (Kunze) Kunze, Linnaea
21:206. 1848.

Psomiocarpa caudata (Kunze) Presl, Epim.
Bot. 162. 1849.

Acrostichum caudatum (Kunze) Hooker,
Species Filicum 5:244. 1864. nomen illeg.
non Hooker (1840).

Polybotrya acuminata Kaulfuss var. villosa
Christ, Prim. Fl. Costar. 3(1):9. 1901. Type:
Costa Rica. Limon: Llanuras de Santa Clara,
500 m, Biolley 10688 (lectotype: US!;
isotype: CR!).

Polybotrya villosula Christ, Bull. Herb. Bois-
sier, II. 6:168. 1906. TyPE: Costa Rica. Santa
Clara, Las Delicias, 1897, 500 m, Pittier
10688 (P?).

Polybotrya costaricensis Brade, Bradea 1:11,
tab. 1, fig. 1. 1969. Type: Costa Rica. Limon:
Hundrisser Ranch, Atlantic shore, Sept.
1909, Brade & Brade 374 (holotype: HB;
isotypes: NY!, UC!).

Stem 0.5—2.5 cm thick, hemiepiphytic; scales
linear-lanceolate to narrowly triangular, 8—20 x
1—2 mm, dull brown, opaque, entire to denticulate,
the base elevated, thickened, curved, and ap-
pressed. Sterile leaves up to 2X 1 m; petioles
30-70 cm long, about 4 the length of the lam-
ina; /amina mostly 2-pinnate-pinnatifid, rarely 3-
pinnate at the base, firm-chartaceous, glabrous to
pilose, the hairs 0.2—1.5 mm long, acicular, the
margins sparsely ciliate, the hairs minute, less
than 0.2 mm long; pinnae up to 20—45(60) x 7—
20(30) cm, subdeltate, acuminate, subequilateral,
slightly more developed on the acroscopic side;
pinnules slightly prolonged acroscopically, the
base truncate to slightly cordate, catadromically
arranged in the medial pinnae; veins free, some-
times with a single vein springing from the costa
between the pinnate groups; axes nearly glabrous
or pubescent to various degrees, the hairs acicular,
whitish; grooves usually pubescent within, often
densely so at the junctures. Fertile leaves similar
in size to the sterile, mostly 2-pinnate, but with
some of the larger pinnules lobed at base; pinnae
caudate, 4—8(11) < 0.5—1.5 cm, apparently sorif-
erous on both surfaces; spores (43)46—50(53) mi-
crons long. n= 41.

Other illustrations: Fée, Mém. Fam. Foug.
(Hist. Acrost.), Atlas, tab. 34, 1845; Vareschi,
FI. Venez., vol. 1, tab. 73. 1968; Brade, Bradea
1(9):63, fig. 1; p. 67, fig. 6.; Croat, Fl. Barro

Vol. 34, Art. 1

Colorado Is., figs. 28 & 29. 1978 (as P. villosula);
Stolze, Fieldiana Bot. n.s. 6, fig. 60a,b. 1981;
Tryon & Tryon, Ferns & Allied Plants, figs. 80.8,
80.9, 80.22, 80.23. 1982.

Polybotrya caudata is one of the most widely
distributed species in the genus (Map 5). It grows
primarily in lowland forests from sea level to 800
m, but specimens from the Andes have occasion-
ally been collected as high as 1900 m. I found
this species at many of the sites where I collected
in Costa Rica, Ecuador, and Peru, but not in Ven-
ezuela. Polybotrya caudata is one of only three
species in the genus that occur on an oceanic is-
land—Cocos Island, about 500 km (310 mi)
southwest of Costa Rica.

Polybotrya villosula was said to differ from
P. caudata only by its long and villous pubes-
cence, but specimens are usually glabrous or fully
pubescent, with few intermediate forms. Since
pubescence does not correlate with any other
character and I found no difference in geographical
range between the two forms, I regard P. villosula
as a synonym of P. caudata.

The juvenile leaves in glabrous plants of this
species may be difficult to distinguish from P.
osmundacea. A good character to separate the two
species is the minutely and sparsely ciliate leaf
margin of P. caudata (Fig. 28a); that of P. osmun-
dacea is always glabrous (Fig. 47). The thick,
dull brown, curved, and appressed stem scales
(Fig. 28b,h) also help distinguish P. caudata from
many other Polybotrya species.

Specimens examined: Mexico. Chiapas: east-
erm highlands near Guatemalan border, Breedlove 3410]
(CAS).

GUATEMALA. Alta Verapaz: near Rio Icvolay,
near Hacienda Yaxcbnal, 5 mi NW of Cubilquitz, 250—
300 m, Steyermark 44675 (F); Cubilquitz, 350 m,
Tuerckheim 148 (US), 358 (P, Z), 78/2 (US), 8040B
(US), 8047 (GH, NY). Izabal: between Bananera and
“La Presa” in Montana del Mico, Steyermark 38229 (F),
38271 (F), 39197 (F); Rio Chacon, 30 m, Johnson 122]
(US); between Los Amates and Izabal, Sierra del Mico,
Kellerman 7354 (F, NY); along Rio Frio, 65 m, Steyer-
mark 39895 (F, US); 2.5 mi N of Rio Dulce on gravel
road to Petén, Dunn & LeDoux 22005 (MO, NY); vi-
cinity EXMIBAL Camp 2 (La Gloria), NW of Lake
Izabal, 400—500 m, Jones & Facey 3246 (NY); vicinity
of Quirigua, Standley 24195 (GH); vicinity of Puerto
Barmios, Standley 25085 (GH, US).

Be.ize. El Cayo: Vallentin, Lundell 6416 (GH,
US). Stann Creek: along road and stream at Dry Creek,
near Dist. of Cayo Border, Croat 2454] (CR, MO);
Middlesex, Gentle 2947 (GH, MICH, US); Schipp 273
(BM, F, GH, MO, UC, US, Z).

November 1987 MONOGRAPH of POLYBOTRA 63

=

t
is
NGS

FiGuRE 28. Polybotrya caudata Kunze. a. leaf margin showing minute cilia; b. stem and basal pinna
(note appressed scales—same scale as f); c—f. acroscopic pinnules, pinnae apex toward the right in
all; g. fertile leaf; h. stem scales, abaxial view at left, side view in center, adaxial view at right.
a,c: Costa Rica, Scamman 7156 (GH). b: French Guiana, Cremers 7369 (CAY). d,g,h: Costa Rica,
Moran 2186 (F). e: Bolivia, Steinbach 7499 (MO). f: Trinidad, Fendler 105 (GH).

64 ILLINOIS NATURAL HISTORY SURVEY

HonburRAS. Atlantida: Lancetilla Valley, near
Tela, Standley 53983 (F, US); near Ceiba, 400 m, Dyer
A225 (US).

NICARAGUA. Chontales: 4 km al oeste de La Liber-
tad, El Comatillo, Gémez & Cittar 6731] (CR); vicinity
of La Libertad, 500-700 m, Standley 9047 (F). Zelaya:
road to Mina Nueva Americana, Stevens & Krukoff
12666 (CR, MO); along new road from Rio Blanco to
Rio Copalar, Stevens & Krukoff 12056 (CR, MO); Cano
Aamora on Rio Rama, Stevens & Krukoff 8827 (CR,
MO); base camp 3.6 km SE Cerro San Isidro, Rio Kama,
Rio Escondido, 0—65 m, Proctor et al. 27091 (F, NY,
VEN); Mosquito Coast, Schramm s.n. (US); Bluefields,
Niell 2610 (CR); area de la Bahia de Bluefields, Rio
Escondido, 0-30 m, Molina R. 1907 (F, US); near
Bluefields, Danneberger s.n. (US); Braggman’s Bluff,
Englesing 29] (F, US); Mina Nueva Americana, 11.3
km N of main road, Pipoly 5299 (CR, MO); 13 mi
above Kururia, Pipoly 3824; Colonia Kururia, Pipoly
3880 (CR, MO), 390] (CR, MO); Neptune Mining Co.,
Stevens & Krukoff 13005 (CR, MO); near junct. of road
to Alimidkanba with road between El Empalme and
Limbiaka, Stevens & Krukoff 12741 (CR, MO); Apz. 5
km de Rama sobre el Rio Rama, Gomez & Cittar 6409
(CR); El Salto along Rio Pis Pis, Pipoly 3542 (CR,
MO); 13 mi above Kururia on road to San Jeronimo,
Pipoly 3817 (CR, MO).

Costa Rica. Cartago: near Turrialba, slope of
the Rio Reventazon behind the Instituto Interamericano
de Ciencias Agricolas, 600 m, Mickel 3368 (NY); Valley
of Rio Reventazon 9 km ENE of Turrialba near Pavones,
650 m, Holm & Iltis 200 (F, NY). Cocos Island: Barclay
2199 (BM); Wafer Bay, Gomez 3354 (CR, F, US, Z);
Gomez 4528 (CR); Chatham Bay, Jiménez M. 3200 (CR,
GH). Heredia: Holdridge’s Finca La Selva, Rio Puerto
Viejo at Quebrada El Sura and Q. El Salto, Rossbach
3710 (GH), Scamman 7482 (GH), Scamman & Hold-
ridge 7998 (CR, GH); 2 km upstream from confluence
of Rio Puerto Viejo with Rio Sarapiqui, Finca “La
Selva,” Burger & Stolze 5865 (CR, F, GH, US), Mickel
3570 (NY). Limon: Llanuras de Santa Clara, Biolley
10688 (CR, US); Hundrisser Ranch, Atlantic slope,
Brade & Brade 374 (NY, UC); 7 km SW of Bribri,
100-250 m, Gomez et al. 20423 (CR, MO, UC); N end
of Tortuguero Natl. Park, Burger & Antonio 1/263 (CR,
F, NY); near Guapiles, at bridge over Rio Guacimo, 50
m, Moran 2182, 2185, 2186 (CR, F, P, PORT); near
banana plantation and Pandora, near Rio Estrella,
Rossbach 3624, 3625 (GH); La Lola, a cacao finca,
near Rio Madre de Dios, Scamman 7156 (GH); Finca
Montecristo, on the Rio Reventaz6n below Cairo, 25
m, Standley & Valerio 48622 (US). Puntarenas: Osa
Peninsula, Mickel 1944 (CR, NY); hills N of Palmar
Norte, along trail to Jalisco, Croat 35176 (MO), Osa,

40 km W of IA rt. 2, Gomez 19489 (CR, MO).
PANAMA. Bocas del Toro: in Laguna de Chiriqui

and its neighborhood, Hart 53 (US). Canal Zone: “Isth-
mo Panama,” Hayes 8 (B, GH, Y, US); hills N of
Frijoles, Standley 27597 (MO, US); Cerro Azul, Tyson

Vol. 34, Art. 1

2109 (MO); along hogback ridge S of Fuertes Cove,
Pearson Peninsula, Croat 8/53 (MO); headwaters of the
Rio Chinilla, above Nuevo Limén, Maxon 6895 (US);
ravine near Frijoles, Killip 2929 (US): along riverbank
between Fort Sherman and Fort Lorenzo, Croat & Porter
15436 (MO); hilly forest around the Agua Clara Reser-
voir, near Gatun, 20-30 m, Maxon 464] (GH, NY,
US); Orange River Trail, Cornman 544 (UC, US); Barro
Colorado Island, Gatun Lake, Maxon et al. 6817 (GH),
Seaverns 56 (F), Starry 9] (F, GH), Croat 5114 (MO),
6850 (MO), 7362 (MO), 8025 (MO), 9000 (MO, NY),
9004 (MO, UC), 9/03 (COL, MO, UC), 10804 (MO),
15256 (MO), 17367 (F, MO), Bailey & Bailey 541 (GH),
Chrysler 4836 (UC); 10 mi from main gate, near Rio
Frijolito, Croat 15081 (F, MO, US); Allison Armour
Trail, Wetmore & Woodworth 130 (GH). Chiriqui:
above Rio San Felix near town of San Felix, ca. 13 mi
N of Rio San Felix bridge, 800-1200 m, Croat 33452
(MO); forest behind Vivero forestal, 12 km N of Los
Planes de Hornito, IRHE Fortuna Hydroelectric Project,
1200-1300 m, Knapp 4966 (MO). Colén: Santa Rita
Ridge, end of road from Transisthmian Highway, ca.
10 mi from road, Porter et al. 4763 (MO); Santa Rita
Ridge, 2.8 mi from Boyd—Roosevelt Hwy., Croat &
Porter 15342 (F, MO); forest along Portobelo—Nombre
de Dios road, 10 km W of Nombre de Dios, Knapp &
Mallet 5402 (MO). Darién: vicinity of Cana, summit
of knoll above Cana, Stern et al. 523 (GH). Panama:
2.5 mi N of Goofy Lake on road to Cerro Azul, Croat
11544 (F, MO); upper Mamoni River, 150—400 m, Pit-
tier 4492 (US); Orange River Valley, E of Juan Diaz,
Killip 2544 (GH, US); along Rio Pirati, foothills of the
Serrania de Maji, Knapp & Mallet 5/33 (MO); in high
ridges of the Serrania de Maji, S of the Choco village
of Ipeti, Knapp et al. 4479 (MO); along Juan Diaz River,
4 mi above Juan Diaz, 0-75 m, Killip 2845 (US). San
José Island: Perlas archipelago, Gulf of Panama, 55 mi
SSE of Balboa, Johnston 452 (GH, US). Veraguas: Isla
de Coiba, Mendez 75 (MO).

TRINIDAD. Aripo road via Arima, Broadway 5717
(F, Z); without locality, Fendler 105 (GH, MICH, MO,
NY, P, PH, UC); without locality, Broadway 5358 (NY);
forest, Brazil, Britton et al. 2144 (GH, NY); Mount
Tamana, Britton et al. 1935 (NY); Oropuche, local road
via Valencia, Broadway 9207 (MO, UC); without local-
ity, Crueger 139 (B); La Sierra, Maraval, Hombersley
s.n. (MO); Arima Ward, Guanapo River Valley, ca. 1
mi SE of La Leja village, Jermy 3/22 (BM); Cumaca
Road, Fay 472 (BM); Brickfield’s Tea Plantation, 3 mi
S of Forestry Rest House, Jermy 2/37 (BM), Valencia
Ward, near Valencia, Jermy /0826 (BM); Hollis Reser-
voir Road, Mickel 9472 (UC).

FRENCH GUIANA. Saint Jean du Mearoni, Benoist
127] (P); Central Guyana, Leprieur s.n. (P); Sommet
Tabulaire, 650-750 m, ca. 45 km SE of Saul, Cremers
6374 (CAY), 6523 (CAY, Z); Bassin du Haut Inipi, 7
km WSW du Pic Coudreau (Monts Bakra), Granville
4012 (CAY, Z); Haut Oyaopock, W of Trois Sauts
Crique Euleupousing, rive gauche du Saut Cambrouse,

November 1987

Granville 1143, 1165 (CAY, NY, Z); S of Tampoc:
Saut Koumakou Soula, Cremers 4460 (CAY, Z); Saut
Tampoc, Granville 484] (CAY); Haute Riv. Mana: Saut
Grous Tigre, Cremers 7552 (CAY, Z); Haute Crique
Baboune, affluent de la Riv. Mana, Cremers 7369
(CAY, Z); Haute Crique Waamahpann, au depart du
chemin indien, Granville 97] (CAY); Frontiere Guyane—
Surinam, Tuma Humac, Granville 99] (CAY, Z); Crique
Gabaret, 25 km de l’embouchure, layon N—W, Oldeman
1933 (CAY, NY); Crique “Roche Fende” (affleunt de
la Comte) a | km environ de son embouchure, Granville
B.4711 (CAY, Z); W of Saul on trail to Monts Galbao,
Boom & Mori 1856 (CAY, NY); Piste allant de Citron
vers le Massif du Decou Decou, Billiet & Jadin 1683
(CAY); Region de Paul Massif du Decou Decou, Crem-
ers 7951 (CAY, Z).

SURINAM. West Rivier, 2-5 km SW of Juliana
Top, 275-300 m, /rwin et al. 54896 (NY); no locality,
Hostmann s.n. (NY); 45 km S of Paramaribo, new road
to Hanover, N of Zanderij, Tryon & Kramer 5611] (GH,
MICH, NY); about 25 km S of Paramaribo, Lindeman
4570 (GH), Kramer 1954 (MO); 3 km S of Juliana Top,
12 km N of Lucie River, 300—325 m, /rwin 5516/ (MO,
NY); Para Dist., Lindeman & Teunissen 1529] (Z);
Suriname River, Plantage “Accaribo,” d’Angremond
s.n. (Z).

GUYANA. Kamuni Ck., Groete Ck., Essequibo
River, Maguire & Fanshawe 22855 (GH, NY); Es-
sequibo River, Persaud 372 (F); Rockstone, Gleason
582 (GH, NY); Demerara, Jenman s.n. (NY); Barima
River, Jenman s.n. (NY); 3 mi S of Chodikas, Guppy
462 (BM); Essequibo River, Moraballi Ck., near Bar-
tica, Richards 803 (BM); lower Cuyuni River, Sandwith
1561 (BM).

VENEZUELA. Bolivar: Dtto. Heres, Campamento
Gun, Fernandez 951 (PORT, UC); Roraima, Schom-
burgk s.n. (B, NY); without locality, anno 1843, Schom-
burgk 1659 (B); E of Cerro El Picacho, N of Las Nieves
and Las Chicharras, 45 km N of Tumeremo, vicinity of
Beborah, Altiplanicie de Nuria, 600—650 m, Steyermark
89120 (NY, VEN); a lo largo de la Quebrada Acarabisi,
limites del Estado Bolivar con la zona dereclamaci6n,
Aymard et al. 952 (PORT, UC). Portuguesa: Depto.
Paez, Pozo Blanco, entre Acarigua y Payara, 190 m,
Ortega 636 (PORT, UC, VEN). Sucre: selva del Rectan-
gulo N del Lago de Tuanoco, Lasser & Vareschi 3926
(VEN); vicinity of Cristobal Colon, Avicagua, Broad-
way 560 (GH, NY, US). Territorio Federal Delta
Amacuro: Depto. Antonio Diaz, 9°15'N, 60°57'W,
upper reaches of riverine forest of Cano Atoiba tributary
of Boca Araguao, Steyermark et al. 115033 (VEN);
Depto. Antonio Diaz, vicinity of Cano Jotajana ( = tierra
alta), tributary of Cano Guiniquina, NW of Epana near
boundary with Depto. Tucupita, 9°15'N, 61°10'W,
50 m, Steyermark etal. 115021 (MO,UC, VEN); Depto.
Pedernales (boundary with Depto. Tucupita): Cano
Simoina, west of Isla Coucuina, S of Barra de Coucuina,
50 m, Steyermark et al. 114331 (MO, VEN), 1/4332
(MO, UC, VEN).

MONOGRAPH of POLYBOTRA 65

COLOMBIA. Amazonas: Trapecio Amazonico,
Loretoyacu River, 100 m, Schultes & Black 8467 (GH,
US). Cauca: Cali, Lehmann 2998 (BM); en la orilla
opuesta Puerto Limén—Rio Caqueta, Mora 4430 (COL);
Gorgona Island, off Narino, Taylor 1223 (MICH, NY);
Rio Timbiqui, B.7. 443 (GH). Chocé: slopes and ridge
of Loma del Cuchillo, ca. 15 km WSW of Chigorod6,
150-400 m, Lellinger & de la Sota 643 (LLP, US);
upper Rio Truando, 2 km SSW of the confluence of
Rio Nercua near the MADUREX Camp, 100 m, Lel-
linger & de la Sota 589 (COL, LP, US); trail to Miniquia
E of Puerto Mutis (Bahia Solano), 20-120 m, Lellinger
& de la Sota 26 (CR, COL, LP, US); Rio San Juan,
3.5 km SW of Andagoya, just NE of the mouth of the
Rio Suruco, 60 m, Lellinger & de la Sota 496 (COL,
LP); Municipio de Rio Sucio, Parque Nacional Los
Catios, alrededor del Campamento de Tilupo, 250-320
m, Forero & Jaramillo 1745 (COL, MO). Cun-
dinamarca: Cordillera Oriental, Mesa Negra, Gazuguan
Valley, 6 km NW of Medina, 580 m, Grant 10436 (F,
US). El Valle: La Cumbre, Cordillera Occidental,
1700-2200 m, Killip 11343 (GH, NY, US); Finca La
Pradera ca. 6 km SW of El Cairo on trail to Rio Blanco,
between El Brillante and Boquerén, Lellinger & de la
Sota 790 (COL, US). Narino: Pambana, between Rio
Pimbi and Rio Cuembi, on Rio Telembi, above Bar-
bacoas, 50 m, Ewan 16846 (BM, GH, UC).

Ecuapor. Los Rios: Rio Palenque biological sta-
tion, km 56 Quevedo—Sto. Domingo road, Evoy 104
(NY). Morona-Santiago: Pachicutza, at “Escuela is-
comisional [sic.] Cardinal Dofner,” km 140 on road
Loja Gualaquiza, 900-1000 m, Holm-Nielsen et al. 4489
(AAU, Q, QCA), 4495 (AAU, Q, QCA); Sucua,
Swingle et al. 70-02-05-2 (UC, US). Napo: casi 10 km
SE de Tena, 3 km este de Puerto Nuevo por camino a
Puerto Misahualli, 300 m, Moran 3534 (F, Q, QCA);
10 km al sur de Tena a Puerto Misahualli, Moran 3588
(Q, QCA); Anangu, Parque Nacional Yasuni, SEF proj-
ect, 260—350 m, @llgaard et al. 38932 (AAU, Q, QCA);
27 km SE of Coca, Moran 3617 (MO, C, QCA). Pas-
taza: Rio Bufeo, northern tributary of Rio Bobonaza,
300 m, Qlilgaard et al. 43798 (AAU, Q, QCA); Rio
Bobonaza, around houses between Huagracachi and
Cachitama, below Montalvo, 300 m, @llgaard et al.
34634 (AAU, Q, QCA); oil exploration camp Chi-
chirota, on the Rio Bobonaza, 300 m, @llgaard et al.
35290 (AAU, Q, QCA). Sur de Santander: vicinity of
Barran ca Bermeja, Haught 1325 (MO).

Peru. Amazonas: Prov. de Bagua, left bank of
Rio Maranon opposite Quebrada Mirana (opposite km
277 of Maranon road above Cascadas de Mayasi), 425—
450 m, Wurdack 2011 (GH, US, USM). Cuzco: Prov.
La Convencién, 73°40'W, 12°30'S, at Camp Zero,
710 m, Dudley 11513 (GH, US); Camp 1, 910-940 m,
in J. Knox’s quadrate, Dudley 10/68 (GH); Prov.
Paucartambo, Mautainiza [?], 800-900 m, Vargas
17800 (GH). Huanuco: Tingo Maria, 615-1100 m,
Allard 22593 (US); Cuchero, Poeppig s.n. (BM); hills

66 ILLinois NATURAL HISTORY SURVEY

E of Tingo Maria, Croat 2119] (MO); 10 km S of Tingo
Maria, Stork & Horton 9509 (F, UC, US); Pampayaco,
Poeppig 201 (B). Loreto: Prov. Maynas, Peter Jensen’s
Explorama Lodge, 50 mi downriver from Iquitos at
Yanamono Ck., Moran 3663 (AMAZ, USM); Prov.
Maynas, about 10 km SW of Iquitos at zoological park,
Moran 3671 (AMAZ, USM); Pebas on the Amazon
River, Williams 1739 (F); Caballo—Cocha on the Ama-
zon River, Williams 2137 (F); Rio Mazan, 100-125 m,
C. Schunke 380 (GH, NY); Prov. Maynas, Rio Itaya,
10 km S of Iquitos, Tryon & Tryon 5204 (GH); lower
Rio Huallaga, 155-210 m, Williams 3999 (F, US);
Quebrada Nawampa, Croat 17620 (MO), Puerto Arturo,
lower Rio Huallaga below Yurimaguas, 135 m, Killip
& Smith 30690 (NY, US); Mishuyacu, near Iquitos, 100
m, Klug 1166 (F, NY, US); Quebrada Tahuayo above
Tamishiyaco, Croat 19717 (MO); Rio Itaya above
Iquitos, Croat 19220 (MO); primary forest 17 km SW
of Iquitos on road to Puerto Almendara, Croat 18388
(MO); 12 km SW of Iquitos, Croat 18219 (MO). Madre
de Dios: 12°49'S, 69°17'W, Prov. Tambopata, Tam-
bopata Nature Reserve, ca. 30 air or 70—80 river km
SSW of Puerto Maldonado at effluence Rio Torre/Rio
Tambopata, 260 m, Barbour 4764, 4790 (F, MO); Par-
que Nacional de Manu, Cocha Casha Biological Station,
Foster P-84-42 (F). Pasco: Puerto Bermudez, 375 m,
Killip & Smith 26637 (NY, US). Puno: San Gavan,
Lechler 2321, 2329 (B). San Martin: San Roque, 1350—
1500 m, Williams 7681 (F, US); Cerro de Campana,
Spruce 4634 (P).

Bo.iviA. La Paz: Prov. Sud Yungas, Limite de
los Deptos. La Paz/Beni, Rio Quiquibey, 320 m, Beck
8037 (LPB); San Carlos, Mapiri, Mapiri River, Tate
422 (LPB, NY); Mapiri, Buchtien 35 (B), 290 (NY,
UC). Santa Cruz: Prov. Sara, Rio Yapaicani, 400 m,
Steinbach 7499 (B, F, GH, MO, Z); bosque del Rio
Surutt, 400 m, Steinbach 3032 (US).

BRaAzIL. Acre: Cruziero do Sul vicinity, Rio Moa
between Igarape Ipiranga and Aquidaba, Prance et al.
12069 (F, LP, NY, UC). Amazonas: prope Sao Gabriel
da Cachoeira, ad Rio Negro, Spruce 2116 (B, BM, P,
RB); Larges, on the Amazon River | km below mouth
of Rio Negro, Conant 940 (F, GH, NY). Para: Serra
dos Carajés, AMAZ camp AZUL, Sperling 5914 (GH,
NY); Belém, Huber 2599 (BM), 7235 (BM).

10. Polybotrya goyazensis Brade (Fig. 29, Map 6).

Polybotrya goyazensis Brade, Bradea 1:24,
tab. 1, fig. 1. 1969. Type: Brazil. Goias:
Goiania, primary forest, December 1936,
A. C. Brade 15373 (holotype: RB!; isotype:
NY!).

Polybotrya macedoi Brade, Bradea 1:24.
1969. Type: Brazil. Goids: Municipio Jatat,
Localidad Balsans, 1 November 1950,
Macedo 2682 (holotype: HB; isotypes: HB,
MO!, SP, US!; paratypes: Brazil. Goias:

Vol. 34, Art. 1

Fazenda Queixado, 8 December 1948,
Macedo 1447 (RB, SP); Fazenda Balsamo,
15 December 1948, Macedo 152] (SP).

Stem 1-2.5 cm thick, hemiepiphytic; scales
spreading, membranaceous, generally 6—12(16)
mm long, dark castaneous or dark reddish with
lighter borders, margins strongly denticulate to
erose, the base cordate to various degrees, often
black and sclerified at the point of attachment.
Sterile leaves up to 1.45 m long; petiole / to V2
as long as the lamina; lamina mostly 2-pinnate-
pinnatifid, lanceolate to ovate, 1.2 x 0.9 m, the
abaxial surface nearly glabrous to densely pubes-
cent, the hairs whitish, acicular, up to | mm long,
glands occasionally present, these reddish, sessile,
resinous; pinnae up to 45 x 23 cm, lanceolate to
narrowly deltate, the suprabasal ones soon becom-
ing pinnatifid; pinnules narrowly deltate, mostly
falcate, catadromically arranged above the base,
4—8(10) X 1-2(3) cm, the acroscopic side slightly
prolonged, the margins entire to crenate, ciliate,
the hairs minute, 4—10 celled; axes nearly glabrous
abaxially or densely pubescent, by tiny, unicellu-
lar hairs, scales very small or lacking, adaxially
pubescent in the central groove with reddish hairs,
0.3—0.8 mm long; grooves usually ciliate on the
ridges. Fertile leaves 2-pinnate, amphiacrosti-
choid; pinnules caudate, often lobed or undulate
at the base; sporangial stalks paraphysate; spores
(44)48—62(70) microns long.

Other illustrations: See original description
of P. goyazensis; Sehnem, FI. Ilust. Catarinense,
pl. 18, 1979.

Polybotrya goyazensis occurs in Paraguay —
the only species of Polybotrya known from that
country—and in the southern half of Brazil but
not in the coastal mountains (Map 6). This range
is unusual because it comprises regions apparently
not occupied by other species of Polybotrya. |
suspect that P. goyazensis evolved from south-
ernmost, isolated populations of P. caudata.

Polybotrya goyazensis has stem scales that
are spreading, shiny, membranaceous, translu-
cent, darkened in the center with lighter borders,
margins strongly denticulate to erose, and the base
cordate around the darkened point of attachment
(Fig. 29f). In contrast, P. caudata has stem scales
that are curved, appressed, thick, dull brown, con-
colorous, margins entire, and attached across the
length of the thickened base (Fig. 28h). The mi-
nute cilia on the margins of the lamina serve to
distinguish these two species from almost all other
species of Polybotrya (Fig. 28a).

November 1987

a SS

FiGureE 29. Polybotrya goyaze
pinnules; f. stem scales. a,b: R
e,f: Argent et al. 6336 (UC).

15 cm

nsis Brade. a. fert
ojas 1043] (B). c: B

fertile leaf; b. sterile leaf; c—e. mal acroscopic
rade 15373 (RB). d: Prance & Selva 59677 (NY).

1cm

68 ILLINOIS NATURAL HISTORY SURVEY

Brade (1969c) distinguished P. macedoi from
P. goyazensis by the former’s pilose hairs on the
axes and its slightly darker scales with lighter bor-
ders and a longer acuminate tip. These differences
are not constant. Various degrees of pubescence
exist, from densely pilose to nearly glabrous. This
intergradation is shown by the type of P. goyazen-
sis: the holotype at RB is nearly glabrous but the
isotype at NY is densely hirsute. Examination of
the stem scales shows a complete gradation of
color, sometimes even on a single specimen.

Specimens examined: BRAZIL. Goias: Goiania,
primary forest, December 1936, A. C. Brade 15373
(RB, NY); Serra do Caiapo, 40 km S of Caiaponia,
Prance & Silva 59677 (NY); Municipio Jatai, Localidad
Balsans, Macedo 2682 (MO, US). Federal District:
“Brasilia,” Glaziou 14456 (B, P). Mato Grosso: 270
km N of Xavantina, 12°54'S, 51°52'W, Ratter et al.
2047 (MO); Municipio Brilhante, Rio Anhandui,
Hatschbach 25118 (UC); Expedition Base Camp, Brejo,
12°49'S, 51°46'W, Argent & Richards 6651 (NY);
49.5 km N along road from base camp, Harley et al.
10922 (NY); Xavantina, ca. 4 km N of ferry, Argent et
al. 6336 (RB, UC). Para: no locality, 1914, Perelot
g.n. (F).

PARAGUAY. Southern Paraguay, Sierra de Amam-
bay, Rojas 10182 (B, BM), /043/ (B, L, LIL, M,
MICH, P), /0804 (B, BM, L); no locality, Hassler
11618 (MICH).

11. Polybotrya pubens Martius (Fig. 30, Map 6).

Polybotrya pubens Martius, Icon. Plant.
Crypt. Brasil. 87, tab. 25. 1834. Type:
Brazil. ad flum. Amazonas prope Serpa, Sep-
tember, Martius s.n. (holotype: M).

Polybotrya pubens Kunze, Linnaea 9:22.
1834. non Martius. TyPE: Peru. Loreto: Prov.
Maynas, ad Yurimaguas, September 1830,
Poeppig s.n. (lectotype: P!).

Acrostichum pubens (Kunze) Hooker &
Baker, Synopsis Filicum 414. 1868.

Acrostichum caudatum (Martius) Hooker var.
pubens (Martius) Baker, Fl. Brasil. 1(2):586,
tab. 38, figs. 1,2. 1870.

Dryopteris guentheri Rosenstock, Feddes
Repert. 25:59. 1928. Type: Bolivia. La Paz:
region of Mapiri, San Carlos, 15 November
1926, Buchtien 260 (holotype: UC!).

Polybotrya crespiana Bosco, Nouvo Giorn.
Bot. Ital., II. 45:145, tab. 9, fig. 1. 1938.
Type: Ecuador. Santiago—Zamora: Indanza,
Crespi s.n. (type not found, known only from
Bosco’s fig. 1).

Vol. 34, Art. 1

Polybotrya decorata Lellinger, Amer. Fern
J. 62:54, figs. 5, 6, 11. 1972. Type: Peru.
Loreto: Iquitos, alt. ca. 100 m, Killip & Smith
26955 (holotype: US!; isotype: NY!).

Stem 1.0—1.5 cm thick, hemiepiphytic; scales
of large mature stems mostly 10-15 x 2-5 mm,
flaccid, membranaceous, concolorous, whitish to
coffee brown or castaneous, spreading or more
commonly matted, margins fimbriate, young or
terrestrial stems with narrowly lanceolate, castane-
ous, denticulate scales. Sterile leaves up to 1.2m
long; petiole up to 21 cm long, about 4 to / the
length of the lamina; /amina to | m long, narrowly
deltate to elliptic, tapered evenly to the apex,
mostly 1-pinnate-pinnatifid, the base occasionally
2-pinnate-pinnatifid, pubescent throughout or
rarely glabrous adaxially, rarely with punctate,
reddish glands; pinnae narrowly triangular, up to
9-19 x 2.5—3.5 cm, serrate to pinnatifid, often
with only the lowest pinnae having a few free
proximal segments; pinnules or segments cata-
dromic, slightly falcate, oblong, the basal ones
slightly longer than the suprabasal and more
deeply cut; veins free, 6—9 per segment; axes con-
spicuously pubescent, the hairs 1-2 mm long, un-
branched, acicular, whitish, 6—15 celled, the sep-
tae often reddish, polyseriate scales lacking, but
scales present as reduced, appressed, reddish, flat-
tened, uniseriate “hairs.” Fertile leaves 2-pinnate,
to 0.8 m long, amphiacrostichoid; axes pubescent
on both surfaces, the hairs ca. 0.1 mm long, col-
orless, subulate; proximal pinnae deeply lobed or
fully pinnate, long-caudate; sporangia with the
capsule walls setose at the apex near the annulus
by two hairs, these 1—3 celled; sporangial stalks
paraphysate; spores (44)48—57(63) microns long.

Other illustrations: Martius, Icones Plant.
Crypt. Brasil. 87, tab. 25. 1834; von Ettings-
hausen, Denkschr. Ak. Wien., tab. 7, figs. 3, 10,
11. 1864; Martius, Fl. Brazil, 1(2):586, tab. 38,
figs. 1, 2. 1870; Brade, Bradea 1(9): 63, fig. 4;
Ibid. 67, fig. 7. 1971.

Polybotrya pubens grows primarily in low-
land forests of the Amazon basin but occurs in
premontane forests up to 1400 m in the Andes
(Map 6). During my fieldwork in Ecuador, I found
this species in well-drained forest soils that were
slightly elevated above the surrounding swamp
forest.

This is one of the most distinct species of
Polybotrya, readily distinguished by its combina-
tion of stem scales, lamina shape and cutting,

November 1987 MONOGRAPH of POLYBOTRA 69

FIGURE 30. Polybotrya pubens Martius. a. pinna of a large, highly divided leaf; b. pinna of typically
cut leaf; c. sterile leaf; d. fertile leaf; e. setose sporangium with paraphysis on stalk; f. setose sporangium;
g. cross section of fertile pinna apex; h. stem scales; i. hairs from sterile lamina. a: Killip & Smith
26955 (US). b: Killip & Smith 26543 (NY). c,h,i: Balslev 4787 (CR). d,g: Killip & Smith 26194 (NY).
e,f: Holm-Nielsen & Jeppsen 663 (AAU).

70 ILLINOIS NATURAL HISTORY SURVEY

pubescence, and long-caudate fertile pinnae. In
addition, P. pubens has, unlike any other species
of Polybotrya, setose sporangial capsule walls
(Fig. 30e,f). The sterile lamina is usually 1-
pinnate-pinnatifid or almost 2-pinnate at the base,
but large leaves that are 2-pinnate-pinnatifid for
much of their length are occasionally collected
(Fig. 30a). These large, more highly divided
leaves appear quite distinct but there are inter-
mediate forms between the extremes of lamina
cutting. The 2-pinnate-pinnatifid form was named
as P. decorata.

Polybotrya pubens and P. glandulosa might
be confused because both have similar laminar
cutting, long, acicular hairs, and both grow in
lowland forests of the Amazon basin. But the
lamina of P. glandulosa is, unlike any other
species of Polybotrya, cuneate and the petiole is
extremely short, only 2.5 cm long. Furthermore,
its fertile leaf is botryoid instead of amphiacros-
tichoid, and its sporangial capsules are glabrous
instead of setose.

Juvenile stems and thin terrestrial stems have
narrow, castaneous, spreading scales that differ
from those on the scandent stems. Dryopteris
guentheri represents a juvenile form of P. pubens
with this scale type. The transition from juvenile
to mature plants and the changes in scale and
lamina morphology are shown in an excellent
series of specimens collected in Ecuador by
Balslev (#4787).

The pubescence of the fertile leaf consists of
colorless, subulate hairs about 0.1 mm long—
these are much reduced in comparison to the hairs
on the sterile leaves. The setae on the sporangial
capsule walls are unique in Polybotrya. They
occur only at the apex of the sporangium near the
annulus (Fig. 30e,f); it is odd that these setae are
not shown on the sporangia illustrated in 1834 by
Martius in his otherwise excellent plate of P.
pubens. The adaptive value (if any) of the hairs
on the capsules is not known.

Unlike all other species of Polybotrya, P.
pubens lacks wide, polyseriate scales on the axes.
Rather, scales are present only in their reduced
form, that is, as tiny, appressed, reddish, jointed,
uniseriate “hairs.”

Both Martius and Kunze published the name
P. pubens in 1834. Kunze’s work appeared in the
first issue of Linnaea, which was published 22-28
June 1834 (Stafleu 1967). Stafleu and Cowan
(1981) report that Martius’s second fascicle of
“Icones plantarum crytogamicarum” was pub-
lished in 1834. Since no month or day is given,

Vol. 34, Art. 1

I cannot determine which work was published
first. I consider Martius the author of P. pubens
because Kunze clearly attributed this name to Mar-
tius.

Specimens examined: COLOMBIA. Amazonas:
Rio Apaporis, Soratama (above mouth Rio Kananari)
and vicinity, Schultes & Cabrera 16053 (US).

Ecuapor. Morona-Santiago: Taisha, 3-4 km
ESE of the military camp, Brandbyge & Asanza C.
31870 (AAU). Napo: Reserva Faunistica Cuyabeno, al
lado norte de Laguna Grande, Balslev 4787 (AAU, CR,
Q, QCA), 4802 (AAU, Q, QCA), Laegaard 51136
(AAU, Q, QCA), 51/50 (AAU, Q, QCA), 5/224 (AAU,
Q, QCA); Parque Nacional Yasuni, in the area of the
SEF project, @llgaard et al. 38969 (AAU), 39039
(AAU), 39040 (AAU); 6 km along Rio Pano, Holm-
Nielsen & Jeppsen 663 (AAU). Pastaza: Rio Bobonaza,
oil exploration camp Chichirota and Destacamento Cabo
Pozo, Mllgaard et al. 34894 (AAU, Q, QCA); Curaray,
SE of the airstrip, Holm-Nielsen et al. 22/21 (AAU);
oil exploration camp Chichirota, on the Rio Bobonaza,
Allgaard et al. 35350 (AAU); Rio Bobonaza, between
Cachitama and the outlet of Rio Bufeo, @llgaard et al.
34703 (AAU). Santiago-Zamora: Taisha, Cazaler &
Pennington 7720 (BM, F, K, NY, US).

PERU. Amazonas: ridge crest of Quebrada Chuivi
(above km 278 of Maranon road), valley of Rio Maranon
near Cascadas de Mayasi, Wurdack 1933 (US). Junin:
E of Quimiri Bridge, near La Merced, Killip & Smith
23916 (NY, US); Pichis Trail, Santa Rosa, Killip &
Smith 26194 (NY, US); Pichis Trail, San Nicolas, Killip
& Smith 25972 (NY, US); Puerto Bermudez, Killip &
Smith 26543 (NY, US). Loreto: above Tamishuyacu,
Croat 1977] (AMAZ, MO); primary forest 17 km SW
of Iquitos, Croat 1/8508 (MO); Iquitos, ca. 100 m, Killip
& Smith 26955 (NY, US); December 1830, Poeppig
s.n. (photo of specimen at V, BM); Prov. Maynas, ad
Yurimaguas, September 1830, Poeppig s.n. (P); La Vic-
toria on the Amazon River, Williams 3/36 (F); lower
Rio Huallaga, Williams 4797 (F, US); Yurimaguas,
Spruce 3880 (P, fragment NY); along Quebrada
Nawampa, Croat 17689 (MO); Santa Rosa, lower Rio
Huallaga below Yurimaguas, Killip & Smith 28760 (NY,
US). Pasco: Prov. Oxypampa, Iscoazin, Foster 7858
(F, USM). San Martin: Monte Campana, Spruce 4740
(photo GH, US); camino a Pushurumbo, 7—8 km al este
del Puente de Palo Blanco, Mariscal Caceres, Tocache
Nuevo, J. Schunke 5789 (NY, US); San Roque, Wil-
liams 7620 (F).

Bo.ivia. La Paz: Prov. Larecaja, Consata 7 km
towards Mapiri, Beck 4924 (F, LPB); region of Mapin,
San Antonio, Buchtien 35 (US), 1/23 (US), 1/24 (NY);
region of Mapiri, San Carlos, Buchtien 260 (UC), 299
(NY); Mapiri, Rusby 442 (NY).

BRaAziL. Amazonas: Rio Negro, 1819, Martius s.n.
(photo of M specimen at BM). Para: Martius s.n. (photo
of M specimen at BM).

November 1987

12. Polybotrya glandulosa Kuhn (Fig. 31, Map 7).

Polybotrya glandulosa Kuhn, Linnaea 36:65.
1869. Type: Brazil. Amazonas: San Gabriel,
no date, Spruce s.n., not 2//6 as indicated
on Morton negative, (holotype: B!).

Polybotrya subelliptica Lellinger, Amer. Fern
J. 62:56. 1972. Type: Peru. Loreto: Mis-
huyacu, near Iquitos, 100 m, Klug 1390
(holotype: US!; isotypes: F!, NY).

Stem 1—2 cm thick, hemiepiphytic; scales dull
brown, thick, opaque, concolorous to bicolorous,
curved-appressed, 8—15 1.5—2.5 mm, the mar-
gins entire, the base thickened and expanded,
slightly elevated, the dorsal surface sometimes
with a medial groove. Sterile leaves up to 75 cm
long; petiole very short, up to 2.5 cm long; lamina
2-pinnate-pinnatifid, subelliptic, 65-73 x 22-28
cm, the base cuneate, about 8 cm wide, the apex
long-attenuate, the abaxial surface with or without
sessile, red, resinous, round glands, the margins
ciliate with hairs similar to those of the veins and
axes; pinnae Ca. 35 pairs, the medial ones 14-16
x 3—3.5(4) cm, sessile or short-stalked, oblong,
the base truncate; pinnules catadromic, oblong,
falcate at the apex, those of the medial pinnae
15-20 x 5—7 mm, the margin entire, crenate or
lobed, generally with 6—10 veins, the basal basi-
scopic margin decurrent; axes and veins densely
pilose on both surfaces, the hairs tawny, pluricel-
lular, acicular, 1.0—2.5 mm long, the scales few
or absent. Fertile leaves narrowly elliptic, 45 x 15
cm, 2-pinnate-pinnatifid, botryoid; medial pinnae
6—8 X 1.5—2 cm; axes sparsely pilose, with scat-
tered, linear, appressed, dark scales; sporangial
stalks paraphysate.

Polybotrya glandulosa, which has been col-
lected only three times, is one of the rarest species
of Polybotrya. It is the only species in the genus
that is endemic to the Amazon basin (Map 7);
however, I suspect it will be found in the adjacent
Guiana Highlands as that region becomes better
explored. I failed to find this species during my
fieldwork in the Iquitos area, in part because the
location of “Mishuyacu” is unknown. The eleva-
tional range for this species is 100—140 m.

This species is unlikely to be confused with
any other Polybotrya since, unlike other species
in the genus, the lamina tapers gradually to an
extremely short petiole (Fig. 3lc). The long,
acicular hairs on the axes and veins further distin-
guish this species from most other congenerics.

MONOGRAPH of POLYBOTRA 71

Polybotrya pubens, however, has long acicular
hairs and leaf cutting similar to P. glandulosa;
since it grows in Amazonian forests, the two
species could be confused. Polybotrya glan-
dulosa, however, can be distinguished from P.
pubens because the former has a botryoid, instead
of an amphiacrostichoid, fertile leaf and a short
petiole.

The specific epithet refers to the red, sessile,
round, resinous glands on the abaxial suface of
the type specimen (Fig. 31d). The two other col-
lections, however, lack such glands. This variabil-
ity is not unusual, since glands are variably present
in all gland-bearing species of Polybotrya.

Specimens examined: VENEZUELA. Territorial
Federal Amazonas: Dept. Rio Negro, 0-3 km N of
Cerro de Neblina Base Camp, on the Rio Mawarinuma,
140 m, Liesner 16293 (MO, UC).

PERU. Loreto: Mishuyacu, near Iquitos, 100 m,
Klug 1390 (F, NY, US).

BRAZIL. Amazonas: Sao Gabriel, no date, Spruce
s.n. (B).

13. Polybotrya lechleriana Mettenius (Fig. 32,
Map 7).

Polybotrya lechleriana Mettenius, Filices
Lechler. 1:4, tab. 1, figs. 1-5. 1854. Type:
Peru. Puno: San Gavan, Lechler 2156 (lec-
totype: B!; isotype: L!; fragments F!, US!;
photo GH! of K).

Acrostichum lechlerianum (Mettenius)
Hooker, Species Filicum 5:246. 1864, nom.
illeg., non Mettenius 1856.

Stem to 1.5 cm _ thick, hemiepiphytic,
mucilaginous?; scales lanceolate, up to 15 x 3
mm, cream to dull brown, thin, appressed, the
margins denticulate to entire. Sterile leaf pubes-
cent throughout, the hairs 3—12 celled, 0.3—1.2
mm long, lax, spreading, colorless; petiole /3—'/2
as long as the lamina, stramineous; lamina fine-
ly divided, (4)3-pinnate-pinnatisect, lanceolate,
both surfaces pubescent, especially along the axes
and veins, eglandular, the apex acute, not long-
attenuate; pinnae lanceolate to ovate, (7)10—15 x
(2)3—7(12) cm, the base truncate, short-stalked,
crowding the rhachis; pinnules catadromic, oblong
with subparallel sides, the base truncate, nearly
sessile, crowding the costa, the apex acute to
rounded; ultimate segments single veined, falcate,
0.5—1 mm wide; axes pubescent abaxially, usually
with a single scale at the juncture of the costa and

2 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

} 4
1cm

i

FicurE 31. Polybotrya glandulosa Kuhn. a. stem scales; b. fertile leaf; c. sterile leaf; d. abaxial
surface of sterile pinna showing sessile reddish glands (as dots) and acicular hairs; e. medial pinna.
a—c,e: Klug 1390 (US, F). d: Spruce s.n. (B).

MONOGRAPH of POLYBOTRA 73

November 1987

11cm

way
Fp ~

Glee t oY

Fi peeRe

wdc,

SS

HL is Ree

SS

11cm

‘ya lechleriana Mettenius. a. sterile leaf; b. rhachis-costa juncture of fertile leaf;

c. costal hairs; d. rhachis-costa juncture of sterile leaf; e. stem scales. a,c: Spruce 4744 (P). b,d: Stiibel

914 (B). e: Dudley 10325 (GH).

FiGure 32. Polybotr

74 ILLiNnois NATURAL HISTORY SURVEY

costule, the scale ovate, appressed, thin, brown;
grooves glabrous or puberulent within, truncated
by the ridges of the next lower order, usually
becoming shallower near the juncture. Fertile leaf
3-4-pinnate, botryoid, pubescent with hairs like
those on the sterile leaf; sori distinct, round, 0.5—
0.8 mm wide, on short stalks, these 1-2 mm long,
the receptacle moundlike; sporangial stalks
paraphysate; spores (50)54—60(65) microns long.

Other illustrations: Hooker, Second Century
of Ferns, tab. 97. 1861; Mettenius’s original de-
scription has an excellent illustration of the type
at Berlin.

Polybotrya lechleriana occurs in rich mon-
tane forests from (100)1000—1500 m. This species
is primarily Andean but has a notable range dis-
junction in the Guiana Highlands at Mt. Roraima
in Guyana, 1900 km from the nearest known popu-
lation in Colombia (Map 7). This disjunction is
best explained as an example of long distance
spore dispersal. Another significant Andean—
Guianan range disjunction occurs in P. frac-
tiserialis (Map 3).

This species is easily recognized by its finely
divided sterile lamina with very narrow, single-
veined segments and lobes (Fig. 32). The finely
cut leaves, so unlike others in the genus, are prob-
ably what prompted Copeland (1947) to remark
that this species “looks out of place here [in
Polybotrya}.” But P. lechleriana has all the fea-
tures of a typical Polybotrya including the unique
stem anatomy of the genus.

Some forms of this plant resemble Polybotrya
stolzei an endemic to the Andes of Colombia, but
P. lechleriana differs by the narrower width of
its ultimate segments or lobes, each of which is
one-veined (see key). Polybotrya lechleriana
probably evolved from a less dissected ancestor
in the P. alfredii group by cessation of the growth
of the marginal meristem to produce a more finely
divided sterile lamina. This species is named in
honor of Wilibald Lechler (1814—1856), aGerman
pharmacist, botanist, and explorer, who sent many
of his collections to Mettenius.

Specimens examined: GUYANA. Mt. Roraima,
Waruma Trail, Persaud 1/4 (NY).

COLOMBIA. Cauca: forests of Rio Timbiqui,
Lehmann B.T413 (GH). Choco: alrededores de San José
del Palmar, Cerro S de la poblacion, Forero & Jaramillo
2469 (COL, MO, NY); NW side of Alto del Buey,
Lellinger & de la Sota 28] (COL, US); Rio Nuqui,
Haught 5498 (COL, F, NY, US). Narino: Municipio
de Altaquer, entre Junin y Buenavista, 650 m, Mora
4287 (COL).

Vol. 34, Art. 1

Ecuapor. Chimborazo: Rio Palora, eastern cor-
dillera of Riobamba, 1400 m, Rimbach 3] (GH, PH,
US). Pastaza: Mera, remnants of primary forest in hills
above town, Plowman & Davis 4542 (GH). Tun-
gurahua: along road between Puyo and Banos at second
bridge W of Mera, ca. 3 km W of Mera, 1160 m, Croat
49718 (MO, UC); Banes—Pintuc, Stitbel 9/3a, 914 (B,
BM).

PERU. Cuzco: Prov. La Convencién, Cordillera
Vilcabamba, Camp 2.5, 1760 m, 73°38’ W, 12°38'S,
Dudley 10325 (GH); Paucartambo, Valle de Pillahuata,
Gerrera 1624 (US). Junin: Prov. Satipo, “Mapiri” ca.
12 km SW of Chequitavo, A.C. Smith 61713 (MO,
USM); Pichis Trail, Porvenir, 1500-1900 m, Killip &
Smith 25921 (NY, US). Puno: San Gavan, Lechler 2156
(B, F, GH, L, US), 2/76 (B); La Pampa, Rio Tavara,
Watkins s.n. (US). San Martin: in monte Guay-
rapurima, prope Tarapoto, Spruce 4744 (L, P; fragment
US). Dept. unknown: entre Los Rios Tintiuiguiato del
Pachin y Tapecia del Tocate, 1500 m, Bues 1743 (US).

BoLiviA. La Paz: Hacienda Simaco sobre el
camino a Tipuani—Tale, 1400 m, Buchtien 5308 (BM,
GH, MO, NY, UC, US, Z).

14. Polybotrya attenuata Moran, sp. nov. (Fig.
33, Map 7).

Polybotrya attenuata Moran, sp. nov. TYPE:
Colombia. Antioquia: Guatape, forests on
slopes above river, 1/77, McAlpin & Kuhn
77-12 (holotype: NY!). The plant from which
the type specimen was taken is cultivated in
the fern greenhouse, New York Botanical
Garden (accession no. 332/78).

Planta hemiepiphytica; caulis ca. 1 cm diam.;
squamulis membranaceis, linearibus, acutis, 9—
12 mm longis, 0.3-—0.7 mm latis, in margine den-
ticulatis vel integris; laminae steriles lanceolatae
vel anguste deltatae, 3-pinnatae, apice longiat-
tenuatae, usque ad 55 cm longae, 26 cm latae,
glabrae vel punctatae, glandulis resinaceis, rotun-
dis; pinnulae catadromicae, 2.0—2.5 cm longae,
0.8—1 .2 cm latae, apice rotundatae; segmenta ter-
tiaria obovata, ad basim cuneata; costae et
rhachides sulcatae pubescentes in sulco, squamis
angustis, fuscatis, denticulatis; folia fertilia bot-
ryoidea; sori rotundi, ca. 1 mm longi, pedicellati.

Stem about | cm wide, hemiepiphytic; scales
light to dark brown, mostly concolorous or rarely
with a narrow hyaline border, thin, linear, 9-12 x
0.3—0.7 mm, the margins subentire to denticulate.
Sterile leaf up to 0.8 m long; petiole ¥3—V4 as
long as the lamina; lamina to 55 X26 cm,
3-pinnate, narrowly lanceolate or triangular, the
apex long-attenuate, the tissue with or without

—

SE, ee

November 1987 MONOGRAPH of POLYBOTRA 75

I,

“ASAD wT og

FiGuRE 33. Polybotrya attenuata Moran. a. sterile leaf; b. pinnules and costa-rhachis juncture (note
the abundance of narrow, dark scales on the axes); c. stem scales; d. two pinnules of fertile leaf;
e. fertile leaf. a—e: McAlpin & Kuhn 77-12 (NY).

76 ILLINOIS NATURAL HISTORY SURVEY

punctate, resinous, spherical, sessile glands; pin-
nae 10-13 X 3.5—4.0 cm long, lanceolate to ob-
long, the base truncate, short-stalked, crowding
the rhachis; pinnules catadromic, 2.0—2.5 * 0.8—
1.2 cm, with only S5—7 lobes or segments, the
apex acute or rounded, the base truncate, sessile
or with a short stalk less than 1 mm long; tertiary
segments obovate, entire to slightly lobed, con-
taining several vein branches, the base cuneate;
axes pubescent within grooves and especially ab-
axially, the hairs 4—12 celled, 0.2-0.3 mm long
jointed, with reddish cross-walls; costal scales
numerous, narrow, dark, denticulate, tortuous, ap-
pressed; grooves pubescent within, the hairs red-
dish, multicellular. Fertile leaves botryoid,
3-pinnate, densely scaly, the scales grading into
jointed hairs (the hairs are actually reduced scales);
sori stipitate, the stalks 1-3 mm long; sporangial
stalks paraphysate.

Polybotrya attenuata, named for its distinc-
tive, long-attenuate leaf apices, is endemic to the
Andes of Colombia (Map 7). This species differs
from P. lechleriana, a similar species, by its long-
attenuate apex, oval tertiary segments, resinous
glands on the lamina, and the darker, narrower
scales on the axes. The pinnules, furthermore, are
shorter and stubbier, having only 5—7 lobes and/or
segments, and these have a cuneate base (Fig.
33a,b).

The presence of glands on the lamina is not
constant. The living plant from which the holotype
was made has always produced leaves having
round, sessile, resinous glands on the undersur-
face. The Lehmann specimen, however, lacks
such glands. Similar glands are also variably pres-
ent in other species, such as P. osmundacea and
P. glandulosa.

Specimens examined: CoLomBiIA. Antioquia:
Guatape, McAlpin & Kuhn 77-12 (NY). Narino: along
river Pipulquer, west slopes of the western Andes,
Lehmann S500B (US).

15. Polybotrya stolzei Moran, sp. nov. (Fig. 34,
Map 9).

Polybotrya stolzei Moran, sp. nov. TYPE:
Colombia. El Valle: Santa Rosa, Dagua Val-
ley, forest along Rio Cabellete, 200—300 m,
22 September 1922, Killip 11549 (holotype:
US!; isotypes: GH!, NY!, PH!).

Vol. 34, Art. 1

Planta hemiepiphytica; caulis 0.6-1.5 cm
diam., (mucilaginus?); | squamae _ fuscatae,
opacae, lineares, integrae, usque ad 12 mm lon-
gae, 0.4 mm latae; lamina sterilis 3-pinnata,
plerumque pubescens in superficiebus ambabus;
rhachis et costae pubescentes, squamis ovatis
tenuibus aliquot; pinnulae catadromicae, pro
parte maxima sessiles, usque ad 33 X 9 cm; seg-
menta tertiaria ovalia vel oblonga integra, (5)6—
8(10) X (2)3—4(5) mm. Folia fertilia ignota.

Stem 0.6-1.5 cm thick, hemiepiphytic,
mucilaginous (?); scales dull brown, appressed,
opaque, linear, 0.4—12.1 x 0.3—1.0 mm, margins
subentire. Sterile leaves up to | m long; petiole
Ys to Y2 as long as the lamina; lamina lanceolate
or subdeltate, 30—65(80) x 19—32(60) cm, 3-pin-
nate, usually pubescent on both surfaces, always
so along the veins beneath, the hairs whitish to
tawny, articulated, 0.1—0.4 mm long, spreading;
pinnae to 33 X 9 cm, narrowly triangular to lan-
ceolate, equilateral or, in the basal pair, with the
basiscopic side slightly prolonged, the base trun-
cate, short-stalked, crowding the rhachis; pinnules
catadromic, oblong, the base truncate, nearly ses-
sile, the apex rounded to acute; tertiary segments
ovate to oblong, (5)6—8(10) x (2)4(5) mm, with
several vein branches, the margins entire to
slightly lobed; axes pubescent abaxially, with a
few thin, ovate, brown scales, especially at the
pinnule junctures; grooves nearly glabrous to mod-
erately puberulent within, never packed with long-
protruding hairs, truncated by the ridges of the
next lower order. Fertile leaf unknown.

This new species is named for Robert G.
Stolze, pteridologist at the Field Museum of Nat-
ural History, Chicago, Illinois. He originally
suggested this genus to me and has given much
help and encouragement during the preparation of
this monograph.

Polybotrya stolzei is endemic to the western
Andes of Colombia (Map 9), occurring in wet,
shaded forests from 200-1750 m. The laminar
cutting of P. stolzei is intermediate between P.
lechleriana and P. alfredii. Extremely large pin-
nules of P. stolzei look like those of P. alfredii
but are distinguished by the pubescence on both
surfaces of the lamina and by the blunter apices
of the medial pinnules. Smaller, more delicate
forms of P. stolzei may look like P. lechleriana
but are distinguished by their oblong tertiary seg-
ments with more than one vein per segment. Un-

November 1987 MONOGRAPH of POLYBOTRA 77

FiGure 34. Polybotrya stolzei Moran. a. costal scales from the pinnule-costa junctures; b. abaxial
view of medial pinnules of “c’; c. medial pinna; d. stem scales; e. basal pinna of large leaf; f. adaxial
view of rhachis-costa juncture. a-d: Killip 11549 (GH). e,f: Lellinger & de la Sota 284 (COL).

78 ILLINoIs NATURAL HISTORY SURVEY

fortunately, none of the collections contain fertile
leaves and this structure, therefore, cannot be com-
pared to the distinctive botryoid fertile leaf of P.
lechleriana.

Specimens examined: CoLomBiA. Choco: NW
side of Alto del Buey, Lellinger & de la Sota 280 (COL,
US), 284 (COL, US); principal ridge and slopes 2 km
E of San José del Palmar, 1550-1600 m, Lellinger &
de la Sota 747 (COL, LP, US), 748 (COL, LP, US).
El Valle: Santa Rosa, Dagua Valley, forest along Rio
Caballete, 200—300 m, Killip 11549 (GH, NY, PH, US).

16. Polybotrya alfredii Brade (Fig. 35, Map 8).

Polybotrya alfredii Brade, Bradea 1:12, tab.
1, fig. 2. 1969. Type: Costa Rica. San José:
Tablazo, 1900 m, July 1908, (Brade’s origi-
nal description states August, apparently an
error.) Brade & Brade 98 (holotype: HB;
isotype: NY!).

Polybotrya alfredii Brade f. carpinterae
Brade, Bradea 1:13. 1969. Type: Costa Rica.
San José: Carpintera, terrestrial, 1500 m, 25
November 1908, Brade & Brade s.n.
(holotype: HB).

Polybotrya gracilis Brade, Bradea, 1:14, tab.
1, fig. 3. 1969. Type: Costa Rica. Cartago:
Tablazo, 1900 m, 28 August 1908, Brade
554 (holotype: HB).

Stem 1—2 cm wide, hemiepiphytic, the apex
in living plants usually covered by a thick mucilage
that disappears upon drying; scales dull, dark to
light brown, opaque to translucent, 8—16(22) x 1—
1.5(2.0) mm wide, narrowly lanceolate, spread-
ing, attached across the width of the slightly thick-
ened base, the margin denticulate or more com-
monly entire. Sterile leaves up to 1.4 m X 0.7 m,
petiole up to 35 cm long; lamina ovate, up to
3-pinnate-pinnatifid, subchartaceous, often drying
greyish green, the proximal margins of the pinnule
bases often ciliate, the hairs less than 0.1 mm
long; pinnae up to 45 X 22 cm, apex acuminate;
pinnules up to 12 X 5 cm, ovate to narrowly trian-
gular, the base truncate and symmetrical, those
of the basal pinnae anadromic or subequal, those
of the medial pinnae catadromous; ultimate seg-
ments ovate to oblong, entire, crenulate, serrate
or lobed, the apex obtuse or acute; axes evenly
pilosulous abaxially, the hairs 0.1—0.2 mm long,

Vol. 34, Art. 1

usually tawny; grooves pubescent within, very
shallow or even disappearing within about 1 cm
of the rhachis so that the costa is almost terete
where it joins the rhachis. Fertile leaves up to
0.8 x 0.35 m, up to 3-pinnate-pinnatifid, ovate,
coenosoric; axes pubescent, the hairs short, 0.1—
0.2 mm long, usually with ovate to lanceolate,
appressed, thin, denticulate to fimbriate scales;
sporangial stalks paraphysate; spores (50)55—
62(66) microns long.

Other illustrations: See the original descrip-
tions cited above.

The altitudinal range of Polybotrya alfredii
is 700-1900 m. In Ecuador, this species grows
in rich montane forests in the eastern Andes. In
Costa Rica, this species occurs primarily in wet,
shaded cloud forests, usually growing alongside
P. gomezii. Polybotrya alfredii produces numer-
ous terrestrial leaves that often predominate the
forest floor. Twice in Costa Rica and once in
Ecuador, I saw fertile leaves produced from the
terrestrial portion of the stem. These are the only
instances in which I saw fertile leaves produced
terrestrially by a scandent species.

Polybotrya alfredii looks like P. lourteigiana
but can be distinguished by its more highly cut
lamina. Since leaves lower on the stem tend to be
smaller and less divided than those higher on the
stem, cutting should be compared using pinnae
over 25 cm long (see couplet 24 in the key). The
two species also differ in the shape of the ultimate
segments or lobes; P. alfredii has relatively more
ovate and shorter segments than P. lourteigiana
(compare Figs. 35 & 37).

Unlike all other species of Polybotrya, P.
alfredii has the stem apex and the lowermost 5 to
15 cm of the petiole covered with a thick, translu-
cent mucilage. Mucilage is totally lost upon drying
and therefore not visible on herbarium specimens.
The function of this mucilage is unknown. More
fieldwork is needed to determine if this unusual
mucilage occurs in two closely related species, P.
lechleriana and P. lourteigiana. Nectaries are
another unusual feature of the morphology of P.
alfredii. During fieldwork in Costa Rica, I saw
nectaries on the rhachis of P. alfredii like those
described by Koptur et al. (1982) for P. osmun-
dacea. See the Morphology and Anatomy section
of this monograph for further information concem-
ing nectaries.

79

MONOGRAPH of POLYBOTRA

November 1987

Fe tb yy .
Minis
FG NOES ye vasssommnls

Zak ANAM UN
INS “LE UA Swi}
v CAAA ont "

SOG cS UTI

Wi Wee, i Wi Ly
a, MiIRNN\ Ty

SR

iy Ne 4
i om mee.

ie Put y' ij

CF)

al
Ay

PUSS.

Y) Aj
Re

: Mig F

NY

7, Lt Lh f , ; 4)
POON

77)

rya alfredii Brade. a. sterile leaf, b. stem scales, the middle shown sideways;

c~g. pinnules, acroscopic side to the right; h. fertile pinnule, adaxial view; i. adaxial su

FiGure 35. Polybot

rface of costa

and pinnule base. a,b,f,h,i: Moran 2442 (CR). c: Moran 3532 (F). d: Buchtien 5164 (U

3214 (CR). g: Dwyer 8337 (MO).

S). e: Moran

80 ILLINOIS NATURAL HISTORY SURVEY

I put P. gracilis in synonymy with P. alfredii,
although the former appears distinct because the
leaf is smaller, only 2-pinnate, and has obtuse
pinnule apices (Fig. 35e). Polybotrya gracilis re-
sembles P. alfredii, however, in such characteris-
tics as scales, pubescence, stem mucilage, habitat,
and range. Furthermore, at several localities in
Costa Rica, I collected plants that exhibited com-
plete intergradation between the typical large
leaves of P. alfredii and the smaller ones of P.
gracilis. Polybotrya gracilis, therefore, represents
a small-leaf form of P. alfredii that has well-
rounded pinnule apices. I chose the name P. al-
fredii over P. gracilis because an isotype has been
seen. This species is named in honor of Alexander
Brade’s brother and field companion, Alfred
Brade.

Specimens examined: NICARAGUA. Rivas: Isla
de Ometepe, NW slope of Volcan Maderas, Stevens &
Krukoff 6518 (CR); Graytown, Camp Murroeal, Shimek
& Smith s.n. (MICH).

Costa Rica. Alajuela: Univ. of San Ramén
biological field station, Moran 3214 (CR, F, GH, MO);
12 km NNW of San Ramon by road on way to San
Lorenzo, 1 km S of Balsa, Liesner & Judziewicz 14855
(CR, MO); San Jerénimo, Wercklé 559 (US); Suerte,
llanuras de Santa Clara, Donnell Smith 6939 (US); Santa
Clara, Cooper 10240 (US); NW of Zarcero, ca. 2 km
W of Zapote on dirt road to Sta. Elena, Lellinger &
White 1361 (US); San Antonio de Zarcero, A.C. Smith
48/293 (US); N of San Ramén, ca. 4 km N of Balsa
along road to Colonia Palmerena, Lellinger & White
1244 (US); Santiago de San Ramon, Brenes 21980 (F),
14248 (GH); Alto de la Palma de San Ramén, Brenes
11676 (F, NY); 7 miles N of San Ram6n square along
ridge road, Evans & Bowers 2944 (MO); Finca La Selva,
San Rafael de Vara Blanca, pendiente N del Volcan
Barba, Jiménez 803 (F); vicinity of La Palma, Maxon
412 (NY); 11 km N of San Ramén, Mickel 2958 (LP,
NY, UC); San Luis de Zarcero, A.C. Smith 778 (NY).
Cartago: road between La Suiza and Tuis, DeWolf 385
(CR); Pacayas, Lankester 653 (US); along Inter-Amer-
ican Hwy. between Cartago and San Isidro del General,
Scamman 5985 (CR), 5986 (CR); Orosi, Finca del Sr.
Valverde, Brade 16824 (CR); El Muneco, S of Navarro,
Standley 33648 (US); Santa Clara de Cartago, Maxon
& Harvey 8242 (US); SE of Platanillo, along Camino
Raiz de Hule, Croat 3676] (MO); Platanillo, de la Sota
5252 (LP, US); foot of Volcan Turrialba, Pacayas, Scam-
man 7155 (CR); along road from Moravia to Quebrada
Platanillo (Tsipiri), Croat 36669 (MO); ca. 22 km E of
Turnalba, high ridge above Platanillo, Mickel 3426 (LP,
NY); near the town of Estrella, Moran 2237 (CR), 2241
(CR); near Alto Patillos, Moran 2442 (CR); Reserva e
Tapanti, Gémez 18906 (CR, MO, UC); Tapanttf,
Grayum & Sleeper 3448 (MO); 12-16 km S of Tapanti,

Vol. 34, Art. 1

along the road above the Rio Grande de Orosi, Stolze
1488 (F, UC). Heredia: 7.5 km N of Vara Blanca, in
vicinity of Rio La Paz Grande, Croat 36007 (MO); Vara
Blanca de Sarapiqui, N slope of Central Cordillera,
Skutch 3236 (GH, US); Santo Domingo de Vara Blanca,
Valerio 2356 (US); Vara Blanca, between Volcan Poas
and Barba, Chrysler & Roever 5095 (US); 4 km N of
Vara Blanca, Croat 35608 (MO); near Rio Las Vueltas,
NE of Volcan Barba, Lent 2642 (F); Virgen del Socorro,
Moran 3168 (CR). Puntarenas: 5 km NE Finca las
Cruces, San Vito de Java, Burch 4505 (NY); near field
station, Rincén de Osa, Burch 4414 (NY); 5 km S of
San Vito de Java, in vicinity of biological field station
at Wilson finca, Mickel 2003 (NY); 5 km S of San Vito
de Java, 1-4 km SW of biological station at Finca Wil-
son, Mickel 3126 (NY). San José: Tablazo, Valerio 63
(US); Biolley 73 (US, Z); Tablazo, Brade & Brade 67
(P), 98 (NY); Alto La Palma, Moran 2325 (CR).

PANAMA. Chiriquf: Palo Alto, just E of Boquete,
Stern et al. 1034 (CR, US); Holcomb trail, Cormman
927 (MICH, UC, US); vicinity of El Boquete, Cornman
1178 (US), 1233 (US), Maurice 699 (US); Rio Caldera,
from El Boquete to the Cordillera, Killip 5202 (CR,
US); valley of Rio Piarnasta, about 5 mi E of Boquete,
Killip 5163 (US), 5386 (US), 5429 (CR, MICH, US).
Coclé: Valle de Ant6n, Ocampo 877 (CR); Cerro Pilon,
Dwyer 8337 (MO); cloud forest at La Mesa above El
Valle, Croat 13431 (MO, UC).

EcuaApDor. Napo: camino Baeza—Tena, 15 km al
norte de Tena, 1100 m, Moran 3532 (F, GH, PORT,
Q, QCA). Pastaza: village of Rio Chico, 8 km from
Puyo, Shemluck 304 (F). Tungurahua: casi 25 km al
este de Banos, detras el pueblo Rio Negro, por Rio
Topo (malempresado “Rio Toro” en el mapa), Moran
3570 (F, GH, Q, QCA).

PERU. Huanuco: SW slope of the Rio Llulla Pichis
watershed on the ascent of Cerro del Sira, Dudley 13258
(GH), 18265A (GH).

BotiviA. La Paz: Hacienda Simaco sobre el
camino a Tipuana, Buchtien 5164 (GH, US).

17. Polybotrya botryoides (Baker) C. Chr. (Fig.
36, Map 8).

Polybotrya botryoides (Baker) C. Chr., Index
Filicum 504. 1906.

Acrostichum botryoides Baker, J. Bot.
19:206. 1881. Type: Colombia. Antioquia:
on trees in the forest, 6000—7000 ft, 22 July
1880, Kalbreyer 1873 (holotype: K; photo
and fragment GH!, US!).

Stem to 2 cm thick, hemiepiphytic; scales dull
brown, about | cm long, with the margins suben-
tire to erose, the base curved and thickened. Sterile
leaves up to 1.5 m (2); petiole densely scaly at

November 1987 MONOGRAPH of POLYBOTRA

0.5 mm

£
£
if 11cm

vay, y SES

ST aes
KR Moran is

FiGurE 36. Polybotrya botryoides (Baker) C. Chr. a. costal scales, the one on the right enlarged to
show the biseriate apex; b. uniseriate “hair” from the costa. Such hairs are actually reduced scales as
shown by their intergradation with the scales and by their characteristics which differ from those of
true hairs; c. sterile pinnule base; d. botryoid fertile pinnule; e. sterile pinnule; f. stem scale, at right
is a side view showing the thickened base; g. petiolar scale, medial portion darkened and margins

erose. a—g: Juncosa 1369 (MO).

82 ILLINOIS NATURAL HisTORY SURVEY

base, the scales similar to those on the stem, but
with wider pale borders and more strongly erose;
lamina 4-pinnate, nearly glabrous on both sur-
faces; pinnae lanceolate, up to 35 x 20 cm, finely
divided all the way to the apex; pinnules up to
10 x 3 cm, catadromic or subequal, lanceolate,
the base often overlapping the costa, subequilat-
eral, i.e., not prolonged acroscopically, short-
stalked (1-2 mm long); tertiary segments oblong,
sessile, up to 2 X 0.9 cm, apex acute or rounded,
base equilateral; quaternary segments oblong to
ovate, sessile, up to 5 x 3 mm, the margins entire
or crenate; axes scaly, the scales numerous, tortu-
ous, thin, reddish, most of these only 2—5 cells
wide, intergrading with hairs, these articulate, red-
dish, 4—8 celled, 0.1—-0.3 mm long; grooves
pubescent within by reddish hairs. Fertile leaves
botryoid, 3-4-pinnate, with the ultimate soriferous
segments globose, 1-2 mm diameter; receptacle
paraphysate by articulate hairs, these often buried
among the sporangia; sporangial stalks paraphy-
sate; spore size unknown.

Polybotrya botryoides is endemic to the
Andes of Colombia (Map 8), occurring in forests
from 1830-2130 m. The above description is
based primarily on the Juncosa collection, which
was the only dried specimen I had available for
study. I could not find enough spores on this speci-
men to make an adequate measurement of spore
s1Ze.

This rare species can be separated from all
other species of Polybotrya by its distinctive com-
dination of botryoid fertile leaf, 4-pinnate sterile
leaf, and axes covered by numerous, linear, tortu-
us, reddish scales (Fig. 36a). No other species
of Polybotrya is as finely divided as this one.
Polybotrya botryoides looks like P. alfredii be-
sause of its large decompound lamina, catadrom-
cally arranged pinnules, subequal pinnule bases,
pubescence of the axes, and dull brown stem
scales. The Venezuelan endemic P. canaliculata
ilso has botryoid fertile leaves, but it lacks hairs
yn the axes, has very different axial scales, and
S less finely cut (Fig. 52).

Specimens examined: COLOMBIA. Antioquia:
Kalbreyer 1873 (photo and fragment GH, US); main
wy. Medellin—Puerto Triunfo, Municipality Cocorna,
‘a. 5 km E of Cocorna Peaje, Quebrada El Biadal, 1830
n, Juncosa 1369 (MO, UC).

18. Polybotrya lourteigiana Lellinger (Fig. 37,
Map 10).

Polybotrya lourteigiana Lellinger, Proc.
Biol. Soc. Washington 89:723, fig. 5. 1977.
Type: Colombia. Choco: Trail along ridge
from the confluence of the forks of the Rio
Mutata above the Rio Dos Bocas to the top
of Alto del Buey, ca. 1450-1750 m, Lellinger
& de la Sota 251 (holotype: US!; isotypes:
COL!, LP).

Stem to 2 cm thick, hemiepiphytic, mucilagin-
ous (?); scales dark brown, concolorous, and en-
tire, or cream-colored with a darker center and
ciliate margins, mostly 8—17 x0.5-2.0 mm.
Sterile leaves up to 1.6 m long; petiole up to 2
as long as the lamina; lamina glabrous on both
surfaces, up to 3-pinnate but mostly 2-pinnate-
pinnatifid throughout, deltate to lanceolate; lower
pinnae mostly 30—45 x 10-18, elliptic, lanceo-
late or narrowly triangular; pinnules arranged
catadromically, subequilateral at base, subsessile,
narrowly oblong, generally 10—15 free pairs, S—
8(11) X 1.5—2.0(3.0) cm, the apex slightly fal-
cate; tertiary segments 5—12(15) X 2—4(6) mm,
arranged catadromically, oblong, the apex
rounded to truncate, the margins entire; axes
pubescent abaxially, the hairs either tawny or red-
dish, 0.5—1.2 mm long, multicellular, lax, or with
the hairs shorter, stiffer, 0.1—0.3 mm long, costal
scales present, inconspicuous, often at the
junctures with the costules, ovate to orbicular,
flaccid, brown; grooves pubescent within, espe-
cially at the junctures, truncated by the ridges of
the next lower order. Fertile leaves 4-pinnate,
coenosoric, or subbotryoid, if the sori discrete,
then not on long stalks; costa pubescent, with or
without ovate, denticulate scales; sporangial
stalks paraphysate; spores (S0)S5—67(70) microns
long.

Other illustrations: See Lellinger’s original
description (cited above) which contains a photo-
graph of a portion of the holotype.

This plant occurs in mid-elevation forests
from 750 to 2250 m in the western Andes of Co-
lombia (Map 10). See comments under Polybotrya
alfredii and P.. pittieri for comparisons with those
species. Considerable morphological variation
exists in the specimens cited below, but I found

Vol. 34, Art. 1

November 1987 MONOGRAPH of POLYBOTRA 83

FiGure 37. Polybotrya lourteigiana Lellinger. a. basal pinnae; b. pinnule bases, apex to the left;
c,d. sterile pinnules, apex to the left; e. stem scales; f. fertile pinnules. a,b,c,e,f: Lellinger & de la
Sota 25] (US). d: Cuatrecasas 15527 (US).

84 ILLINOIS NATURAL HISTORY SURVEY

no constant characters to separate the extreme
forms. Pubescence varies tremendously; the Lel-
linger and de la Sota specimens are mostly long
pilose on the axes whereas the Cuatrecasas speci-
mens have shorter, stiffer hairs. Intermediates
exist, and no other characters correlate with pubes-
cence.

The type collection consists of six sheets, of
which US #2748016 is here excluded; it actually
represents P. stolzei.

Specimens examined: CoLomsiA. Choco: Trail
along ridge from the confluence of the forks of the Rio
Mutata above the Rio Dos Bocas to the top of Alto del
Buey, ca. 1450-1750 m, Lellinger & de la Sota 251
(COL, LP, US); principal ridge and slopes 2 km E of
San José del Palmar, Lellinger & de la Sota 768a (COL,
LP, US); 0.3 km E of the Ciudad Bolivar—Quibd6 road
across the suspension bridge at ca. km 141, Lellinger
& de la Sota 899 (COL, LP, US); Mojarras de Tado,
8.5 km E of Istmina, Lellinger & de la Sota 387 (CR,
COL, LP, US). Chocé-Antioquia: principal ridge W
of La Mansa, at ca. km 105.5 of the Ciudad Bolivar—
Quibd6 road, Lellinger & de la Sota 948 (COL, LP,
US). Choc6—EI Valle: Serrania de Los Paraguas, along
the trail from El Cairo to Rio Blanco, ca. 8 km SW of
EI Cairo, Lellinger & de la Sota 843 (COL, LP, US).
EI Valle del Cauca: Hoya del Rio Cali, La Margarita,
Duque-Jaramillo 1868 (COL); Cordillera Occidental,
vertiente occidental, del Rio Sanquinini, La Laguna,
Cuatrecasas 15527 (US); Cordillera Occidental, ver-
tiente occidental, monte La Guarida, filo de la cordillera
sobre La Carbonera, entre Las Brisas y Alban, Cuatre-
casas 22143 (F, US). Narino: Municip. de Altaquer,
Carretera entre Altaquer y Junin, Guyambe, Mora 4160
(COL).

19. Polybotrya pittieri Lellinger (Fig. 38, Map
11).

Polybotrya pittieri Lellinger, Proc. Biol. Soc.
Washington 89:725, fig. 6. 1977. TyPE: Co-
lombia. Cauca: Dagua Valley, Cordoba,
Pacific Coastal Zone, 30-100 m, Pittier 587
(holotype: US!).

Stem | cm thick, hemiepiphytic; scales 9-14
x 1.0-1.4 mm, spreading, reddish, membrana-
ceous, the margin denticulate. Sterile leaves up
to 1.3 m long; petiole to 38 cm long; lamina trian-
gular at base, to 60 x 90 cm, 3-pinnate at the base,
2-pinnate-pinnatifid medially, nearly glabrous on
both surfaces, the apex long-tapered; pinnules ar-
ranged catadromically, up to 5 X 2 cm, relatively
short and stubby, the base subequilateral, the apex
obtuse or slightly acute; tertiary segments up to

Vol. 34, Art. 1

1.0 X 0.5 cm, oblong; axes pubescent on the ab-
axial surface, the hairs short, stiff, ca. 0.2 mm
long; grooves truncated by the ridges of the next
lower order or only indistinctly admitted to the
groove, pubescent within, the hairs reddish or
tawny, 0.2—0.5 mm long. Fertile leaves subtri-
angular, 4-pinnate, botryoid, the apex long-
attenuate; pinnules to 2.1 X 1 cm; sori stipitate,
round, the stalks 1-4 mm long, arranged pin-
nately; sporangial stalks paraphysate; spores
(62)65—70(77) microns long.

Other illustrations: Lellinger’s original de-
scription has a photograph of the type.

Polybotrya pittieri, known from only two col-
lections, is endemic to the western Andes of Co-
lombia (Map 11). This species occurs at low ele-
vations, from 30—100 m, unlike its most closely
related species, P. attenuata and P. botryoides,
which occur at higher elevations. It is named after
Henry F. Pittier (1857-1950), a Swiss botanist
and civil engineer who collected prolifically in the
American tropics.

Polybotrya pittieri looks most like P. lour-
teigiana. Lellinger (1977) observed only that P.
pittieri had a rather stiffly pilose indument on the
thachis and costae, whereas P. lourteigiana had
a lax, hirtous indument. From the greater number
of specimens of P. lourteigiana available to me,
however, pubescence type does not appear con-
stant. Nevertheless, P. pittieri seems to be a dis-
tinct species, and the best way to distinguish it is
by its botryoid fertile leaves with prominently
stipitate sori (Fig. 38b). P. lourteigiana, on the
other hand, has coenosoric fertile leaves and sub-
sessile sori (Fig. 37f). Another difference, al-
though one that may prove inconsistent when more
specimens are known, is the broadly deltate lamina
of P. pittieri; P. lourteigiana is slightly reduced
at the base. Using these characters, I find that the
paratypes of P. pittieri actually belong to P. lour-
teigiana.

Polybotrya pittieri might also be confused
with P. canaliculata since both have botryoid fer-
tile leaves, but P. pittieri differs by its broadly
deltate lamina, pinnule cutting, and pubescent
axes that lack scales. Its spores are extremely large
compared to others in the genus, suggesting that
P. pittieri is a polyploid.

Specimens examined: CovompBia. Cauca:
Dagua Valley, Cérdoba, Pacific Coastal Zone, 30-100
m, Pittier 587 (US); El Tambo, von Sneidern 1590 (US)

85

MONOGRAPH of POLYBOTRA

November 1987

Sh7
ee.

Aj

it
it
a
iP

FiGuRE 38. Polybotrya pittieri Lellinger. a. sterile leaf; b. fertile leaf; c. stem scales; d. sterile basi-

scopic pinnule from basal pinna. a—d: Pittier 587 (US).

86 ILLINOIS NATURAL HISTORY SURVEY

20. Polybotrya cylindrica Kaulfuss (Fig. 39,
Map 16).

Polybotrya cylindrica Kaulfuss, Enum.
Filicum 56. 1824. Type: Brazil. Santa
Catarina: Island of St. Catharina, Chamisso
s.n. (LE).

Polybotrya incisa Link, Hort. Berol. 2:135.
1833. Type: Brazil. No collector or date
given (holotype: B!).

Acrostichum incisum (Link) Hooker, Species
Filicum 5:245. 1864.

Acrostichum cylindricum (Kaulfuss) Hooker,
Species Filicum 5:246. 1864.

Polybotrya frondosa Fée, Crypt. Vasc. Bré-
sil. 1:15. 1869. Type: Brazil. No exact lo-
cality given, Glaziou 2428 (holotype: K;
photo GH!).

Polybotrya osmundacea_ var. cylindrica
(Kaulfuss) Luetzelb. Estudo Bot. Nordeste
Brazil 3:245. 1923.

Polybotrya osmundacea var. frondosa (Fée)
Luetzelb. Estudo Bot. Nordeste Brazil 3:245.
1923.

Polybotrya osmundacea var. incisa (Link)
Luetzelb. Estudo Bot. Nordeste Brazil 3:245.
1923.

Stem 1—2 cm thick, hemiepiphytic; scales dull
brown, opaque, concolorous or with a very narrow
hyaline border, entire, generally 6—12 mm long,
the base thickened, curved, appressed. Sterile
leaves up to 1.7 m long; petiole 30—SO cm long;
lamina to 1.3 m long, 3-pinnate-pinnatifid, gla-
brous or pubescent with small, unicellular, whitish
hairs, the margins sparsely to moderately ciliate
by jointed hairs, these ca. 0.1 mm long; pinnae
narrowly lanceolate to subdeltate, the lowermost
to 52 x 28 cm; pinnules generally 9-14 x 3.5 x
5.5 cm, anadromic proximally, becoming cata-
dromic distally, the acroscopic side slightly pro-
longed, proximal ones with a short 3—5 mm stalk;
tertiary segments ca. 7—11 per pinnule, ovate to
oblong, the largest usually pinnatisect, the apex
acute or rounded; vein tips slightly clavate and
prominulous; axes abaxially sparsely to densely
pubescent, the hairs colorless, clear, stiff, subulate
ca. 0.1—0.3 mm long, with the cross walls not
easily visible, mixed with appressed, linear,
scraggly, inconspicuous scales; grooves filled with

Vol. 34, Art. 1

reddish hairs. Fertile leaves to 90 X 62 cm, 3-4-
pinnate; ultimate segments often oblong and ap-
pearing cylindrical due to folding back of the seg-
ment margins; sporangial stalks paraphysate;
spores (55)60—66(70) microns long.

Other illustrations: Fée, Mém. Fam. Foug.
2 (Hist. Acrost.), tab. 36. 1846; op. cit. tab. 35
(as P. incisa); Brade, Bradea, 1(9):66, figs. 1 &
2; 67, figs. 16 & 17. 1969; Sehnem, Flora Ilustrada
Catarinense, plate 19; plate 20 (as P. cylindrica
var. frondosa). 1979.

Polybotrya cylindrica, endemic to the coastal
mountains of southeastern Brazil (Map 16), is the
largest and most finely divided species of Polybot-
rya in Brazil. It occurrs in wet, shaded, primary
forests from 0 to 775 m. This species differs further
from other Polybotrya species in southeastern
Brazil by its stem scales that are dull brown with
entire margins and the thickened and curved bases.
The curved scale base orients the rest of the scale
parallel to the surface of the stem, so that the
scales are appressed (Fig. 39f,g). Another helpful
feature in identification is the laminar margin,
which is sparsely ciliate by jointed hairs less than
0.1 mm long.

Brade (1971) distinguished P. frondosa from
P. cylindrica by the amount of lamina dissection.
But lamina dissection exhibits complete intergra-
dation from 2-pinnate to 3-pinnate-pinnatifid and
no other characterisitcs correlate with it (Fig.
39a-d).

Although I have not seen the type of this
species, the original description by Kaulfuss is
sufficient to ascertain his plant’s identity. He spe-
cifically mentions the ciliate margins that distin-
guish this plant from others in southeastern Brazil.
Furthermore, this species is the only Polybotrya
that grows on the Island of Santa Catarina, where
the type was collected. The specific epithet prob-
ably refers to the fertile segments which, upon
drying, reflex so that the sori appear on all sur-
faces, thereby imparting a cylindrical shape to the
segments.

Specimens examined: BRaAziL. Espirito Santo:
Municipio de Cachoeira de Itapeminm—Vargem (?),
Brade 19972 (HB, MO, NY). Parana: Mun. Guaratuba,
Pedra Branca de Araraquara, Hatschbach 8/41 (US);
Pedra Branca de Araraquara, Hatschbach 7424 (US);
no locality, Hatschbach 10749 (NY); no locality, Dusén
13658 (GH); Porto Dom Pedro Il, Dusén 4423(BM);

November 1987

MONOGRAPH of POLYBOTRA

iE; 2D
XN

B;
SS
Ns

\

aE

FiGureE 39. Polybotrya cylindrica Kaulfuss. a. sterile pinna; b—d. sterile pinnules showing variation
in amount of dissection; e. fertile pinnules; f. stem scales, top view; g. stem scale, side view

a,e: Goedas 217 (NY). b: Mosen 3050 (B). c,f,g: Schmalz 158 (UC). d: Dusén 6926 (F).

88 ILLINOIS NATURAL HISTORY SURVEY

Volta Grande, Dusén 6984 (BM); Porto de Cima, Dusén
6926 (F, GH, LIL, MO, NY, PH, UC, Z); Tacarehy,
Dusén 15351 (BM, F, PH), 15353 (GH, MO, PH, UC,
Z). Rio de Janeiro: without locality, Hunnewell 18511
(MICH); Distrito Federal, Guanabara, Represa Camori,
Brade 12576 (RB); Rezende, 80 mi WNW of Rio, on
Paraiba River, Rio Palmital, Castellanos 25710 (F);
Serra de Itatiaia, Brade 12614 (NY , RB); Brene, Glaziou
955(RB); vicinity of Meio da Serra, Smith & Brade 2286
(GH); Estrada Velha de Barrira a Teresopolis, Duarte
& Pereira 65323 (F, LP); Corcovado, Miers s.n. (NY);
Teresopolis, Brade 9365 (NY, UC); Serra da Carioca,
Brade 13982 (RB). Rio Grande do Sul: S. Leopoldo,
Lehnun 2562 (LIL); Porto Alegre, Fargens 25] (UC).
Santa Catarina: Municip. [birama, Horto Florestal,
Smith & Klein 7546 (US); Horto Florestal, I.N.P.,
Ibirama, Reitz & Klein 11/01 (US), 1102 (US), 2665
(US), 3466 (US); Blumenau, Warnow, Goedas 217
(NY, UC, US); Itajai, Reitz /5/ (LIL); Municipality
Brusque, Azambruja, Smith and Reitz 6135 (COL, GH,
MO, US); Reserva Florestal, dos Piloes, Duarte 3/79
(LP, NY); Mina Velha, Garuva, Sao Francisco do Sul,
Reitz & Klein 4635 (US), 4682 (US); Sabia, Vidal
Ramos, Reitz & Klein 5108 (US); Passo Mansa, Haer-
chen 124 (NY); Joinville, Schmalz 78 (MO), 158 (NY,
UC); Tres Barras, Garuva, Sao Francisco do Sul, Reitz
& Klein 5549 (US); Sta. Catherine 1834, Gaudichaud
73 (P); Azambuja—Brusgui, Reitz 2825 (LIL, P);
Blumenau, Muller 6039 (MICH). Sao Paulo: Sao Bento,
Luetzelburg 254 (MICH, US); Serra da Bocaina, Brade
20931 (LP); Morro das Pedras, Iguape, Brade s.n. (US);
Paranapicaba, Biological Station, Handro 1229 (GH,
US); Agua funda, Handro 2228 (GH, US); Serra do
Mar, Wacket 21556 (GH); Campo Grande to Alto da
Serra, 40 km SE of Sao Paulo, Tryon & Tryon 659]
(GH); Santos, 1874, Mosen 3050 (B, P). State un-
known: Glaziou 2428 (photo of K specimen at GH); no
locality, Claussen s.n. (P); Micken s.n. (NY); Capao do
Fradie, Schneus 3412 (GH); Morro do Antao, Sehnem
3092 (GH).

21. Polybotrya hickeyi Moran, sp. nov. (Fig.
40, Map 13).

Polybotrya hickeyi Moran, sp. nov. TYPE:
Bolivia. Cochabamba: Prov. Chapare, road
from Cochabamba to Villa Tunari, in thick
rain forest, very heavily shaded, on trunk of
Ficus 1700 m, 23 March 1980, Hickey 80]
(with Eshbaugh) (holotype: GH!; isotype:
MU).

Caulis hemiepiphyticus, 1-2 cm diam.; squa-
mae atrobrunneae, vulgo opacae, leviter nitidae,
concolores vel marginibus tenuioribus et pal-

Vol. 34, Art. 1

lidioribus ad apicem; lamina usque ad 3-pinnata-
pinnatifida, deltata, usque ad | m longa, leviter
pubens inferne, pilis cylindricis, appressis, usque
ad 0.1 mm longis; pinnae usque ad 46 X 25 cm,
deltatae, paribus proximalibus acroscopice pro-
longatis; pinnulae catadromicae supra basin, del-
tatae, oblongae vel lanceolatae; costae sine sulcis
prominentibus; venae prominulae superne, apice
leviter incrassatae. Sori discreti, rotundi, oppositi
vel suboppositi.

Stem 1-2 cm thick, hemiepiphytic; scales
dark chocolate brown, mostly opaque, concolor-
ous, or the apex with thinner, lighter borders, the
margins denticulate to entire, the base thickened,
attached across its full width. Sterile leaves up to
1.5 m (?); petiole with thin, ovate to lanceolate
scales; lamina to 3-pinnate-pinnatifid, deltate, to
about 1 m long, the abaxial surface slightly pu-
berulent, the hairs cylindrical, appressed, up to
0.1 mm long; pinnae up to 46 X 25 cm, deltate,
the proximal pair basiscopically prolonged, the
distal ones gradually becoming acroscopically pro-
longed; pinnules catadromic above the base, del-
tate to oblong-lanceolate, the basiscopic margins
thickened and decurrent on the costa, the base
slightly prolonged acroscopically, the apices
acuminate; tertiary segments also with a thick-
ened, decurrent, light-colored basiscopic margin;
veins prominulous adaxially, the tips thickened
and ending just behind the margin; costules bor-
dered by a raised, erect flap or wing of tissue;
axes often not prominently grooved, pubescent by
ca. 0.1 mm long, colorless, erect hairs, the scales
appressed, denticulate, flexuose. Fertile leaves
botryoid; sori with few fusions, arranged mostly
oppositely or suboppositely.

Polybotrya hickeyi grows in high elevation
forests in the Andes of Colombia and Bolivia; it
has not yet been collected from Ecuador or Peru
(Map 13). This species looks most like P.
puberulenta—see that species for comparison.
This plant is named for one of the collectors, Dr.
R. James Hickey, who has made important con-
tributions to the study of /soetes and Lycopodium.

The Colombian specimens look like the Bo-
livian type specimen except for the stem scales.
The Bolivian type has dark, chocolate brown
scales with denticulate margins and lighter borders
apically (Fig. 40g). The Colombian specimens
have merely dull brown, concolorous scales with

November 1987 MONOGRAPH of POLYBOTRA 89

FiGure 40. Polybotrya hickeyi Moran. a. fertile pinnules; b,c,d. sterile pinnules, acroscopic side to
the right; e. medial pinna; f. basal pinna (note that the two proximal basal pinnules are fertile); g. stem
scales (Bolivia); h. stem scale (Colombia); i. abaxial surface of distal pinnules. a,d: Ludwig 262 (NY).
b,h: Grant 10283 (F). c: Killip & Smith 20210 (US). e,f,g,i: holotype, Hickey 80/ , with Eshbaugh (GH).

90 ILLINOIS NATURAL HISTORY SURVEY

entire margins and a prominently curved and thick-
ened base; therefore, they appear appressed (Fig.
40h). This thickened and curved scale base appears
somewhat elevated, like those of P. caudata and
P. cylindrica . The lamina of P. hickeyi turns a
peculiar yellowish green upon drying.

Specimens examined: CoLcomBia. Cauca: W
slope of W Cordillera, above Carpinteria, Alston 8278
(BM). Cundinamarca: Cordillera Oriental, Toquiza,
Gazaunta Valley, Cordillera de Helicona, 15 km NW
of Medina, 2200 m, Grant 10283 (F, US); Fusagasuga,
Ludwig 262 (BM, MO, NY). Norte de Santander: Pica-
Pica Valley, above Tapata (N of Toledo) 2100-2400
m, Killip & Smith 20210 (GH, NY, US).

Bolivia. Cochabamba: Proy. Chapare, road from
Cochabamba to Villa Tunari, 1700 m, Hickey 801 with
Eshbaugh (GH).

22. Polybotrya puberulenta Moran, sp. nov.
(Fig. 41, Map 13).

Polybotrya puberulenta Moran, sp. nov.
Type: Ecuador. Napo: camino Baeza—Tena,
5 km al sur de Cosanga. Cordillera Oriental,
bosque perturbado, 1100 m, 4 de enero 1984,
Moran 3528 (holotype: F!; isotypes: COL!,
GH!, LPB!, MO!, NY!, Q!, QCA!, UC!,
US!, VEN!).

Caulis hemiepiphyticus; squamae aurantiacae
sordidae vel luteae, membranaeae, late patentes,
centro leviter fuscatae, marginibus denticulatis vel
erosis; lamina 2-pinnata-pinnatisecta perlate
ovata, crassa, flavovirentes pallide, pubents utrin-
que, pilis patentibus, flexuosis, ca. 0.1 mm longis;
pinnae usque ad 34 X 18 cm, infimus basiscopice
prolongatae; pinnulae catadromicae oblongae, in
margine incrassatae et decurrentes; lobi tantum
crenati, apice truncati vel rotundati; venae pro-
minulae superne, apice leviter incrassatae. Folia
fertilia 4-pinnata, axibus dense pubescentibus,
pilis 3—5 mm longis et squamis linearibus, appres-
sis vel leviter patentibus; segmenta opposita vel
subopposita, receptaculo pubescenti, pilis 1-2 mm
longis, articulatis.

Stems 1-2 cm thick, hemiepiphytic; scales
yellow to sordid orange, darker in the center with
light yellow borders, membranous, spreading, I—
3 mm wide, the margins erose to denticulate, at-
tached across the width of the narrowed base, in
fresh material somewhat rugose transversely.

Vol. 34, Art. 1

Sterile leaves up to 1.2 m long; petiole up to 27
cm long; lamina to 2-pinnate-pinnatisect, broadly
ovate, thick and somewhat fleshy in living mater-
ial, color light yellowish green, puberulent on both
surfaces, the hairs spreading, flexuous, ca. 0.1
mm long, intergrading with the scales on the lower
surface; pinnae up to 34 * 18 cm, the proximal
pairs basiscopically prolonged, the distal ones grad-
ually becoming equilateral or slightly acroscopi-
cally prolonged; pinnules catadromic throughout,
oblong, the basiscopic margin thickened, decur-
rent on the costa, the apices obtuse to acute, base
cuneate and slightly prolonged acroscopically;
costules bordered by a raised flap of lighter colored
tissue; lobes of the largest pinnules merely crenate,
the margins thickened, the apices obtuse or trun-
cate; veins prominulous adaxially, the tips slightly
thickened and ending just behind the margin; axes
evenly pubescent abaxially, the hairs 0.1 mm long,
tawny, erect, scales caducous, usually lacking;
grooves pubescent within, becoming shallow and
flat at the junctures, not admitted to the groove
of the next lower order. Fertile leaves 4-pinnate,
the axes densely pubescent, the hairs jointed, 3-5
mm long, the scales S—10 mm long, linear, ap-
pressed to slightly spreading; sori opposite or sub-
opposite; sporangial stalks paraphysate; recepta-
cle pubescent by jointed hairs, 1-2 mm long, that
protrude beyond the sporangia; spore size un-
known.

Polybotrya puberulenta grows at high eleva-
tions in the Andes of Ecuador, Bolivia, and prob-
ably Peru (Map 13). The specific epithet refers to
the sterile lamina, which is puberulent on both
surfaces. Another distinctive feature of this species
is the stem scales, which are relatively wide (I-3
mm), thin, spreading and yellowish when viewed
with transmitted light. The lamina is relatively
thick and fleshy; the mesophyll cells apparently
collapse when dry and the veins become promin-
ulous. The lamina, which is always broadest at
the base, has a peculiar light greyish green color
when dry.

This species differs from P. hickeyi in several
characteristics of the stem scales, pinnule arrange-
ment, lamina dissection, pubescence, paraphysis
length, and scalyness of the major axes. Both
species, however, have thickened, decurrent,
basiscopic margins. The medial, less cut pinnules
of P. hickeyi may approach in outline those of P.
puberulenta, but the largest pinnules of the basal

November 1987 MONOGRAPH of POLYBOTRA

a
4

SSNS j
Supp SS

tig y) Nee, anil

me

Poy

IGURE 41. Di eh otrya ee rulenta Mor: nules;
surface of costa and pin as Mor 1 3528 ©.

92 ILLINOIS NATURAL HistORY SURVEY

pinnae are much more highly divided in P. hickeyi
(compare Figs. 40f and 41a).

Unfortunately, Rusby’s Bolivian collection
consists only of the fertile leaf, but I believe it
belongs to this species because of its cutting, in-
dument type, and yellowish stem scales. The long,
protruding paraphyses are easily seen with a hand
lens and offer an excellent character to distinguish
this species from P. hickeyi.

Specimens examined: ECuApDoR. Napo: camino
Baeza—Tena, 5 km al sur de Cosanga, Cordillera Orien-
tal, bosque perturbado, 1100 m, Moran 3528 (COL, F,
GH, LPB, MO, NY, Q, QCA, UC, US, VEN).

BOLiviA. La Paz: Yungas, 6000 ft, “only the fertile
frond collected,” Rusby 443 (NY).

23. Polybotrya alata Moran, sp. nov. (Fig. 42,
Map 14).

Polybotrya alata Moran, sp. nov., TYPE:
Panama. Panama: Campo Tres, 3 mi NE of
Altos de Pacora, primary forest, alt 5|00—800
m, epiphytic, the caudex closely appressed,
sterile, 10 March 1973, Croat 22706
(holotype: MO!; isotype: L!).

Caulis hemiepiphyticus usque ad 2.5 cm
diam.; squamae lineares, 15-25 X 0.8—1.2 mm,
chrysobrunneae, concolores vel centris leviter fus-
catis, appressis, ascendentibus, marginibus den-
ticulatis. Lamina 2-pinnata-pinnatifida, superne
glabra, inferne pubens, pilis circa 0.1 mm longis,
niveis, subulatis; pinnae usque ad 30 X 8 cm, an-
guste triangulares, apicibus attenuatis, zonis
mediis alatis secus costam; pinnulae catad-
romicae, usque ad 6 X 2 cm, lobis catadromicis,
crenatis; rhachis et costae pubescentes et
squamatae. Folia fertilia ignota.

Stem 2.5 cm thick, hemiepiphytic; scales
linear, 15—25 x 0.8-1.2 mm, membranaceous,
golden brown, concolorous or witha slightly dark-
ened central stripe, appressed-ascending, margins
denticulate. Sterile leaves up to 1.3 m long; petiole
to 45 cm long, scaly, pubescent with 0.2 mm long,
unicellular, colorless hairs; lamina 2-pinnate-
pinnatifid, the adaxial surface glabrous, the ab-
axial surface lightly to moderately pubescent, the
hairs unicellular, less than 0.1 mm long, whitish,
subulate; pinnae up to 30 X 8 cm, narrowly trian-
gular with a long tapering apex, 3—4 times as long
as broad, the lowest becoming pinnatifid in the

Vol. 34, Art. 1

apical one-third, the transition zone between pin-
nate and pinnatifid marked by a narrow, straight,
decurrent, green wing parallel to the costa; pin-
nules catadromically arranged in the medial pin-
nae, lobed almost to the costule, inequilateral at
base, up to 6 X 2 cm, the proximal acroscopic
ones longer than the proximal basiscopic; pinnule
lobes catadromically arranged, the margins cre-
nate, the apices rounded; axes pubescent ab-
axially, the hairs 0.1—0.2 mm long, subulate, col-
orless, often difficult to see on the lamina, inter-
mixed on the costae with long, scraggly, subclath-
rate, denticulate scales; groove usually glabrous,
abruptly truncated by the raised ridge of the costal
groove, the junctures short-pubescent. Fertile
leaves unknown.

Polybotrya alata has been collected only
twice, both times in the mountains of Panama
(Map 14), but I suspect it also occurs in the moun-
tains of southern Costa Rica. Distinctive features
of this species are the narrowly triangular pinnae
with long attenuate apices and the pinnules that
are lobed almost to the costule, thereby making
the lamina 2-pinnate-pinnatifid (Fig. 42a). The
transition zone from the free pinnules to the pin-
natifid apex differs from all other species of
Polybotrya by the thin, basally decurrent, green
wing, which imparts an alate appearance to the
costa (Fig. 42c). The stem scales are also distine-
tive by their long length (15—25 x 0.8—1.2 mm)
and shiny, golden brown color.

The short, even pubescence on the abaxial
surface of the leaf separates P. alata from all other
Central American Polybotrya, except P. caudata,
which can also have pubescent leaves. The laminar
hairs of P. caudata, however, are longer (1—3mm)
than those of P. alata and are 4—10 celled. The
two species also differ sharply by their stem scales.
The only other Central American species with
which P. alata could be confused is P. osmun-
dacea, but that species is easily distinguished by
its anadromic pinnule arrangement and nonalate
costae. Furthermore, the costular groove of P.
alata is truncated by the ridges of the costa; in
contrast, P. osmundacea has uninterrupted, decur-
rent grooves (Fig. 47h).

Specimens examined: PANAMA. Panama: Cam-
po Tres, 3 mi NE of Altos de Pacora, Croat 22706 (MO,
L). Veraguas: 5 mi W of Santa Fe on road past Escuela
Agricola Alto Piedra on Pacific side of divide, 800—1200
m, Croat 2301/1 (MO).

November 1987 MONOGRAPH of POLYBOTRA

mm

FIGURE 42. Polybotrya alata Moran. a. sterile pinna; b. margin of stem scale: c. abaxial surface of
distal pinnules (note pubescence and decurrent pinnule bases); d. stem scale. a—d: Croat 22706 (MO).

93

94 ILLINOIS NATURAL HISTORY SURVEY

24. Polybotrya aequatoriana Moran, sp. nov.
(Fig. 43, Map 15).

Polybotrya aequatoriana Moran, sp. nov.
Type: Ecuador. Napo: Cordillera Oriental,
camino Baeza—Tena, 34 km al sur de Baeza,
2100 m, 30 de diciembre 1983, Moran 3512
(holotype: F!; isotypes: AAU!, COL!, GH!,
LPB!, MO!, NY!, Q!, QCA!, UC!).

Caulis hemiepiphyticus, 1.5—3.4(4.0) cm
diam.; squamis sordide aurantiaco-luteis, nitidis,
membranaceis, implexis, lanatis, in margine mi-
nute denticulatis, 1—2(3) * 12—20(25) mm; lamina
lanceolata, usque ad 1.40.7 m, 2-pinnata-
pinnatifida, modice vel sparse puberula subter,
pilis appressis, 0.1—0.3 m longis; pinnae longidel-
tatae, 25—35 X 13—18(20) cm; pinnulae catad-
romicae, acroscopice prolongatae, basiscopice
valde obliquae; lobi integri vel vade crenati;
rhachis et costae pubescentes, pilis rigidis albidis,
sulcis profundis pubescentibus. Pinnulae fertiles
lineares, vulgo 4—6 X 0.5—1.0 cm, segmentis op-
positis vel suboppositis circularibus vel oblongis,
3-5 x 1-2 mm.

Stem 1.5—3.5(4) cm thick, hemiepiphytic;
scales dingy orange yellow, mostly darkening
upon drying, shiny, membranous, tangled and
woollike, the margins minutely denticulate, 1—
2(3) X 12—20(25) mm. Sterile leaves up to 1.6 m
long; /amina lanceolate, to 1.4 x 0.7 m, 2-pinnate-
pinnatifid, moderately to lightly pubescent below,
the hairs fine, appressed, 0.1—0.3 mm long, grad-
ing into scales along the axes, often with a few
scattered resinous, circular glands (use at least
30), the margins somewhat thickened and
lighter colored, often becoming revolute upon dry-
ing; pinnae long-triangular, 25—35 x 13—18(20)
cm; pinnules arranged catadromically above the
base, prolonged acroscopically, reduced and very
oblique basiscopically; lobes entire or rarely shal-
lowly crenate; axes pubescent below with un-
evenly scattered, whitish, erect hairs 0.1—0.2 mm
long, scaly by linear, flexuous, darkened, often
appressed scales, these most abundant at the
junctures; grooves deep and distinct, filled with
hairs, these sometimes clavate, ca. 0.1 mm long,
reddish, the ridges well defined, stramineous, con-
tinuous with those of the costules but truncated
by the ridges of the rhachis. Fertile leaves bot-
ryoid; pinnules linear, mostly 4—6 X 0.5—1.0 cm;
sori arranged oppositely or suboppositely, round

Vol. 34, Art. 1

to oblong; sporangial stalks paraphysate; spore
size unknown.

Polybotrya aequatoriana occurs in the cloud
forests of Ecuador and Bolivia (Map 15), from
2100-2410 m, the highest known altitudinal range
of any species of Polybotrya. Polybotrya aequa-
toriana is named for Ecuador, where I collected
the type.

Polybotrya aequatoriana can be easily sepa-
rated from P. hickeyi and P. appressa by its stem
scales, which are thin, yellowish orange, and
membranaceous (Fig. 43b); see P. appressa for
further comparison with that species. Polybotrya
aequatoriana has highly reduced basiscopic lobes
on the sterile pinnules (Fig. 43c), and the fertile
pinnules are much narrower than those of other
congenerics.

Specimens examined: ECUADOR. Napo: Cor-
dillera Oriental, camino Baeza—Tena, 34 km al sur de
Baeza, 2100 m, Moran 3512 (AAU, COL, F, GH, LPB,
MO, NY, Q, QCA, UC), 3585 (F, Q, QCA); road
Baeza—Lago Agrio, ca. 114 km from Lago Agno, 1750
m, @llgaard et al. 35776 (AAU, QCA).

Bo.iviA. La Paz: Prov. Sud Yungas, Huancane,
7.5 km hacia el sud sobre el camino nuevo, 2410 m,
Beck 3108 (LPB); same locality, but at 6.5 km, 2280
m, Beck 3060 (LPB, NY).

25. Polybotrya appressa Moran, sp. nov. (Fig.
44, Map 17).

Polybotrya appressa Moran, sp. nov. TYPE:
Ecuador. Napo: Cordillera Oriental, camino
Baeza—Tena, 34 km al sur de Baeza, bosque
virgen, 2100 m, 24 de enero 1984, Moran
3586 (holotype: F!; isotypes: GH!, MO!,
NY!, Q!, QCA!, US!).

Caulis hemiepiphyticus, 1-2 cm diam.;
squamae concolores, obscure brunneae, integrae,
appressae, vulgo 1-2 X 0.1—0.2 cm; lamina ovata
vel late deltata, 2-pinnata-pinnatifida, superne
glabra, inferne pilosa, pilis disnersis, ca. 0.1 mm
longis; pinnae usque ad 37 X 18 cm; pinnulae pro-
longatae acroscopice, latis basiscopicis reductae;
lobis rotundatis vel plusminusve truncatis; rhachis
et costae aequaliter pubescentes, pilis usque ad
0.1 mm longis, sulcis glabris. Folia fertilia ignota.

Stem 1-2 cm thick, hemiepiphytic; scales
concolorous, dull brown, thick, opaque, entire,

November 1987 MONOGRAPH of POLYBOTRA 95

: yas!
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d
ie

i
% f
$ #
f i
f s
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eh rt Oh
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Ficure 43. Polybotrya aequatoriana Moran. a. sterile leaf; b. stem scales; c. costa and pinnule bases,
abaxial view (note only scattered hairs); d. rhachis, costa, pinnule bases, adaxial view, groove minutely
pubescent, thin. a—d: Moran 3512 (F).

96 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

FiGurE 44. Polybotrya appressa Moran. a. sterile leaf; b. stem scales; c. costa and pinnule bases,
abaxial view; d. costa and pinnule bases, adaxial view. a—d: Moran 3586 (F).

November 1987

appressed, mostly 1-2 x0.1—0.2 cm. Sterile
leaves up to 1.5 m long; lamina ovate to broadly
deltate, dark green above, slightly paler below,
2-pinnate-pinnatifid, up to 1.3 x 0.74 m, gla-
brous adaxially, abaxially with a few scattered, ap-
pressed hairs, these less than 0.1 mm long, inter-
grading with scales on the costae; pinnae to 37 X
18 cm, widest at the base, the apex long-tapering,
the acroscopic basal pinule usually prolonged
beyond the rest; pinnules arranged catadromically,
prolonged acroscopically and reduced basiscopi-
cally at the base, the proximal ones with acuminate
apices, the distal ones with falcate and truncate
apices; lobes rounded to somewhat squarish, entire
or rarely minutely serrate at the apex; axes evenly
pubescent abaxially, the hairs short, less than 0.1
mm long, the costal scales present or caducous
and absent, linear to lanceolate, appressed, light
brown, thin; grooves sparsely pubescent within,
not very deep or prominent, bordered by broad,
shallow ridges, not admitted to the groove of the
next lower order. Fertile leaf unknown.

Polybotrya appressa is known only from the
eastern Andes of Ecuador (Map 17), where it
grows in cloud forests at high elevations. This
new species is named for its distinctive, appressed
stem scales (Fig. 44b), which help separate it from
similar species, many of which have spreading,
membranous stem scales. Polybotrya caudata has
similiar dull, thick, opaque stem scales, but that
species does not grow at high elevations.

Polybotrya appressa differs from P. aequa-
toriana, a closely allied Ecuadorian species, by
its stem scales, groove architecture and pubes-
cence, and lamina shape. Polybotyra hickeyi is
also very closely related, especially by its short,
even, abaxial, costal pubescence, and the indis-
tinct adaxial grooves that are glabrous within; but
it differs primarily by its more finely cut lamina.
In addition, the lamina of P. appressa is dark
green, in contrast to the pale yellowish green
laminae of P. hickeyi and P. aequatoriana.

Specimens examined: ECuApDoR. Napo: Cor-
dillera Oriental, camino Baeza—Tena, 34 km al sur de
Baeza, Bosque virgen, 2100 m, 24 de enero 1984, Moran
3586 (F, GH, MO, NY, Q, QCA, US); Cordillera Orien-
tal, 5 km al sur de Cosanga, camino Baeza—Tena, 1000
m, 4 de enero 1984, Moran 3527 (Q, QCA); Cerro
Huacamayos, on road Baeza—Tena, ca. 34 km from
Baeza, Mllgaard et al. 53824 (AAU, QCA).

MONOGRAPH of POLYBOTRA 97

26. Polybotrya altescandens C. Chr. (Fig. 45,
Map 12).

Polybotrya altescandens C. Chr., Index
Filicum 7. 1905. nom. nov. for Acrostichum
chrysolepis Sodiro, non Fée 1869.

Acrostichum chrysolepis Sodiro, [Anal.
Univ. Quito (X1)77:561. 1894.] Crypt. Vasc.
Quit. 485. 1893. nom. illeg., non Fée 1869.
Type: Ecuador. Pichincha: selva tropical, de
Pilaton-Toachi, septiembre 1892, Sodiro s.n.
(holotype: P!).

Stem (1)2—3(4) cm thick, hemiepiphytic;
scales golden to yellowish, mostly concolorous,
shiny, tangled and forming a dense “wool” around
the stem, linear, 0.6—1.3 * 15—25(32) mm, base
cordate and darkened at the point of attachment.
Sterile leaves up to 2 m long; lamina to 1.65 m
long, lanceolate, to 2-pinnate-pinnatifid or rarely
3-pinnate, usually moderately pubescent, the hairs
fine, appressed, more or less cylindrical, multicel-
lular, 0.2—0.4 mm long, intergrading with scales
on the costa; pinnae evenly tapering to a truncate
base, the acroscopic side slightly prolonged, espe-
cially in the distal pinnae, 14—25(35) 3—10(17)
cm, soon becoming pinnatifid in the distal por-
tions; pinnules anadromous or subequal proxi-
mally, catadromous distally, entire to coarsely ser-
rate, the basal acroscopic one almost always con-
spicuously prolonged beyond the others; axes
pubescent abaxially, the hairs subulate, colorless,
0.1—0.2 mm long, unicellular, the scales few,
scattered, denticulate; grooves slightly pubescent
within, those of the costule decurrent into those
of the costa, costal groove gradually becoming
shallow and expanded before the juncture with the
rhachidial groove; veins in pinnate groups, in large
pinnae and pinnules the lowermost basiscopic
veinlet often springing directly from the costa or
costule, appearing as a single isolated veinlet be-
tween the main pinnate groups. Fertile leaves 3-
pinnate, pubescent by jointed hairs 0.5—1.0 mm
long, these grading into linear, tortuous scales;
sori obovate or oblong; sporangial stalks para-
physate; spores (54)59-69(72) microns long.
n=41,

This species has an altitudinal range of
(800)1200—2500 m. Most of the collections come
from the western cordillera of Ecuador, but two
apparently disjunct locations occur in Colombia

Vol. 34, Art. 1

ILLINOIS NATURAL HISTORY SURVEY

15 cm

cm

FiGure 45. Polybotrya altescandens C. Chr. a. stem scales; b. fertile leaf; c. fertile pinnae; d. sterile
pinnule (note isolated veinlet between the main pinnate groups); e. fertile pinnule (note obovate seg-
ments). a,b: Moran 3559 (CR). c,e: Rimbach 9] (F). d: Steyermark 528/]2 (F).

November 1987

and Peru (Map 12). I suspect that this species
occurs in many other valleys of the Colombian
and Peruvian Andes, and that lack of collecting
accounts for the apparent disjunctions. My field
experience in Ecuador showed this species to be
quite common in the western cordillera but absent
from the eastern cordillera.

Polybotrya altescandens can be immediately
recognized by its bright golden to yellowish scales
that thickly cover the stem. These scales are longer
and narrower than those of any other species of
Polybotrya (Fig. 45a). The lamina is less cut than
that of similar Andean species, and the pinnae
soon become pinnatifid in their distal portions, a
characteristic that further accentuates this less-
divided look. The sori are usually clavate or short-
oblong, in contrast to the more elongated sori of
other species (Fig. 45e). An unusual tendency of
the venation further distinguishes this species: in
large pinnae and pinnules the lowermost basi-
scopic vein migrates onto the costa or costule and
therefore appears as a single isolated veinlet be-
tween the main pinnate groups (Fig. 45d).

Specimens examined: COLOMBIA. Antioquia:
bosque bajo la cumbre cerca de Boquer6n, camino entre
Medellin y Palmitas, 2300-2500 m, Hodge 6592 (GH).

Ecuapor. Azuay: rich dense jungle between
Chacanceo and Rio Blanco, on road to Molleturo, be-
tween Rio Putucay and Rio Norcay, 1220 m, Steyermark
528/2 (F). Carchi: environs of Maldonado, 1450 m,
Madison et al. 485] (F). Chimborazo: mountains in
Sacramento area, Wiggins //073 (NY). Cotopaxi: road
between Quevedo and El Coraz6n, 6.4 km NW of El
Coraz6n, 67.5 km SE of Quevedo, Croat 55844 (MO).
Guayas: valley of Rio Chimbo, 800 m, Rimbach 9] (F,
US). Imababura: in the vicinity of the Rio Verge, ca.
5 km SW from the village of Mani, Rio Cachaco, 1300
m, Sperling & Bleiweiss 5034 (GH, Q, QCA).
Pichincha: tropical forest de Pilaton-Toachi, September
1892, Sodiro s.n. (P); road El Paraiso—Saguangal, 11
km from El Paraiso, @llgaard et al. 37702 (AAU),
37867 (AAU); road El Paraiso—Saguangal, 3 km from
El Paraiso, 1500 m, @llgaard et al. 37820 (AAU); selva
topical, Valle de Nanegal, Sodiro s.n. (P, US); Mindo,
Sydow 339 (US); road from Sto. Domingo de los Co-
lorados to Aloag, 2.5 km E of Cornejo Astorga, 1200
m, Moran 3544 (F, GH, Q) QCA), Tinalandia Resort,
ca. 25 km E of Sto. Domingo de los Colorados, N side
of Rio Toachi, 1000 m, Moran 3559 (CR, F, GH, NY,
Q, QCA); ca. 30 km WNW of Quito, 67 km on road
to Mindo, 2200 m, Moran 3564 (F, Q, QCA, US).
Province unknown: western cordillera, 800 m, forest
region, Rimbach 3/2 (US); without locality, Jameson
33 (P).

MONOGRAPH of POLYBOTRA 99

Peru. Huanuco: La Divisoria, ca. 25 km NE of
Tingo Maria, Moran 3688 (F, GH, MO, USM). Loreto:
Prov. Coronel Portillos, Dtto. Padre Abad., divisoria
cerca al Rio Chino, J. Schunke 10200 (MO).

27. Polybotrya gomezii Moran, sp. nov. (Fig.
46, Map 14).

Polybotrya gomezii Moran, sp. nov. TYPE:
Costa Rica. Alajuela: ca. 20 km N of San
Ramon, at the Univ. of San Ram6n’s Biolog-
ical Field Station, 1100 m, cloud forest, 17
July 1983, Moran 324] (holotype: CR!;
isotypes: F!, GH!, MO!, NY!, UC!, US!).

Caulis hemiepiphyticus, 1-2 cm diam.;
squamae appressae vel ascendentes, brunneae,
obscurae, opacae, anguste lanceolatae, vulgo 10—
20 x 1-2 mm, marginibus vulgo integris; lamina
usque ad 1.5 X0.75 m, ovata vel deltata, ad-
modum glabra, 2-pinnata-pinnatifida, coria-
cea, inferne anadromica, superne catadromica;
pinnae usque ad 45 X 20 cm, anguste deltatae,
apicibus longiacuminatis; pinnulae usque ad 12 X
2.5 cm, acroscopice leviter prolongata. Folia fer-
tilia usque ad 0.75 m, 3-pinnata; sporae (50)54—
60(64) micrometralae.

Stem 1-2 cm thick, hemiepiphytic; scales ap-
pressed to spreading, dull brown, opaque, thick-
ened in the middle, narrowly lanceolate, 10—20 x
1—2 mm, margins entire or with a few apical den-
ticulations. Sterile leaf up to 1.5 m long; lamina
up to 1.2 x 0.75 m, 2-pinnate-pinnatifid or very
rarely 3-pinnate, coriaceous, ovate-triangular,
nearly glabrous, usually anadromic below, becom-
ing catadromic in the distal one-third, the change
from anadromic to catadromic marked by a small,
reduced, basiscopic pinnule or segment; pinnae
up to 45 x 20 cm, narrowly triangular, the apex
long acuminate, the lower ones pinnatifid in the
apical third, giving a broadened appearance to the
pinnae; pinnules up to 12 * 2.5 cm, apex of prox-
imal ones acuminate to long acuminate, gradually
becoming curved and then truncate in the pin-
natifid apex, the base inequilateral with the acro-
scopic side slightly more developed, the margins
serrate to pinnatifid; axes usually glabrous abax-
ially or with unicellular, fine, whitish hairs, scaly,
the scales caducous, appressed, up to 2 mm long,
light brown, with a long narrow apex; grooves
mostly glabrous within or nearly so, sometimes

100 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

5 cm

FiGuRE 46. Polybotrya gomezii Moran. a. acroscopic basal pinnule, pinnae apex toward the left (note
catadromous venation); b. sterile leaf (note that the transition zone from anadromic to catadromic is
marked by reduced basiscopic pinnules); c. fertile leaf; d. stem scales; e. basal pinna. a,d: Moran 324]
(CR). b,e: Moran 244] (F). c: Lent 3537 (F).

November 1987

pubescent at the junctures, the hairs short, less
than 0.1 mm long and inconspicuous. Fertile
leaves up to 0.75 m long, 3-pinnate, coenosoric;
axes with scattered scales similar to those on the
sterile leaf, junctures pubescent, the hairs numer-
ous, short, less than 0.2 mm long; sporangial
stalks paraphysate; spores (50)54—60(64) microns
long.

This species is named in honor of Luis D.
Gomez P., pteridologist, formerly at the Museo
Nacional de Costa Rica, who has encouraged my
study of Polybotrya and extended much hospitality
during my thesis research in Costa Rica. Polybot-
rya gomezii is endemic to Costa Rica (Map 14),
where it occurs from sea level up to 1500 m,
reaching its greatest frequency and abundance in
cloud forests. Most of the specimens were col-
lected in the mountains around San José. I found
P. gomezii at five localities, always with P. al-
fredii; no hybrids were found.

This species can be distinguished from others
in Central America by its nearly glabrous, only
2-pinnate-pinnatifid leaves, and the nearly gla-
brous axes. The pinnae soon become pinnatifid in
the apical half, which also gives a more broad,
less finely cut appearance to the leaf (Fig. 46b).
The opaque, dull brown stem scales with entire
or sparsely denticulate margins (Fig. 46d) also
help separate this species from others found in
Costa Rica. The veinlets of the pinnules are always
arranged catadromically, even in pinnae having
the pinnules arranged anadromically. This order-
ing is unusual because the disposition of the
veinlets usually reflects the pinnule arrangement;
that is, if the pinnules are anadromic, the veinlets
are also anadromic.

Specimens examined: Costa Rica. Alajuela:
ca. 20 km N of San Ramon, at the Univ. of San Ramén’s
Biological Field Station, cloud forest, Moran 3241] (CR,
F, GH, MO, NY, UC, US); forest between Quebrada
Quicuyal and Quebrada Arrayanes, Cariblanco, Lent
3537 (F). Cartago: near Alto Patillos, Moran 2441 (CR,
F, GH, MO, NY, US); cloud forest near the entrance
to Parque Nacional Tapanti, 1270 m, Moran 3338 (CR,
F, GH, NY, US); 3 km SE of Tapanti, lower slopes of
Alto Patillos, Lent 1084 (F); La Hondura, 1400 m, Val-
erio s.n., (CR); Navarro, 1400 m, Wercklé s.n. (GH,
P). Heredia: about 10 km on the road towards Virgen
de Socorro, 1000 m, Moran 3/60 (CR). Puntarenas:
Monteverde Cloud Forest Reserve, 1560 m, Fiedler &
Koptur 51 (UC). San José: Parque Nacional Braulio
Carrillo, ca. 1 km along road from entrance, Moran
3271 (CR).

MONOGRAPH Of POLYBOTRA 101

28. Polybotrya osmundacea Willd. (Figs. 47 &
48, Map 18).

Polybotrya osmundacea Willd., Species Pl.,
ed. 4, 5:99. 1810. Type: Venezuela.
Monagas: Caripe, Humboldt 459a (fertile),
459b (sterile), Herb. Willd. 19507-1, 19507-
2 (holotype: B; photos F!, GH!).

Acrostichum osmundaceum (Willd.) Hooker,
Species Filicum 5:246. 1864.

Polybotrya aristeguietae Brade, Bradea 1:19,
tab. 1. 1969. Type: Venezuela. Miranda:
Santa Teresa-Altogracia de Orituco, June
1953, Aristeguieta 1780 (holotype: VEN!).

Polybotrya vareschii Brade, Bradea 1:20, tab.
2. 1969. Type: Venezuela. Aragua: Rancho
Grande, Dependiente Norte, selva nublada,
800 m, Vareschi & Gessner 1875 (holotype:
VEN!).

Stem 1—2.5(4) cm thick, hemiepiphytic;
scales extremely variable, with plants from
Amazonia tending to have thick, opaque, dark
brown, subentire, somewhat squarrose scales and
plants from elsewhere tending to have shiny, as-
cending, spreading, bicolorous scales, often with
a dark central stripe and lighter borders, margins
highly erose or, less commonly, denticulate,
mostly 1.0—2.2 x 0.8—1.9(2.3) cm. Sterile leaves
up to 1.8 m long; /amina deltate, ovate, or lanceo-
late, to 3-pinnate-pinnatifid, chartaceous, rarely
with scattered, round, sessile, resinous glands on
the abaxial surface, the margins glabrous; pinnae
narrowly deltate, equilateral, (7)13—18 pairs; pin-
nules arranged anadromically or rarely catadrom-
ically (Peru), up to 14 x 4 cm, the largest deeply
cut at the base, apex acuminate; basal acroscopic
tertiary segments ovate, lanceolate or rhombic,
margins entire, crenate or lobed, the apex often
serrulate; axes glabrous or pubescent abaxially,
the hairs whitish, 0.2—-0.5 mm long, the scales
absent or few and inconspicuous; grooves decur-
rent into those of the next lower order, usually
filled with short, less than 0.7 mm long, reddish
or brownish hairs. Fertile leaves to 3-pinnate-
pinnatifid, deltate, coenosoric; sporangial stalks
paraphysate; spores (50)54—62(66) microns long.

Other illustrations: See original descriptions
of P. aristeguietae and P. vareschii; Humboldt et
al., Nov. Gen. Sp. Pl., tab. 2. 1815; Proctor,
Ferns of Jamaica, fig. 107. 1985.

102 ILLINOIS NATURAL HisTORY SURVEY Vol. 34, Art. 1

_15 cm

tem

SS Sy)
AML

Ficure 47. Polybotrya osmundacea Willd. a. sterile leaf (Costa Rica); b. stem scale, dark, opaque
type (Amazon basin of Ecuador); c. stem scales, thin, transparent type (Andes of Colombia); d. sterile
pinnule (northern Venezuela); e. sterile pinnule, same scale as d (Costa Rica); f. basal pinnae of fertile
leaf, same scale and plant as a (Costa Rica); g. two basal tertiary segments (Colombia); h. rhachis-costa
juncture; note decurrent axial grooves (Ecuador). a,f,e: Moran 2/67 (F). b: Moran 3547 (F). c.g: Killip
& Smith 15341 (US). d: Vareschi & Gessner 1875 (VEN). h: Moran 3592 (F).

November 1987

Polybotrya osmundacea, the type species of

the genus, is one of the most widely distributed
species of Polybotrya (Map 18). It occurs in wet,
shaded forests from sea level to 2100 m. The
specific epithet refers to the fancied resemblance
of the fertile leaves to the fertile apex of Osmunda
regalis (Willdenow 1810).

This species varies tremendously in scales,
cutting, and pubescence. In plants from
Amazonia, the scales are usually thick, dark,
Opaque, more or less squarrose, and not thickly

MONOGRAPH of POLYBOTRA 103

investing the stem so that the yellowish stem
aerophores are easily visible in fresh material (Fig.
47b). Plants from other parts of the range may be
similar, but more often the scales are thin, lighter
colored, transparent, often bicolorous, and more
thickly investing the stem (Fig. 47c). The stem
scales, however, do not correlate with any other
characteristics.

Laminar cutting is another variable charac-
teristic. The type specimens of P. aristeguietae
and P. vareschii, both collected in northern Ven-

“

FiGurE 48. Geographical variation in the pinnule cutting of Polybotrya osmundacea Willd. The elon-
gated side of the pinnule is always acroscopic. Clockwise, starting from Bolivia: Buchtien 298 (UC);
Killip & Smith 2399] (F); Moran 3642 (F); Moran 3618 (PORT); Lellinger & de la Sota 213 (COL);
Moran 2167 (CR); H. Smith 1050 (US); Vareschi & Pannier 1686 (US); Killip & Lasser 37756 (US);
Murillo 2580 (F); Broadway 5589 (F); Steyermark 107148 (MO); Granville 3865 (CAY); Berg et al.

P18138 (NY).

104 ILLINOIS NATURAL HISTORY SURVEY

ezuela, scarcely differ from each other in cutting.
These two specimens, however, seem to differ
from P. osmundacea elsewhere in its range by
their less-cut leaves (Fig. 47d; Fig. 48, pinnules
from northern Venezuela). From the greater
number of specimens available to me, it appears
that these extremes of cutting are connected by
intermediates and that no other characters correlate
with these extremes. Accordingly, I have placed
P. aristeguietae and P. vareschii in synonymy
with P. osmundacea.

Specimens examined: GUATEMALA. Izabal: vi-
cinity of Quirigua, Standley 24195 (NY, US).

HONDURAS. Proy. unknown: near Lake Yojoa,
Steeves & Ray 504 (GH, US); near Cockscomb Mts.,
Schipp 8101 (US); Maya Mounds, Schipp 8108 (F).

NICARAGUA. Comarca del Cabo: 40-50 km SW
of Waspan, Atwood 37/1 (VT). Zelaya: Cerro la
Pimienta, no. 1, summit area, Pipoly 5/46 (CR); costado
S del Cerro La Pimienta, Grijalva 300 (CR).

Costa Rica. Cartago: forests of Las Vueltas,
Tucurrique, Tonduz 13337 (US), 18879 (US); forests of
Tuis, Pittier 12416 (CR, US); Aragon, Pittier 9076 (Z);
Chitaria, Valerio 328 (US); Chitaria, forest near old
jailhouse, Moran 2167 (CR, F); Turrialba, Aragon, Ton-
duz 9006 (US), 9016 (US); vicinity of Pejivalle, Skutch
4637 (CR, US); Peralta, Lankester s.n. (US); vicinity
of Pejivalle, Standley & Valerio 47127 (US); Turmialba,
Instituto Interamericana, Ganaderia, Croat 690 (MO),
738 (MO); Florecia, Turrialba, Jiménez 3299 (F), 3304
(CR, F); Reventazon below Turmialba, Hatch 90 (F);
Turnalba, Pittier 9016 (Z); 3 km W of Turrialba, Mickel
2624 (LP, NY); Turrialba, near the Interamerican Inst.,
Scamman 7152 (GH), 7687 (GH); Turrialba, Lent 299
(GH); valley of Rio Reventaz6n, 3 km SE of Turrialba,
Holm & Iltis 200 (MICH). Cocos Island: Stewart 241
(US); Fisher s.n. (US); Klawe 1474 (US), 1504 (US).
Limon: SW of Siquirres, on road to Turrialba between
Moravia and Guayacan | km of Guayacan, Lellinger &
White 1436 (F, US); near banana plantation and Pandora,
Rio Estrella, Rossbach 3628 (GH); Los Diamantes,
USDA Rubber Plant Station, Scamman 5987 (GH), 7153
(GH); hills of Atlantic slope, Tonduz 14568 (P); vicinity
of Guapiles, Standiey 37096 (US). Puntarenas: Coto,
at sea level, Valerio 333 (F); San Vito, de la Sota 518]
(LP); Finca Loma Linda 1 mi SW of Camas Gordas,
Croat 22260 (MO); road to Puerto Jiménez, Osa, 40
km W of I.A. route 2, Gomez 19489 (CR, UC); Osa
Peninsula, 15 km S of Tropical Science Center field
station and ca. 20 km S of Rinc6én de Osa, Mickel 2803
(NY), 28/7 (NY); Finca Las Cruces, San Vito de Java,
Burch 4613 (MO, NY); ca. 10 mi. SE of Rincén de Osa
along road to Pacific, Evans & Bowers 2792 (MO). San
José: vicinity of El General, Skutch 30/8 (F, MO); San
Isidro de El General, Scamman 5984 (GH).

PANAMA. Chiriqui: 10—11 miles W of Puerto Ar-
muelles in vicinity of San Bartolo Limite, Croat 2/987

Vol. 34, Art. 1

(MO); vicinity of Gualaca ca. 10.7 miles from Planes
de Horito, La Fortuna on road to dam site, Antonio
5133 (MO, UC); 4.5—5.0 km N of dam over Fortuna
Lake, Croat & Grayum 60022 (MO); Fortuna Dam area,
Quabrada Bonito to N of reservoir, Churchill 5776
(MO). Darién: vicinity of airstrip at Cana gold mine,
Croat 38047 (MO). Panama: Cerro Jefe, near summit,
Croat 22682 (MO).

CuBA. Oriente: Sierra Maestra on ascent from Rio
Yara, Ekman 14210 (US); Bahia de Taco, on the ndge
below Santa Maria and Rio Jiguani on the path to the
“Tberia” mines, Ekman 3769 (NY); Trail Navas to Camp
Buena Vista, Shafer 4458 (NY); La Perla, Maurel &
Maurel 3816 (NY); La Perla, Shafer 8889 (NY); near
Monte Verde, Wright 786 (BM, F, GH, MO, NY, PH,
UC, US); Eggers 5324 (F); Finca Guadalupe, Placetas,
Las Villas, Acuna 17531 (US); Sevilla Estate, near San-
tiago, trail from Magdalena to Sierra Maestra, Taylor
447 (NY); Loma del Gato and vicinity, Sierra Maestra,
Hioram & Clement 6424 (GH, US); La Prenda, Hioram
2495 (COL, P, US, UC); Santiago, Loma del Gato-
Cobre, Clement 725 (US).

Jamaica. Clarendon: “Second Breakfast Spring”
below Tweedside, Underwood 1612 (NY). Portland:
Dollwood, Watt 160 (GH, P, US), 7276 (BM); Mabess
River, below Vinegar Hill, Underwood 1323 (NY); trail
from Vinegar Hill to Mabess River, Underwood 1246
(NY), 1250 (NY); Mabess, Fisher 132 (NY); Blue Mts.,
Stony River to Macungo River, Morely & Whitefoord
655 (BM), 675 (BM); Blue Mts., Stony River Base
Camp, Morely & Whitefoord 695 (BM, MO); John Crow
Mts., E slope, 1.5 mi SW of Ecclesdown, Wilson &
Webster 549 (MICH); above Moore Town, Clute 259
(NY, US); valley of the Rio Grande, 8 air miles S of
Port Antonio, Gastony 43 (GH); vicinity of Thomsons
Gap, Maxon & Killip 756 (F, GH, NY, US); Spur of
John Crow Mts. opposite Mill Bank, Maxca 9366 (GH,
NY, PH); vicinity of Mill Bank, Maxon & Killip 148
(F, GH, NY, US); along trail to waterfall N of Hardwar
Gap, Proctor 16506 (MO), 22269 (GH). St. Andrew:
Blue Mts. near Cinchona, 15 km from Kingston, Fisher
s.n. (P); Catherine Peak, Faull 12583 (GH). St. Ann:
Blue Mts., Trafalgar, Perkins 1163 (GH). St.
Catharine: vicinity of Hollymount, Mount Diablo,
Maxon 2293 (NY, US). St. Thomas:Mansfield and ad-
joining properties, near Bath, Maxon 2470 (NY, US);
Manchester Blue Mt., Day s.n. (NY); forested ndge E
of Cuna Cuna Gap, Maxon 9464 (NY, US); Bath, Gilbert
s.n. (GH); upper southern slopes and summit of Mac-
casucker Bump, Maxon 9522 (GH, NY, PH); SE slopes
of Stone Hole Bump, Maxon 8978 (GH, NY, PH); Com
Puss Gap and vicinity, trail W over Blue Mts., Wilson
& Murray 564 (BM, GH, MICH); Corn Puss Gap, Proc-
tor 3969 (PH, US).

Harti. Massif du Nord, Chavary, Ekman 4734
(US); Camp Perrin, Ekman 5214 (US).

GRENADA. Grand Etang, Beard 1252 (UC, US);
no locality, Fraser s.n. (P); in Mirabeau Mts., Broadway

——

November 1987

2520 (Z); St. Georges, Azimas, 1896, Broadway s.n.
(NY); without locality, Sherring 13 (BM).

TRINIDAD. Pass, Arima Valley, Fleming & Fleming
52 (NY); no locality, Fendler 69 (BM, F, GH, MICH,
MO, NY, P, PH, UC); Hart 229 (P); Mount Tocuche,
Britton et al. 1267 (GH, NY, US); Tacarigua Ward, El
Tocuche, Walker T10995 (BM); Morne Bleu, Britton et
al. 2276 (GH, NY, US); St. George, Blanchisseuse,
Las Lapas trace, Barnard et al. 41] (BM, MO); Blan-
chisseuse saddle, Richardson 2037 (US); Blanchisseuse
road, 11-mile post, Broadway 6902 (F, MO, Z); Blan-
chisseuse road, 10-mile post, Broadway 5589 (F, MO);
Heights of Aripo, Broadway 9948 (F, NY, US), 9950
(F, GH, NY, US), 9957 (GH, NY, US); Las Lapas
road, Broadway 6459 (BM); Arima—Blanchisseuse
road, 13-mile post, Jermy 11195 (BM), 9-10-mile post,
Jermy 2368 (BM); Morne Bleu ridge, Jermy 2846 (BM);
Arima—Blanchisseuse road, 10-mile post, Fay 373
(BM); Maracas Valley, Las Cuevas trail, Fay 345 (BM);
10.5 mi N of Arima, Crosby 76 (MICH).

FRENCH GUIANA. Sommet tabulaire, zone centrale,
about 40 km SE of Saul, Granville 3865 (CAY, Z).

GUYANA. Region of Mt. Raywa, Jenman s.n.
(NY).

VENEZUELA. Anzoategui: Dtto. Bolivar, Fila El
Gacharo, ridge of the fila above Los Chorros and El
Cielo, Serrania de Turumiquire, Davidse & Gonzdlez
19444 (MO). Apure: Reserva Forestal San Camilo, SW
of caserio San Camilo (El Nula), Steyermark et al.
101552 (US, VEN). Aragua: selva nublada de Rancho
Grande, Dependiente Norte, Vareschi & Gessner 1875
(VEN); Parque Nacional, Dos Ritos, Killip & Lasser
37756 (US, VEN). Barinas: Dtto. Bolivar, along road
from Barinitas to Mérida, near one land bridge at San
Isidro, 30 km NW of Barinitas, Moran 3718 (PORT,
VEN); Dtto. Bolivar, San Isidro, ca. 5 mi NW of La
Soledad along Barinas—Sto. Domingo road, A.R. Smith
et al. 1388 (PORT, UC, Z). Distrito Federal: virgin
wet forest on slopes along old road between “Por-
tachuelo” and “Penita” (Petaquire) and Carayaca, be-
tween Colonia Tovar—Junquito road and Hacienda El
Lim6n, 6-8 mi below junction of Junquito—Colonia
Tovar road, Steyermark & Nevling 95930 (GH, VEN).
Falc6n: Cerro Azul, Wingfield 6914 (VEN). Falcén/
Lara: Cerro Socopa, Liesner et al. 8359 (MO, VEN).
Lara/Yaracuy: Dtto. Urdaneta y Bolivar, la fila Azul
y Hda. El Jaguar, Ortega & Smith 2387 (PORT); Sierra
de Aroa, 10-13 mi NW of Urachiche (Edo. Yaracuy)
along dirt road leading NW from Urachiche to Duaca
(Edo. Lara), A.R. Smith et al. 1347 (UC, PORT, Z).
Miranda: Santa Teresa—Altogracia de Orituco, Aris-
teguieta 1780 (VEN); Cerros del Bachiller, near E end
of virgin evergreen forest, above Quebrada Corozal, S
of Santa Cruz, 10 km (by air) W of Cupira, Steyermark
& Davidse 116499 (MO, UC, VEN). Monagas: Caripe,
Humboldt 459a,b (P, photo F, GH); Cerro de Gaucharo,
of Guacharo, Steyermark 6201/5 (F). Nueva Esparta:
Cerro Copey, Sugden 115] (UC). Territorio Federal

MONOGRAPH Of POLYBOTRA 105

Amazonas: Sierra Parima, vecinidades de Simarawochi,
Rio Matacuni, a unos 6—7 km al oeste de la frontera
Venezolana—Brasilera, Steyermark 107148 (MO, NY,
VEN); Depto. Rio Negro, 0-2 km E of Cerro La Neblina
Base Camp on Rio Mawarinuma, Liesner 16/33 (MO,
UC); environs of Neblina Base Camp, Plowman &
Thomas 13678 (F, UC). Sucre: Peninsula de Paria, Dtto.
Marino, camino Mundo Nuevo—Manacal, 18-20 km N
de Irapa, Dumont et al. 7439 (NY, VEN); alrededore
de Manacal, Murillo 2580 (F, NY, VEN); Peninsula de
Paria, Cerro de Humo, NE de Irapa, Steyermark 94952
(F, GH, VEN); Peninsula de Paria, arriba de Mundo
Nuevo, oeste de Cerro de Humo, Steyermark & Rabe
96145 (GH, VEN); Peninsula de Paria, vicinity of Ma-
nacal 15 km (by air) NW of Irapa, Steyermark & Liesner
120634 (MO, UC, VEN); Peninsula de Paria, arriba de
Mundo Nuevo, oeste de Cerro de Humo, Steyermark &
Rabe 71756 (VEN). Tachira: Dtto. Uribante, along road
from La Siberia to entrance to Las Cuevas Represa, van
der Werff & Gonzalez 5202 (MO, UC). Yaracuy: Dtto.
Bruzual, selva siempreverde, Montana de Maria
Lionoza, Quebrada Quibayo, desde abajo hasta casi la
cumbre, Steyermark et al. 125039 (UC, VEN); Cerro
“Chimborazo,” Sierra de Aroa, Vareschi & Pannier
2715 (US, VEN); en la selva que cubre la fila “La
Enjalma” al sur de Chivacoa, Vareschi & Pannier 2660
(VEN). Zulia: Parija, Vareschi 3147 (VEN); Dist. Mara,
NW wooded slopes of Cerro Negro, 5.5 km SW of
Rancho 505, S of Rio Guasare, Steyermark et al. 122814
(MO, VEN); Dtto. Bolivar, Cuenca del Embalse Burro
Negro (Pueblo Viejo), sector entre Quiros—El Pensado
y el pie de Cerro Socopo, en el area aprox. 10 km en
linea recta al este de Churugauarita, Bunting 9516
(VEN); 15 km de El Vigia, carretera Panamericana,
Vareschi & Pannier 1686 (US).

CoLomBIA. Antioquia: Rio Leon, Bendix site,
Cain 74 (MICH); carretera al mar cerca de Villa Arteaga,
Gutiérrez & Barkley 170109 (GH); Municip. Anori, Pro-
videncia, Soejarto 2805 (COL). Boyaca: Muzo, Lindsay
262 (BM). Cauca: Agua Clara, along hwy from Buena-
venturea to Cali, Killip & Cuatrecasas 38902 (F, US);
Costa del Pacffico, Rio Micay, brazo Noanamito, orilla
derecha, El Chachajo, Cuatrecasas 14246 (US). Chocé:
0.5—2.5 km N of the INDERENA Camp on Rio Truando
near Caserfo La Teresita, Lellinger & de la Sota 553
(COL, US); NW side of Alto del Buey, Lellinger & de
la Sota 213 (COL, US), 250 (COL, LP, US); trail from
Rio Mecana to Alto de Mecana, Gentry & Juncosa41021
(MO, UC). Magdalena: Sierra Nevada de Santa Marta,
region del Campano, Barkley & Gutiérrez V. 1897
(MICH); Santa Marta, near Las Partidas, 3500 ft, H.H.
Smith 1050 (B, F, NY, MICH, MO, PH, US, VT);
Forest Boca Toma, El Recuerdo, 2500 ft, Bennett 23
(F); Santa Marta Mts., trail beyond falls, El Recuerdo,
Niemeyer 44 (US). Meta: Villavicencio, Alston 7641
(BM). Santander: Barbosa, Henri-Stanislas 1710 (US),
Mesa de los Santos, Killip & Smith 15341 (COL, GH,
NY, US); between Lebrija and San Vicente, Alston 7341

106 ILLINOIS NATURAL HISTORY SURVEY

(BM). Prov. unknown: Municip. de Marsella, Vereda
La Nona, Finca Palermo, cerca al caserio Caracas, Cor-
dillera Central, vertiente occidental, /drobo et al. 10116
(COL).

EcuApor. Los Rios: Rio Palenque Biological Sta-
tion, km marker 56 N of Quevedo, Moran 3600 (Q,
QCA). Napo: 27 km SE de Coca, alrededor de pozo de
petroleo Auca 4, Moran 3618 (PORT, Q, QCA); 12 km
SW of Coca, por el camino se llama “Los Zorros,”
Moran 3612 (Q, QCA); 73 km NE de Baeza, propiedad
de Inecel, “Cascada de San Rafael,” Moran 3592 (F,
MO, Q, QCA); Puerto Francisco de Orellana (Coca),
17 km SW of the town at road along Rio Napo (Los
Zorros), Balslev & Madsen 10649 (AAU, Q, QCA);
Rio Napo, Panachocha (Oasis), Harling et al. 7535 (F,
GH); Anangu, Parque Nacional Yasuni, SEF project
area, Mllgaard et al. 38845 (AAU, Q, QCA), 38894
(AAU, Q, QCA), 39086 (AAU, Q, QCA). Pastaza:
Lorocachi, zone oeste del campamento militar a 3 km
del Rio Curaray, Jaramillo et al. 30783 (AAU, Q,
QCA). Pichincha: Pululahua, Sodiro s.n. (US); Chim-
borazo, Spruce 5685 (P); Los Colorados, Sodiro s.n.
(P); 15 km E of Sto. Domingo de los Colorados, road
behind Brasilia a Toachi, along Rio Toachi, Moran 3547
(F, Q, QCA). Santiago-Zamora: Cordillera Cutuct,
ridge just S and W of Rio Itzintza, Camp 1298 (NY),
1359 (NY). Tungurahua: Banos-Jivaria de Pintuc,
Stiibel 875 (B). Prov. unknown: Junganza, Crespi s.n.
(US); San Miguel, Sodiro 8] (UC).

PERU. Huanuco: SW slope of the Rio Llulla Pichis
watershed, on the ascent of Cerros del Sira, Camp 3
(Laguna), Dudley 13005 (GH); Tingo Maria, Allard
21609 (US), 21997 (US). Junin: Pichis Trail, Yapas,
Killip & Smith 25452 (NY, US); Chanchamayo Valley,
C. Schunke 164 (F), 705 (F), 1341 (F), 1395 (F), 1451
(F); E of Quimiri bridge, near La Merced, Killip &
Smith 2399] (F, NY, US). Loreto: Prov. Maynas, Peter
Jensen’s Explorama Lodge, 50 mi downriver on the
Amazon at Yonamono Ck., Moran 3642 (AMAZ, F,
USM); Altura Tuta Pishco on Rio Napo, Croat 20287
(MO); Gamitanacocha, Rio Mazan, J. Schunke 275 (F,
GH, NY, UC, US), 380 (F, GH, UC, US); Pumayacu,
between Balsapuerto and Moyobamba, Klug 3208 (F,
GH, MO, US); Veradera de Mazan, Croat 20786 (MO);
17 km SW of Iquitos, Croat 18476 (MO); Rio Napo
near Entrada de Isla Inayuga, Croat 20543 (MO). Madre
de Dios: Prov. Manu, Vargas 1/7743 (GH). San Martin:
Camino a Pushurumbo, 7—8 km E del puente de Palo
Blanco, Mariscal Caceres, Tocache Nuevo, J. Schunke
5785 (COL, NY, US). Ucayali: Rio Aguaytia above
mouth of Quebrada Yurac—Yacu, Croat 2085] (MO).

BouiviA. La Paz: Regi6n de Mapiri, Buchtien 298
(NY, UC, US); Mapiri, San Carlos, Buchtien 1066 (LP);
Yungas, Mururata, 1839, Pentland s.n. (P).

BRAziL. Amazonas: Manaus—Caracarai road, km.
148, Berg et al. P18138 (F, NY, VEN). Para: Serra
dos Carajas, AMAZ Camp Azul, Sperling et al. 5915
(GH, NY).

Vol. 34, Art. 1

29. Polybotrya cyathifolia Fée (Fig. 49, Map
19).

Polybotrya cyathifolia Fée, Mém. Fam.
Foug. 6. (Hist. Acrost.) tab. 2. 1866. TYPE:
Guadeloupe. “near the house of Mr. Bovie,”
L’Herminier s.n. (holotype: P!; other L’Her-
minier specimens that are probably types are
at GH!, L!, fragment NY!).

Stem 1-2 cm thick, hemiepiphytic; scales
dull, light reddish brown, concolorous, spreading,
the margins subentire to denticulate. Sterile leaves
up to 1.5 mlong; lamina to 3-pinnate, ovate to lan-
ceolate, up to 1.2 x 0.8 m, membranaceous, al-
most always with round, sessile, reddish, punctate
glands; pinnae up to 40 x 23 cm, narrowly deltate,
pinnatifid to the very apex, stalked 5-10 mm;
pinnules catadromous to anadromous but mostly
subequal, narrowly deltate, stalked 1-3 mm, the
base subequilateral, slightly prolonged acroscopi-
cally but not oblique or cuneate on the basiscopic
side; tertiary segments serrate to entire, the proxi-
mal ones of the largest pinnules oblong, slightly
gibbous at the base on both the basiscopic and
acroscopic sides; axes abaxially sparsely to mod-
erately pilose with hairs similar to those above;
grooves densely packed with multicellular reddish
or tawny hairs, these 0.3—0.8 mm long and con-
spicuously exserted from the groove, evident to
the unaided eye, especially at the pinnae junctures.
Fertile leaves 3-pinnate-pinnatifid, coenosoric;
sporangial stalks paraphysate; spores (47)50—
59(63) microns long.

Other illustrations: See original description;
Plumier, Tr. Foug. Amér. tab. 32. 1705.

Polybotrya cyathifolia is endemic to the Car-
ribean islands of Guadeloupe and Martinique (Map
19). Few collections have been made of this fern,
but the original collection by L’Herminier is well
prepared and has many duplicates.

Polybotrya cyathifolia is very much like P.
osmundacea, but it differs by the shape of its
tertiary segments, the resinous punctate glands,
and by the long-pilose hairs on the axes. The best
way to distinguish P. cyathifolia is by the costal
groove hairs that copiously fill and protrude from
the groove, especially at the junctures (Fig. 49c).
Polybotrya osmundacea, on the other hand, usu-
ally has very short hairs (0.1—0.2 mm long) in
the groove. An important tendency in lamina cut-
ting is for the basal tertiary segments to be slightly

November 1987 MONOGRAPH of POLYBOTRA 107

Era ASAD TA RIT Aad ey ed BAe ESL

5 cm

FiGurE 49. Polybotrya cyathifolia Fée. a. stem scales; b. basal pinna; c. rhachis-costa-costule
junctures; d. fertile pinnule; e~g. sterile pinnules; h. abaxial view of lamina showing resinous glands.

a,c,h: L’Herminier s.n. (P). b,g: L’Herminier s.n. (GH). d: collector unknown (GH). e,f: collector
unknown (B).

108 ILLINOIS NATURAL HISTORY SURVEY

gibbous at its base on both sides (Fig. 49c). This
gives the pinnules a characteristic appearance, al-
though it is somewhat difficult to describe.
Polybotrya osmundacea rarely has these lobules
and, if so, they are developed only on the acro-
scopic side. Most specimens of P. cyathifolia have
the abaxial surface of the lamina covered with
punctate, resinous glands (Fig. 49h); P. osmun-
dacea rarely has such glands.

Specimens examined: GUADELOUPE. “near the
house of Mr. Bovie,” L’Herminier s.n. (P, GH, L, frag-
ment NY).

MARTINIQUE. Duss 1503 (NY, US), 3897 (NY),
4719 (F, US); Hahn s.n. (P); Belanger s.n. (F).

30. Polybotrya latisquamosa Moran, sp. nov.
(Fig. 50, Map 19).

Polybotrya latisquamosa Moran, sp. nov.
Type: Colombia. Meta: Cordillera La Maca-
rena (extremo nordeste), macizo Renjifo, al-
rededores, alt. 1300—1900 m, 6—20 de enero
1951, Idrobo & Schultes 1106 (holotype: US!;
isotype: COL?).

Caulis 2 cm diam.; squamae castaneae,
lineares vel anguste lanceolatae, 7-12 X 0.8-2.0
mm; petiolus ad basim squamatus, squamis cas-
taneis, longioribus, 12-16 X 5—6 mm, deltatis vel
late ovatis, marginibus pallidis erosis; lamina late
deltata, 3-pinnata-pinnatifida, glabra in super-
ficiebus utrinque, usque ad 13.5 X 5 cm, petiolulis
usque ad 8 cm; segmenta tertiaria valde ascenden-
tia basiscopice. Folia fertilia ignota.

Stem 2 cm thick, hemiepiphytic; scales cas-
taneous, linear to narrowly lanceolate, ascending-
spreading, darker and opaque in the center with
lighter borders, margins erose, 7-12 X 0.8-2.0
mm. Sterile leaves up to 1.5 m (?); petiole scaly
at the base with scales similar to those of the stem
but much larger, 12-16 x 5—6 mm, deltate to
broadly ovate; lamina broadly deltate (?), 3-pinnate-
pinnatifid, glabrous on both surfaces; pinnae ca.
10-12 free pairs (?), alternate, the largest 45 x 26
cm; pinnules arranged anadromically, the proxi-
mal ones with stalks up to 8 mm long, the base
strongly prolonged acroscopically, basiscopically
shortened and strongly ascending, up to 13.5 x 5
cm; tertiary segments up to 3.5 X 1.3 cm, lanceo-
late, cuneate at the base, pinnatifid, 3—5 free be-
low the pinnatifid apex, the basiscopic side strong-

Vol. 34, Art. 1

ly oblique, often with the first proximal 1—3 lobes
completely suppressed; costae glabrous abaxially
or with a few scattered, inconspicuous, less than
0.1 mm long, whitish, subulate hairs; grooves
filled with reddish, inconspicuous hairs less than
0.1 mm long. Fertile leaf unknown.

The type location is a remote area in south-
central Colombia, the Cordillera de Macarena,
separated from the main Andean chain (Map 19).
Since the holotype consists of only a portion of
the stem and several incomplete pinnae, additional
collections of this species would add greatly to
the incomplete description.

Polybotrya latisquamosa resembles P. osmun-
dacea because of its large decompound leaves with
anadromically arranged pinnules but differs by its
exceedingly large, broad, castaneous scales at the
base of the petiole (Fig. 50d); thus the specific
epithet. The stem scales differ from those of P.
osmundacea by their deep castaneous color. The
holotype is larger and more finely cut than the
average P. osmundacea specimen, and the basi-
scopic sides of the pinnules and tertiary segments
are extremely reduced and oblique-ascending (Fig.
50a). Finally, the major axes of P. latisquamosa
are glabrous (Fig. 50e), unlike those of P. osmun-
dacea, which are often pubescent.

31. Polybotrya sessilisora Moran, sp. nov. (Fig.
51, Map 20).

Polybotrya sessilisora Moran. TYPE: Colom-
bia. Vaupés: Rio Vaupés, Mitt y alrededores,
250 m, 8 September 1951, Schultes & Ca-
brera 13963 (holotype: US!; isotypes: COL!,
GH!).

Caulis hemiepiphyticus, 5-10 mm diam.;
Squamae appressae, ascendentes, lineares, 8—12
mm longae, atrocastaneae, marginibus pallidis et
valde vehementer denticulatis; lamina usque ad
3-pinnata-pinnatifida, glabra, coriacea, pallide
viridis; pinnae anguste deltatae vulgo 17-36 X 7-
16 cm; pinnulae vulgo 3.5-10 X 1.2-3.0 cm,
anadromicae, apicibus persaepe obtusis; rhachis
et costae sparsim pubescentes, squamis fuscatis,
appressis, flexuosis; venae conspicuae et pro-
minulae. Folia fertilia usque ad 3-pinnata;
lamina anguste alata viridis, marginibus leviter
incrassatis; sori ca. | mm longi, discreti, circu-
lares.

November 1987 MONOGRAPH of POLYBOTRA

A

VG

D
(Z

g
NZ
if? J AP’
ye

10 cm

Z
SA

~

FiGure 50. Polybotrya latisquamosa Moran. a. sterile pinnule; b. apex of sterile leaf; c. stem scale;
d. scale from petiole base; e. pinnule from medial pinna. a—e: Idrobo & Schultes 1106 (US).

110 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

FiGurE 51. Polybotrya sessilisora Moran. a. distal half of sterile leaf; b. abaxial surface of costa and
pinnules (note the thickened, lighter colored margin and dark tortuous scales); c. fertile pinnule, abaxial
view (note the thickened margins and discrete sori); d. stem showing straight, appressed scales;
e. medial pinnules of basal pinna, acroscopic side is up. a,c,d,e: Schultes & Cabrera 13963 (COL).
b: Prance et al. 15332 (NY).

November 1987

Stems 5—10 mm thick, hemiepiphytic; scales
appressed, ascending, linear, 8—12 mm long, dark
castaneous with lighter narrow borders and
strongly denticulate margins. Sterile leaves up to
75 cm long; petiole 2 to as long as the lamina,
scaly with scales similar to those of the stem, but
more tortuous and spreading, the base mostly cor-
date; lamina to 40 cm long, up to 3-pinnate-
pinnatifid, narrowly deltate, the tissue glabrous,
coriaceous, light green, the margins glabrous; pin-
nae narrowly deltate, 17-36 x 7—16 cm, the prox-
imal acroscopic pinnule or segment slightly pro-
longed above the rest; pinnules to 3.5—10 * 1.2—3
cm, arranged anadromically, the basiscopic mar-
gin thickened and decurrent on the costa, the apex
of the less cut medial pinnules merely acute or
obtuse; veins conspicuous and prominulous abax-
ially; axes moderately to sparsely pubescent abaxi-
ally, the hairs colorless, tawny, 0.5—1.0 mm long;
grooves of axes decurrent on those of lower order,
not interrupted, pubescent within by tiny, less than
0.2 mm long, reddish, jointed hairs, covered with
dark, spreading, tortuous, denticulate scales like
those of the petiole. Fertile leaves 3-pinnate, bot-
ryoid, the margins slightly thickened; sori sessile,
round, about 1 mm long; spores (44)46—52(56)
microns long.

Polybotrya sessilisora grows in lowland
forests of the northern Amazon basin (Map 20).
It is probably more common than the number of
collections suggests, since the northern Amazon
basin is poorly collected. I suspect that this species
also occurs in the adjacent Guiana Highlands.

The specific epithet refers to the botryoid sori
that are sessile instead of short-stalked as in other
species of Polybotrya. Moreover, the sori are em-
bedded in the lamina, which is not completely
reduced to the axis (Fig. 51c). As evidenced by
outgroup comparison to other dryopteroid ferns,
the ancestor to Polybotrya surely had fertile leaves
with discrete, round sori from which the lamina
was reduced. Because the lamina of P. sessilisora
is not fully reduced, I interpret its fertile leaf as
the most primitive in the genus. No other species
in the genus has this distinct kind of fertile leaf.

Another distinctive feature of P. sessilisora
is the dark castaneous scales that contrast sharply
with the light green lamina. The scales of the axes
are spreading and tortuous, whereas those of the
stem are appressed and straighter (Fig. 51b,d).

MONOGRAPH of POLYBOTRA 111

These scales become smaller and narrower in the
distal parts of the lamina until they become uni-
seriate, appressed hairs. The lamina is always
broadest at the base, in contrast to the closely
related P. osmundacea, which is usually reduced
at the base. Polybotrya osmundacea also tends to
be much more highly dissected when leaves of
equal size are compared.

Specimens examined: COLOMBIA. Vaupés: Rio
Vaupés, Miti y alrededores, 250 m, Schultes & Cabrera
13963 (COL, GH, US).

BraAziL. Amazonas: Tapuruquara, beside road to
airport, Prance et al. 15332 (NY); Reserva Experimen-
tal, km 60, Manaus—Caracarai road, Conant 1016 (GH),
1482 (GH); Reserva Ducke, km 26, Manaus—Itacoatiara
road, Conant 1080 (GH).

BRAZIL—GUYANA BOUNDARY: Akarai Moun-
tains, height of land between drainage of Rio Mapuera
(Trombetas tributary) and Shodikar Creek (Essequibo
tributary), dense forest 600-800 m, A.C. Smith 2984
(GH, NY).

32. Polybotrya canaliculata Klotzsch (Fig. 52,
Map 19).

Polybotrya canaliculata Klotzsch, Linnaea
20:429. 1847. Lectotype: Venezuela. Ar-
agua: Colonia Tovar, 1846, Moritz 278 (lec-
totype: B!; isolectotypes: F!, GH!, L!, NY!,
P!, US!; photo GH! of L). LECTOPARATYPE:
Colonia Tovar, 1846, Karsten (Coll. II) no.
13 (B!; isolectoparatypes; BM!, L!; photo of
L specimen, NY!).

Acrostichum canaliculatum (Klotzsch) Hook-
er, Species Filicum 5:247. 1864.

Stems 0.5—2 cm thick, hemiepiphytic; scales
ascending with spreading tips, 12—20 x 0.7—1.2
mm, dull brown to dark purple brown, concolor-
ous or rarely with a very narrow hyaline border,
the margins entire to denticulate, opaque to
slightly thickened at the center and base. Sterile
leaves up to 1.8 m long; petiole up to 45 cm long,
shorter than the lamina, scaly at the base or
throughout, the scales becoming progressively
smaller and more ovate-lanceolate and erose-
denticulate upwards, with a blackened basal point
of attachment; /amina glabrous, lanceolate to
triangular or subtriangular, 50-80 cm wide, to
(4)3-pinnate-pinnatifid, subcoriaceous, the abax-
ial surface lighter in color than the adaxial; basal

112 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

-

at

oN ae tae a5 .
Wwe . e aa ~ : s eS i
Ae Zane ; ™ ee

Ne
nia ae

antl
aw

[>a

ys
‘

ZV

i)
H
an

my
2»)
\
aX) yy.
~My)
11cm

[VA

|

FIGURE 52. Polybotrya canaliculata Klotzsch. a. fertile leaf; b. abaxial surface of costa and pinnule;
c. adaxial view of fertile pinnule (the main axis is the costa—same scale as b); d. stem scales; e—f. pin-
nules; g. basal pinna; h. medial pinna, from same leaf as in g. a—d: Moritz 278 (B). e,f: Karsten 13(B).
g,h: van der Werff 3486 (MO).

November 1987

pinnae triangular to subtriangular, up to 40 x 28
cm, the lowermost pinnate throughout except pin-
natifid at the extreme apex; pinnules arranged
catadromically or anadromically, up to 16 x 6.5
em, triangular-lanceolate, the acroscopic side usu-
ally prolonged; costules adaxially deeply grooved
and bordered by a raised, erect to spreading flap
of tissue; fertiary segments entire to pinnatifid
(deeply so in large leaves), equilateral at the base
with a rounded to acute apex; axes scaly, the scales
scattered, flaccid, membranaceous, ovate to lan-
ceolate, denticulate, 1—3.5 mm long, the abaxial
surface usually glabrous; grooves pubescent
within, and especially at segment junctures, the
hairs tiny, less than 0.2 mm long, reddish. Fertile
leaves botryoid, 0.8 x 0.6 m, 4-pinnate; axes with
numerous septate hairs that often grade into narrow
scales; pinnae up to 30 cm long, subtriangular;
sori round to obovate, distinctly stalked, the stalk
about 1 mm long; spores (50)55—66(70) microns
long.

This species grows in cloud forests in the
Andes of northern Venezuela from 1200 to 1600
m (Map 19). Van der Werff and Smith (1980)
state that this species can be abundant locally, but
hat the plants rarely bear fertile leaves.

The specific epithet refers to the raised flap
of tissue that borders the costules and accentuates
the depth of the central groove. This character is
not diagnostic, since other decompound Polybot-
rya species also have raised costular ridges, al-
though in P. canaliculata these ridges tend to be
relatively taller and more well developed.

When available, the botryoid fertile leaves
Fig. 52a,c) distinguish this species from P. os-
mundacea and its allies. Distinguishing vegetative
sharacters are the dull brown, concolorous stem
scales, and the major axes with prominent, ovate
‘0 lanceolate, flaccid scales (Fig. 52b). The pin-
nule arrangement of P.. canaliculata may be either
anadromic or catadromic. This variation is pecu-
liar because the pinnule arrangement is usually
sonstant within most species of Polybotrya. A
similarly variable species in this respect is P.
gomezii, an endemic from Costa Rica. I interpret
P. canaliculata as a primitive species of Polybot-
rya because of its decompound lamina and bot-
ryoid fertile leaf.

I have chosen the Moritz collection as the
lectotype because of its wide distribution in her-
baria and the excellent quality of the specimens.

Specimens examined: VENEZUELA. Aragua:

Colonia Tovar, 1846, Moritz 278 (B, BM, F, GH, L,

MONOGRAPH of POLYBOTRA 113

NY, P, US); Colonia Tovar, 1846, Karsten /3 (B, BM,
L; photo of L specimen, NY); Parque Nacional Henn
Pittier, bosque de Rancho Grande, Tschudi 166 (VEN);
Colonia Tovar, 1854—5, Fendler 262 (GH, MO, NY,
P, PH, US). Fale6n: Sierra de San Luis, arriba de Sta.
Maria, alt. 1200 m, 5 June 1979, van der Werff 3486
(MO, UC); Sierra de San Luis, selva nublada, entre La
Chapa y Uria, Steyermark 99/85 (VEN). Yaracuy:
Dtto. Bolivar, entre las Parchitas, Tierra Fria y Ojo de
Agua, Ortega & Smith 2498 (PORT), 25//] (PORT).
State unknown: Andes of Venezuela, 1889, Goebel
s.n. (P).

33. Polybotrya semipinnata Fée (Fig. 53, Map
18).

Polybotrya semipinnata Fée, Crypt. Vasc.
Brésil. 1:16. 1869. Type: Brazil. Rio de
Janeiro: Yacuacanga, 15 June 1869, Glaziou
2427 (K, P!, RB!, US!; photo of K specimen
at US!).

Aspidium scandens Raddi, Plant. Brasil.
1:34, tab. 49. 1825. Type: Brazil. Raddi s.n.
(FI; isotype: K, photo GH!).

Polybotrya scandens (Raddi) Christ, Bull.
Herb. Boissier, II. 4:965. 1904. nom. illegit.,
non Fée 1852.

Stem 1—1.5 cm thick, hemiepiphytic; scales
membranous, spreading, mostly 9-12 x 0.5—1.2
mm, bright castaneous, concolorous or with a dark
central stripe and lighter borders, the margins
denticulate to strongly erose. Sterile leaves up to
1 m long; petiole 3 to 2 the length of the lamina;
lamina ovate to lanceolate to 3-pinnate but mostly
2-pinnate-pinnatifid throughout, generally 45—
60(75) x 30—55(64) cm, coriaceous, glabrous ex-
cept on major axes; pinnae broadly triangular to
ovate, mostly 15—30 x 11-17 cm, divided at base
and soon becoming pinnatifid distally; pinnules
5—10(13) X 2-3. cm, lanceolate, anadromic
throughout, the bases mostly cuneate, unequal,
the basiscopic side more narrowly cuneate and the
acroscopic side slightly prolonged, proximal ones
with a 3—5 mm long stalk, rarely with the lower-
most acroscopic segment cut to the costule; axes
pubescent abaxially with hairs less than 0.1 mm
long, stiff, colorless, subulate, the scales few or
absent; grooves glabrous or nearly so within, decur-
rent on those of the next lower order. Fertile leaves
smaller than the sterile, 3-pinnate (-pinnatifid),
coenosoric; tertiary segments oblong, generally
3-7 mm long; sporangial stalk paraphysate,;
spores (48)50—62(66) microns long.

114 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

WSS,

S

S

ANS

iG
AY

S
x - \
SN

A,
Z
ZZ,

—

LZ=Z4

1com

FiGuRE 53. Polybotrya semipinnata Fée. a. sterile pinnules; b. sterile leaf; c. middle pinnae of fertile
leaf; d. stem scales. a—c: Brade 806] (PH). d: Duarte et al. 65322 (F).

November 1987

Other illustrations: Raddi, Plant. Brasil. tab.
49. 1825; Mettenius, Filices Hort. Lips., 23, tab.
2, figs. 1-6. 1856 (as P. acuminata); Brade,
Bradea 1:64, fig. 1; 67, fig. 9. 1971 (as P. scan-
dens).

Polybotrya semipinnata is one of five
Polybotrya species that are endemic to the coastal
mountains of southeastern Brazil (Map 18, Table
2), a distribution that emphasizes the biogeo-
graphic distinctness of the Serra do Mar Moun-
tains. The altitudinal range of P. semipinnata is
from 800 to 1000 m.

This plant, easily recognized by its lamina
cutting, is not readily confused with other species
of Polybotrya. The pinnule bases are distinctive
because they are stalked and more narrowly
cuneate on the basiscopic side. The pinnae are
stouter and much less divided compared to those
of other decompound Polybotrya species; the dis-
tal portions soon become pinnatifid, a characteris-
tic that accentuates this less-cut appearance.

An earlier published name, P. acuminata
Link, has often been applied to this species. Since
the type specimen cannot be located and I cannot
discern from Link’s description the species he had
in mind, I treat P. acuminata as a name of uncer-
tain application.

Specimens examined: BRAZIL. Rio De Janeiro:
Yacuacanga, Glaziou 2427 (P, RB, US; photo of K
specimen at US); Estrada Velha da Bocaina, Teresopolis,
Duarte et al. 65322 (F, LP); Serra dos Orgaos,
Teres6polis can. Quebrafrasco, 1000 m, Brade 16456
(RB). Sao Paulo: Piruhyba, Loefgren & Duvall 36217
(RB); prope Rio Grande ad Sao Paulo Railway, 800 m,
Wettstein & Schiffner s.n. (P); Alto da Serra, Lueder-
waldt s.n. (BM, NY), 2/547 (NY); Iguape, morro das
Pedras, Serrinha Peroupara, Brade 806] (NY, PH, UC,
US); Pilar, Gerdes 102 (NY, UC). Sta. Catarina: with-
out locality, Schwacke s.n. (P).

34. Polybotrya speciosa Schott (Fig. 54, Map
21).

Polybotrya speciosa Schott, Genera Filicum
tab. 7. 1834. NEOTYPE: (here chosen) Schott,
Genera Filicum tab. 7, based on material from
“Brasiliae provincia Sebastianopolitana.”

Polybotrya tomentosa Brade, Arq. Inst. Biol.
Veg. Rio de Janeiro 1:224, fig. 2, plates 2
and 3. 1935. Type: Brazil. Minas Gerais:
Serra do Itatiaia Maromba, 25 June 1930,
Brade 10351 (holotype: RB!).

MONOGRAPH of POLYBOTRA 115

Polybotrya osmundacea Willd. var. cris-
popaleacea Rosenst., Feddes Repert. 21:349.
1925. Type: Brazil. Sao Paulo: Alto da Serra,
11 February 1925, Brade 5838 (S; isotype:
UC!).

Polybotrya litoralis Brade, Bradea 1:26, tab.
1, fig. 2. 1969. Type: Brazil. Rio de Janeiro:
Angra dos Reis, Serra do Mar, 29 June 1935,
Brade 14943 (holotype: RB!).

Polybotrya rosenstockiana Brade, Bradea
1:27, tab. 1, fig. 3. 1969. Type: Brazil. Rio
de Janeiro: Serra dos Orgaos, Corrego Beija-
flor, Brade 16579 (holotype: RB!; isotype:
LP!).

Stem 1—3 cm thick; scales commonly brick
red, rarely dull brown, concolorous or with a dark
central stripe, generally 8—20(27) x 0.5—1.5(2.0)
mm, spreading, membranous, the margins den-
ticulate to strongly erose. Sterile leaves up to 1.4
m long; petiole Vio—Y the length of the lamina;
lamina to 1.2 X 0.8 m, lanceolate to ovate, to
3-pinnate but mostly 2-pinnate-pinnatifid through-
out, tomentose to glabrous, the margins sparsely
ciliate to glabrous; pinnae to 40 x 15 cm, free
pinnules usually 5—9; pinnules acroscopic, short
to long triangular, mostly 4.5—8.0(10.0) x 1.5—
3.5 cm, the base stalked, the stalk 2-4 mm long,
the acroscopic side prolonged, the basiscopic side
oblique; tertiary segments generally oblong, the
margins entire to crenulate or dentate; axes tomen-
tose to glabrous or subglabrous, the hairs usually
0.1—0.5 mm long, the scales few, appressed, tor-
tuous, narrow; grooves pubescent within, the hairs
reddish. Fertile leaves coenosoric, 3-pinnate; spo-
rangial stalks paraphysate; receptacle glabrous or
with multicellular, branched paraphyses, these as
long as or slightly longer than the sporangia;
spores (56)60—75(82) microns long.

Other illustrations: See original descriptions
cited above; Brade, Bradea, tab. 3, fig. 4 (as P.
littoralis); tab. 4, fig. 1 (as P. rosenstockiana);
tab. 4, fig. 3; tab. 6, figs. 12, 13 & 15. 1971.

Polybotrya speciosa is endemic to the Serra
do Mar Mountains along the coast of southeastern
Brazil (Map 21). It differs from the four other
species of Polybotrya there by its combination of
strongly denticulate, red stem scales, finely cut
lamina, and multicellular, branched paraphyses
(Fig. 54). The pubescence of the abaxial surface
varies from densely tomentose to nearly glabrous.

Vol. 34, Art. 1

116 ILLINOIS NATURAL HIsTORY SURVEY

FicurE 54. Polybotrya speciosa Schott. a. sterile and fertile leaves; b. branched paraphyses; c- pin- .
nules; d,e. stem scales. a,b,e: Brade 16579 (RB). c,d: Brade 10351 (RB).

November 1987

Five of the specimens examined had the adaxial
surface slightly pubescent whereas the remaining
were completely glabrous. No other species of
Polybotrya, except P. pilosa, has branched para-
physes.

The spore size varies greatly, perhaps reflect-
ing different ploidy levels. The averages from
eight collections, with 15 spores measured from
each, are as follows (in microns): 56, 57, 57, 58,
69, 69, 79, 82. This problem needs further study
of additional collections and cytological samples.

I have been unable to locate Schott’s type,
but I feel quite certain that Schott’s excellent plate
represents this species. Schott’s specimens be-
came part of the Cardinal Hynald herbarium now
located in Budapest, Hungary (BP). The curator
of the fern collection at Budapest, Mr. Tibor Szer-
dahelyi, informed me (in litt.) that much of
Schott’s type material had been destroyed during
World War II and that he could not find the type.

My placement of the three species described
by Brade in synonymy with P. speciosa requires
comment. I find no differences between the types
of P. tomentosa and P. litoralis—the two might
well have been collected from the same individual.
Given this likeness, I find it odd that Brade (1969c)
did not mention P. tomentosa in his discussion
after the description of P. litoralis. I also place
P. rosenstockiana and P. osmundacea var. cris-
popaleacea in synonymy, although they differ
slightly from most specimens of P. speciosa by
their somewhat broader stem scales (Fig. 54, com-
pare d & e). They also differ by their leaves,
which are less pubescent, primarily so along the
axes and veins. Both these characters, however,
intergrade and do not correlate with any others.

Specimens examined: BRAZIL. Rio de Janeiro:
Serra dos Orgaos, Corrego Beijaflor, Brade 16579 (LP,
RB); Serra do Taquaral, Brade 17464 (MO, NY); There-
zopolis, Brade 9773 (BM, UC), 9843 (NY); Parque
Nacional de Serra dos Orgaos, de la Sota 2343 (LP);
Serra dos Orgaos, Morro Assu, Luetzelburg 6858 (US);
Organ Mts., Rose & Russell 20790 (US); Corcavado,
collected by the U.S. South Pacific Exploring Expedi-
tion, 1838—42 (NY, US); Mt. Tijuca, Cuyler 4796 (US);
Angra dos Reis, Serra do Mar, Brade 14943 (RB);
Guanabara, Estrada do Sumaro, Pabst et al. 6772 (LP).
Minas Gerais: Serra do Itatiaia, Maromba, Brade 10351
(RB); Itatiaia, Maromba, Brade 202/4 (F, LP, MO,
NY); same locality, Brade 12616 (BM); Tijuca, Alston
899] (BM). Sao Paulo: Alto da Serra, Brade 5838 (UC).
State unknown: “Brazil,” Webb 26 (GH).

MONOGRAPH of POLYBOTRA 117

35. Polybotrya pilosa Brade (Fig. 55, Map 20).

Polybotrya pilosa Brade, Bradea 1:27, tab.
1, fig. 4. 1969. Type: Brazil. Rio de Janeiro:
Teresopolis, Varesea, 1000 m, 27 October
1929, Brade 9787 (holotype: RB!; isotypes:
NY!, UC!).

Differing from P. tomentosa only by charac-
teristics of the pubescence; hairs 1—2(2.5) mm
long, pilose, acicular, pluricellular, whitish or
tawny, disposed along the abaxial surface of the
veins and major axes, rarely on the intervening
laminar tissue between the veins, also occurring
on the adaxial surface and protruding from the
costal and rhachidial grooves. Paraphyses present,
of both branched and unbranched types.

Polybotrya pilosa is endemic to southeastern
Brazil (Map 20). This species may be subject to
reinterpretation in the future, as it differs from P.
speciosa only by its pubescence, which shows no
intergradation between the two species. The hairs
of P. speciosa are less than 1 mm long, somewhat
tortuous, and occur on the veins and intervening
laminar tissue. The hairs of P. pilosa are more
than | mm long, straightish, and acicular; they
occur only along the veins, not on the intervening
laminar tissue (Fig. 55a). These differences are
readily apparent to the unaided eye. The rein-
terpretation of P. pilosa will require fieldwork
designed to study intra- and interpopulational vari-
ation in southeastern Brazil.

Other illustrations: See Brade’s original de-
scription, cited above.

Specimens examined: BRAZIL. Rio de Janeiro:
“near Rio de Janeiro,” collected by U.S. Exploring Ex-
pedition, 1838-42 (US); Tijuca, Brade 20733 (NY,
UC); “Rio Janeiro,” 1851, Andersjon s.n. (LP); Organ
Mts., Wagner s.n. (GH); Teres6polis, Varesea, Brade
9787 (NY, RB, UC); Serra Estrela, Weddell 952 (P);
“Brasilia,” Riedel 8] (GH).

118 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

Ficure 55. Polybotrya pilosa Brade. a. costa and pinnule with characteristic long, acicular hairs;
b. sterile leaf; c. stem scales. a: Wagner s.n. (GH). b,c: Brade 9787 (RB).

November 1987

Names of Uncertain Application

Polybotrya acuminata Link, Hort. Berol. 2:135.
1833. Psomiocarpa acuminata (Link) Presl,
Epim. Bot. 162. 1849. Type: Brazil. Collector?
I have been unable to locate the type and cannot
discern from the original description the taxon to
which this name applies. The type specimen may
have been lost during World War II; I have, how-
ever, Link’s other type specimens from Berlin.
Mettenius (1856, tab. 2) has an excellent illustra-
tion of P. semipinnata, which he refers to as P.
acuminata, but I do not know whether he saw
type material.

Polybotrya fulvostrigosa Christ, Bull. Herb. Bois-
sier, Il. 1:70. 1901. Type: Peru. Loreto: Cerro
de Canchahuaya, Huber 1448. | cannot find the
type and am uncertain from Christ’s description
to what species this name belongs.

Polybotrya lomarioides Mettenius, Filices Lech-
ler. 2:5. 1858. Type: Peru. Puno: San Gavan,
Lechler s.n.. | have not seen the type and cannot
be certain from the description if this name applies
to a Polybotrya species or to another genus.

Polybotrya nutans Kunze, Linnaea 9:24. 1834.
Type: Peru. “Sylvae flor. Peruv. ad Pampayaco

MONOGRAPH of POLYBOTRA 119

in cortice arborum vetustarum parasitica, Jul.
1829,” Poeppig s.n. (B!, P!). The fertile leaf of
the type specimen came from a species of Polybot-
rya, but I do not know which one. The sterile leaf
of the type specimen is a tree fern, perhaps a
species of Trichipteris.

Polybotrya scandens Fée, Genera Filicum 47.
1852. Type: Venezuela. Lagunetta, Galeotti. |
have not seen the type and cannot ascertain from
Fée’s description the species to which this name
applies. I suspect it is P. osmundacea. Windisch
(1982) reported Fée’s specimen of P. scandens to
be at the Botanical Gardens in Rio de Janeiro
(RB). Although I received other Fée specimens
from RB, P. scandens was not among them. Fée
gives the location as “Lagunetta, Mexico,” but
this site is certainly an error; see the discussion
under P. serratifolia.

Polybotrya trapezoides Link, Filicum Species
164. 1841. Type: Cultivated plant at Berlin of
unknown origin (B!). I received a specimen from
Berlin (B) labelled as species but cannot be sure
that this is the type. This specimen, however, is
certainly not a species of Polybotrya. Because it
is so young, I am uncertain what genus it repre-
sents.

120 ILLINOIS NATURAL HIistORY SURVEY

Excluded Taxa
Excluded Subgenera

Polybotrya subg. Ectoneura Fée, Mém. Fam.
Foug. (Hist. Acrost.) 75. 1845 (nomen illegit.).
= Bolbitis (fide Hennipman 1977).

Polybotrya subg. Egenolfia Fée, Mém. Fam.
Foug. (Hist. Acrost.) 14. 1845. = Bolbitis (fide
Hennipman 1977).

Excluded Sections

Polybotrya sect. Arthrobotrya v.A.v.R., Handb.
725. 1908. =Lomariopsis sect. Polyseriatae
(fide Holttum 1978).

Polybotrya sect. Egenolfia Diels, E&P Nat. Pfl.
Fam. 1:195. 1900. =Bolbitis (fide Hennipman
1977).

Polybotrya sect. Lomagramma Kuhn, Ann. Mus.
Bot. Ludg.-Bat. 4:198. 1899. =Lomagramma
(fide Holttum 1978).

Polybotrya sect. Teratophyllum Christ, Farnkr.
Erde 42. 1897. =Lomariopsis sect. Poly-
seriatae (fide Holttum 1978).

Excluded Species

Polybotrya acrostichoides Kuhn, Fil. Afr. 52.
1868. =Bolbitis acrostichoides (Sw.) Ching
(fide Hennipman 1977).

Polybotrya apiifolia Kunze, Farnkr. 1:142, tab.
62. 1844. =Psomiocarpa apiifolia (Kunze)
Presl.

Polybotrya appendiculata (Willd.) J. Smith, Jour.
Bot. 4:150. 1841. =Bolbitis appendiculata
(Willd.) Iwatsuki, (fide Hennipman 1977).

Polybotrya arfakensis Gibbs, Arfak 71. 1917.
=Alsophila biformis Rosenstock (fide Holttum
1963).

Polybotrya articulata Fée, Mém. Fam. Foug.
(Hist. Acrost.) 74, tab. 37. 1845. = Teratophyl-
lum articulatum (Fée) Kuhn (fide Holttum 1978).

Polybotrya aspidioides Grisebach, Cat. Pl. Cub.
276. 1866. = Atalopteris aspidioides (Grisebach)
Maxon & C. Chr.

Polybotrya asplenifolia (Belanger) Presl, Tent.
Pterid. 231. 1836. =Bolbitis appendiculata
(Willd.) Iwatsuki (fide Hennipman 1977).

Polybotrya aurita Blume, Fl. Jay. Fil. 15, tab. 1.
1828. =Stenosemia aurita (Sw.) Presl (fide
Christensen 1905).

Vol. 34, Art. 1

Polybotrya bifurcata (L.f.) J. Smith, Jour. Bot.
4:150. 1841. =Elaphoglossum _ bifurcatum
(Jacq.) Mickel (fide Mickel 1980).

Polybotrya blumeana (Fée) Mettenius, Fil. Lips.
24, tab. 2, fig. 10. 1856. =Leptochilus lomari-
oides Blume.

Polybotrya cervina (L.) Kaulf., Enum. Fil. 55.
1824. =Olfersia cervina (L.) Kunze, Flora
7:312. 1824.

Polybotrya cicutaria Blume, Enum. Fil. 100.
1828. =Stenosemia aurita (Sw.) Presl (fide
Christensen 1905).

Polybotrya duplicato-serrata Hayata, Ic. Fl. For-
mosa 5:305, fig. 123. 1915. =Bolbitis rhizo-
phylla (Kaulf.) Hennipman (fide Hennipman
1977).

Polybotrya exaltata Brackenridge, U.S. Expl.
Exp. 16:78. 1854. =Bolbitis rhizophylla
(Kaulf.) Hennipman (fide Hennipman 1977).

Polybotrya filiculifolia (L.) Farw., Amer. Midl.
Nat. 12:303. 1931. =Anemia?

Polybotrya flabellata (Willd.) Mettenius, Ann.
Sci. Nat., Paris, V, 2:206. 1864. = Peltapteris
peltata (Sw.) Morton.

Polybotrya fraxinifolia (Presl) Mettenius, Fil.
Lechler. 2:8. 1959. =Bolbitis serratifolius
(Kaulf.) Schott (fide Hennipman 1977).

Polybotrya furcata (L.f.) Kuhn, Fil. Afr. 52.
1868. = Elaphoglossum bifurcata (Jacq.) Mickel
(fide Mickel 1980).

Polybotrya gaudichaudiana (Gaudich.) Moore,
Index Fil. XV. 1857. =Bolbitis appendiculata
(Willd.) Iwatsuki (fide Hennipman 1977).

Polybotrya hamiltoniana (Wallich) Fée, Mém.
Fam. Foug. (Hist. Acrost.) 78. 1845. =Bolbitis
appendiculata (Willd.) Iwatsuki (fide Hennipman
1977).

Polybotrya helferiana Kunze, Farnkr. Erde 2:35,
tab. 94. 1848. = Bolbitis appendiculata (Willd.)
Iwatsuki (fide Hennipman 1977).

Polybotrya intermedia Fée, Mém. Fam. Foug.
(Hist. Acrost.) 76, tab. 40, fig. 1. 1845. = Bol-
bitis rhizophylla (Kaulf.) Hennipman (fide Hen-
nipman 1977).

Polybotrya lomarioides (Blume) Kuhn, Ann.
Mus. Ludg.-Bat. 4:295. 1869. =Lomagramma
lomarioides (Blume) J. Smith (fide Holttum
1978).

November 1987

Polybotrya marattioides Brackenridge, U.S.
Expl. Exped. 16:79. 1854. =Asplenium marat-
tioides (Brackenridge) C. Chr. (fide Christensen
1905).

Polybotrya marginata Blume, Enum. PI. Jay. 100.
1828. = Bolbitis appendiculata (Willd.) Iwatsuki
(fide Hennipman 1977).

Polybotrya montana (Gaudich.) Moore, Index Fil.
350. 1862. =Bolbitis appendiculata (Willd.)
Iwatsuki (fide Hennipman 1977).

Polybotrya nana Fée, Mém. Fam. Foug. (Hist.
Acrost.) 75, tab. 38, fig. 1. 1845. =Blechnum
lanceolatum (R. Br.) Sturm. (fide Christensen
1905).

Polybotrya neglecta Fée, Mém. Fam. Foug. (Hist.
Acrost.) 75, tab. 39, fig. 2. 1845. =Bolbitis
rhizophylla (Kaulf.) Hennipman (fide Hennipman
1977).

Polybotrya nieuwenhuisenii Raciborski, Bull. Int.
Ac. Cracovie 57. 1902. =Heterogonium steno-
semioides (Baker) Christ (fide Holttum 1975).

Polybotrya nodiflora Belanger, Voy. Ind. Or. Bot.
2:17. 1833. =Bolbitis appendiculata (Willd.)
Iwatsuki (fide Hennipman 1977).

Polybotrya orientalis Blume, Enum. Pl. Jav. 99.
1828. =Stenosemia aurita (Sw.) Presl (fide
Christensen 1905).

Polybotrya peltata (Sw.) J. Smith, Jour. Bot.
4:150. 1841. = Peltapteris peltata(Sw.) Morton.

Polybotrya plumieri (Fée) Moore, Index Fil. 13.
1857. =Elaphoglossum plumieri Moore (fide
Proctor 1977).

Polybotrya polyphylla (Brackenridge) C. Chr.,
Index Fil. 505. 1906. = Lomagramma polyphylla
Brackenridge.

Polybotrya prolifera Mettenius, Fil. Lips. 24, pl.
2, fig. 11. 1856. =Bolbitis subcrenata (Hooker
& Grev.) Ching (fide Hennipman 1977).

Polybotrya pteroides (J. Smith) Kuhn, Ann. Mus.
Bot. Lugd.-Bat. 4:295. 1869. =Lomagramma
pteroides J. Smith (fide Holttum 1978).

Polybotrya quercifolia (Retz.) Mettenius, Fil.
Lechler. 2:12. 1859. =Leptochilus zeylandicus
(Houtt.) C. Chr. (fide Christensen, 1905).

MONOGRAPH of POLYBOTRA 121

Polybotrya rhizophylla (Kaulf.) Presl, Tent.
Pterid. 231. 1836. =Bolbitis rhizophylla
(Kaulf.) Hennipman (fide Hennipman 1977).

Polybotrya serrulata Fée, Mém. Fam. Foug.
(Hist. Acrost.) 76, pl. 39, fig. ii. 1845. = Bol-
bitis rhizophylla (Kaulf.) Hennipman (fide Hen-
nipman 1977).

Polybotrya sinensis (Baker) C. Chr., Index Fil.
57. 1913. =Bolbitis sinensis (Baker) Iwatsuki
(fide Hennipman 1977).

Polybotrya sorbifolia (L.) Keyserling, Pol. Cyath.
Herb. Bung. 32. 1873. nomen __ illegit.

=Lomariopsis sorbifolia (L.) Fée (fide Proctor
1977).

Polybotrya stenosemioides_ (Baker) Copel.,
Polypod. Philipp. 40. 1905. =Heterogonium
stenosemioides (Baker) C. Chr. (fide Holttum
1975).

Polybotrya subquinquefida (Fée) Mettenius, Fil.
Lechler. 2:12. 1859. =?Leptochilus latifolium
(Meyen) C. Chr. (fide Christensen 1905).

Polybotrya taccaefolia (J. Smith) Mettenius, Fil.
Lechler. 2:12. 1859. =Leptochilus latifolius
(Meyen) C. Chr. (fide Christensen 1905).

Polybotrya tenuifolia (Desv.) Kuhn, Fil. Afr. 52.
1868. =Stenochlaena tenuifolia (Desv.) Moore
(fide Christensen 1905).

Polybotrya teysmanniana (Baker) Posthumus,
Rec. Trav. Bot. Neerl. 33:872. 1930. =
Stenosemia teysmanniana (Baker) Diels (fide
Christensen 1905).

Polybotrya trilobata (J. Smith) Mettenius, Fil.
Lips. 24. 1856. = Leptochilus latifolius (Meyen)
C. Chr. (fide Christensen 1905).

Polybotrya tripartita (Hooker & Grev.) J. Smith,
Jour. Bot. 4:150. 1841. = Peltapteris tripartita
(Hooker & Grey.) Morton (fide Morton 1955).

Polybotrya vivipara Hooker, Exotic Flora, 2: pl.
107. 1825. =Bolbitis appendiculata (Willd.)
Iwatsuki subsp. vivipara (Hooker) Hennipman
(fide Hennipman 1977).

Polybotrya wilkesiana Brackenridge, U.S. Expl.
Exped. 16:80, tab. 10. 1854. =Teratophyllum

wilkesianaum (Brackenridge) Holttum (fide
Holttum 1978).

122 ILLINOIS NATURAL HistoRY SURVEY

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Gomez, L.D. 1976. Contribuciones a la pteridologia
centroamericana, I. Enumeratio Filicum Nicara-
guensium. Brenesia 8:41—57.

HENNIPMAN, E. 1977. A monograph of the fern genus
Bolbitis (Lomariopsidaceae). Leiden Botanical
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HoutrruM, R.E. 1959. Vegetative characters distin-
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Dryopteris of Christensen’s Index Filicum and other
ferns of similar habit and sori. Gardens’ Bulletin
(Singapore) 17:361—367.

Vol. 34, Art. 1

HoLttuM, R.E. 1963. Cyatheaceae. Flora Malesiana,
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HoLtruM, R-E. 1975. The fern genus Heterogonium
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Ho.ttuM, R.E. 1982. The continuing need for more
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Tectaria, 1. A commentary on recent schemes of
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Koptur, S., A.R. SMITH, and I. BAKER. 1982. Nec-
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LELLINGER, D.B. 1972. Five new species of South
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Moran, R.C. 1986. The neotropical fern genus Olfer-
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November 1987

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MONOGRAPH of POLYBOTRA 123

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124

ILLINOIS NATURAL HisTORY SURVEY

Taxa and Distribution of
Polybotrya

Numbers correspond to the species numbers as-
signed in the taxonomic treatment.

1.

Polybotrya serratifolia (Fée) Klotzsch: Trini-
dad, Venezuela.

Polybotrya polybotryoides (Baker) Christ:
Mexico, Belize, Guatemala, Honduras, Costa
Rica, Panama, Colombia, Ecuador, Peru.
Polybotrya suberecta (Baker) C. Chr.: Co-
lombia, Ecuador, Peru.

. Polybotrya andina C. Chr.: Ecuador.

5. Polybotrya sorbifolia Kuhn: Costa Rica, Ven-

ezuela, Colombia, Brazil.

. Polybotrya fractiserialis (Baker) J. Smith:

French Guiana, Surinam, Guyana, Ecuador,
Peru, Bolivia.

. Polybotrya crassirhizoma Lellinger: Colom-

bia, Ecuador, Peru, Bolivia, Brazil.

8. Polybotrya espiritosantensis Brade: Brazil.

9. Polybotrya caudata Kunze: Mexico, Guate-

mala, Belize, Honduras, Nicaragua, Costa
Rica, Panama, Trinidad, French Guiana,
Surinam, Guyana, Venezuela, Colombia,
Ecuador, Peru, Bolivia, Brazil.

. Polybotrya goyazensis Brade: Brazil, Para-

guay.

. Polybotrya pubens Martius: Colombia, Ecua-

dor, Peru, Bolivia, Brazil.

. Polybotrya glandulosa Kuhn: Venezuela,

Peru, Brazil.

. Polybotrya lechleriana Mettenius: Guyana,

Colombia, Ecuador, Peru, Bolivia.

. Polybotrya attenuata Moran: Colombia.
. Polybotrya stolzei Moran: Colombia.
. Polybotrya alfredii Brade: Nicaragua, Costa

Rica, Panama, Ecuador, Peru, Bolivia.

We

18.
19.
20.
21.

Pap

23.
24.

DS:
26.

27.
28.

29:

30.
31.

32:

33.
34.
35:

Vol. 34, Art. 1

Polybotrya botryoides (Baker) C. Chr.: Co-
lombia.

Polybotrya lourteigiana Lellinger: Colombia.
Polybotrya pittieri Lellinger: Colombia.
Polybotrya cylindrica Kaulfuss: Brazil.
Polybotrya hickeyi Moran: Colombia, Bo-
livia.

Polybotrya puberulenta Moran: Ecuador,
Bolivia.

Polybotrya alata Moran: Panama.

Polybotrya aequatoriana Moran: Ecuador,
Bolivia.

Polybotrya appressa Moran: Ecuador.

Polybotrya altescandens C. Chr.: Colombia,
Ecuador, Peru.

Polybotrya gomezii Moran: Costa Rica.

Polybotrya osmundacea Willd.: Guatemala,
Honduras, Nicaragua, Costa Rica, Panama,
Cuba, Jamaica, Haiti, Grenada, Trinidad,
French Guiana, Guyana, Venezuela, Colom-
bia, Ecuador, Peru, Bolivia, Brazil.

Polybotrya cyathifolia Fée: Guadeloupe,
Martinique.

Polybotrya latisquamosa Moran: Colombia.

Polybotrya_sessilisora Moran: Colombia,
Brazil.

Polybotrya canaliculata Klotzsch: Vene-
zuela.

Polybotrya semipinnata Fée: Brazil.
Polybotrya speciosa Schott: Brazil.

Polybotrya pilosa Brade: Brazil.

Distribution Maps

The small open dots on the twenty-one distribution
maps that follow indicate towns. Other symbols
are explained in the legends.

November 1987

Map |. Distribution of Polybotrya suberecta (Baker) C. Chr. (diamond) and P.

Klotzsch (circle). Map 2. Distribution of Polybotrya polybotryoide
andina C. Chr. (square).

MONOGRAPH of POLYBOTRA

serratifolia (Fée)
S (Baker) Christ (circle) and P.

Vol. 34, Art. 1

ILLINOIS NATURAL HIsTORY SURVEY

Map 3. Distribution of Polybotrya sorbifolia Kuhn (diamond) and P. fractiserialis (Baker) J. Smith

(circle).

Map 4. Distribution of Polybotrya crassirhizoma Lellinger.

MONOGRAPH of POLYBOTRA

November 1987

Map 6. Distribution of Polybotrya pubens Mar-

Map 5S. Distribution of Polybotrya caudata Kunze.

tius (circle) and P. goyazensis Brade (diamond).

128 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

Map 7. Distribution of Polybotrya lechleriana Mettenius (circle), P. attenuata Moran (triangle), and
P. glandulosa Kuhn (square). Map 8. Distribution of Polybotrya alfredii Brade (circle) and P.
botryoides (Baker) C. Chr. (triangle). Map 9. Distribution of Polybotrya stolzei Moran. Map 10.
Distribution of Polybotrya lourteigiana Lellinger. Map 11. Distribution of Polybotrya pittieri Lel-
linger.

November 1987 MONOGRAPH of POLYBOTRA 129

Map 12. Distribution of Polybotrya altescandens C. Chr.
Moran (circle) and P. puberulenta Moran (triangle).
Moran (circle) and P. alata Moran (triangle). MAP
Moran. Map 16. Distribution of Polybotrya cylindrica
rya appressa Moran.

Map 13. Distribution of Polybotrya hickeyi
Map 14. Distribution of Polybotrya gomezii
15. Distribution of Polybotrya aequatoriana
Kaulfuss. Map 17. Distribution of Polybot-

130 ILLINOIS NATURAL HISTORY SURVEY Vol. 34, Art. 1

Map 18. Distribution of Polybotrya osmundacea Willd. (circle) and P. semipinnata Fée (triangle).
Map 19. Distribution of P. latisquamosa Moran (solid square), P. canaliculata Klotzsch (triangle),
and P. cyathifolia Fée (open square).

November 1987 MONOGRAPH of POLYBOTRA 131

Map 20. Distribution of Polybotrya sessilisora Moran (circle) and P. pilosa Brade (triangle).
Map 21. Distribution of Polybotrya speciosa Schott (circle) and P. espiritosantensis Brade (triangle).

132 ILLINOIS NATURAL History SURVEY

Index to Collectors’ Numbers

Numbers in parentheses refer to the species num-
bers assigned in the taxonomic treatment.

Acosta Solis 6358 (2).
Acuna 17531 (28).
Aguilar 947 (6).
Alfaro 8073 (5).
Allard 20661 (6); 21609 (28); 21997 (28);
22305 (6); 22526 (6); 22593 (9).
Alston 7341 (28); 7641 (28); 8278 (21);
8991 (34).

Alverson et al. 342 (2).

Anderson 10152 (5).

Antonio 4019 (2); 5133 (28).

Argent & Richards 6651 (10).

Argent et al. 6336 (10).

Aristeguieta 1780 (28); 3963 (1).

Armond 298 (2).

Atwood 3711 (28).

Aymard et al. 952 (9).

B.T. 443 (9).

Bailey & Bailey 541 (9).

Balsley 4787 (11); 4802 (11).

Balslev & Madsen 10649 (28).

Barbour 4764 (9); 4790 (9).

Barclay 2199 (9).

Barkley & Gutiérrez V.

Barnard et al. 411 (28).

Beard 1252 (28).

Beck 1635 (7); 3060 (24); 3108 (24);
4924 (11); 8037 (9).

Bennett 23 (28).

Benoist 1271 (9).

Berg et al. P18138 (28).

Berry 948 (1).

Billiet & Jadin 1683 (9).

Biolley 73 (16); 10688 (9).

Boom & Mori 1856 (9).

Boutin & Schlosser 5902 (2).

Brade 372 (2); 554 (16); 5838 (34);
8061 (33);9365 (20);9773 (34);9787 (35);
9843 (34); 10351 (34); 12576 (20);
12614 (20); 12616 (34); 13982 (20);
14943 (34); 15373 (10); 16456 (33);
16579 (34); 16824 (16); 17464 (34);
18224 (8); 19972 (20); 20214 (34);
20733 (35); 20931 (20).

Brade & Brade 67 (16); 98 (16); 374 (9).

Brandbyge & AsanzaC. 31870 (11);
32881 (7).

Brandbyge et al.
33684 (7).

1897 (28).

32544 (7); 32617 (7);

Vol. 34, Art. 1

Breedlove 34101 (9).
Breedlove & Smith 21886 (2).

Brenes 11676 (16); 14248 (16); 21980 (16).
Britton et al. 1267 (28); 1935 (9); 2144 (9);
2276 (28).

Broadway 560 (9); 2520 (28); 5358 (9);

5589 (28); 5717 (9); 6459 (28);
6902 (28); 9207 (9); 9947 (1);
9948 (28); 9949 (1); 9950 (28);

9951 (28).

Buchtien 11 (6);13 (6);35 (9&11); 260 (11);
290 (9); 298 (28); 299 (11); 1015 (6);
1066 (28); 1123 (11); 1124 (11); 2138 (6);
5164 (16); 5308 (13).

Bues_ 1743 (13).

Bunting 9516 (28).

Burch 4414 (16); 4505 (16): 4613 (28).

Burger & Antonio 11263 (9).

Burger & Stolze 5865 (9).

Cain 74 (28).

Camp 1298 (28); 1359 (28).

Cardenas 1249 (6).

Castellanos 25710 (20).

Cazalet & Pennington 7720 (11).

Chrysler 4836 (9).

Chrysler & Roever 5095 (16).

Churchill 5776 (28).

Churchill & de Nevers

Clement 725 (28).

Clute 259 (28).

Conant 940 (9); 1016 (31); 1080 (31):
1482 (31).

Cooper 10240 (16).

Cornman 544 (9); 927 (16); 1178 (16);
1233 (16).

Cremers 4460 (9); 6374 (9); 6523 (9):
7369 (9): 7552 (9); 7951 (9).

Croat 690 (28); 738 (28); 4266 (2):
5114 (9); 6850 (9); 7362 (9);

8025 (9); 8153 (9); 9000 (9);
9004 (9); 9103 (9); 10804 (9);
11544 (9); 12114 (2); 12143 (2);
13431 (16); 14687 (2); 14742 (2):
14786 (2); 15081 (9); 15256 (9);
17161 (2); 17367 (9); 17620 (9);
17689 (11); 18219 (9); 18388 (9):
18476 (28); 18508 (11); 19220 (9);
19717 (9); 19771 (11); 20287 (28);
20543 (28); 20551 (7); 20763 (7):
20786 (28); 20851 (28); 21014 (6);
21152 (6); 21191 (9); 21987 (28);
22260 (28); 22682 (28); 22706 (23);
22789 (2); 23011 (23); 23244 (2);

4993 (2).

November 1987

24541 (9); 24566 (2); 27648 (2);
33452 (9); 35176 (9); 35608 (16);
36007 (16); 36669 (16); 36761 (16);
36800 (2); 37757 (2); 37794 (2);
38047 (28); 41637 (2); 49718 (13);
51154 (6); 55844 (26); 56510 (3).

Croat & Folsom 34051 (2).

Croat & Grayum 60022 (28).

Croat & Porter 15342 (9); 15436 (9).

Crosby 76 (28).

Crueger 139 (9).

Cuatrecasas 14246 (28); 15527 (18);
22143 (18).

Cuyler 4796 (34).

Davidse & Gonzalez 19444 (28).

DeWolf 385 (16).

Dodson 7380 (3).

Dodson et al. 8679 (3).

Donnell Smith 6939 (16).

Duarte 3179 (20).

Duarte & Pereira 65323 (20).

Duarte et. al. 65322 (33).

Dudley 10168 (9); 10325 (13); 11513 (9);
13005 (28); 13258 (16); 13290D (2);
18265A (16).

Dumont et al. 7439 (28).

Dunn & LeDoux 22005 (9).

Duque-Jaramillo 1868 (18).

Dusén 4423 (20); 6926 (20); 6984 (20);
13658 (20); 15351 (20); 15353 (20).

Duss 1503 (29); 3897 (29); 4719 (29).

Dwyer 8337 (16).

Dyer A225 (9).

Eggers 5324 (28).

Ekman 3769 (28); 4734 (28); 5214 (28);
14210 (28).

Englesing 291 (9).

Esposto 10928 (3).

Evans & Bowers 2792 (28); 2944 (16);
3152 (2).

Evoy 104 (9).

Ewan 16846 (9).

Fargens 251 (20).

Faull 12583 (28).

Fay 345 (28); 373 (28); 472 (9); 859 (1).

Fendler 69 (28); 105 (9); 235 (1); 261 (1);

262 (32).
Fernandez 951 (9).
Fiedler & Koptur 51 (27).
Fisher 132 (28).
Fleming & Fleming 52 (28).
Folsom 3590 (2).
Forero & Jaramillo 1745 (9); 2469 (13).

MONOGRAPH of POLYBOTRA

Foster P-84-42 (9); P-84-91 (6); 85-37 (3);

85-162 (2); 4020 (6); 7455 (6); 7858 (11);

7937 (6); 9284 (7).

Foster & Foster 854 (8).

Foster & Kennedy 1814 (2).

Fournier 357 (2).

Gardner 1901 (5).

Gastony 43 (28).

Gaudichaud 73 (20).

Gentle 2947 (9).

Gentry & Juncosa 41021 (28).

Gentry et al. 27365 (6); 27880 (7).

Gerdes 102 (33).

Gerrera 1624 (13).

Gill 47 (7).

Glaziou 955 (20); 2427 (33); 2428 (20);
14456 (10).

Gleason 582 (9).

Goedas 217 (20).

Goémez 3324 (2); 3354 (9); 4528 (9);
6914 (2); 7027 (2); 7122 (5);
18064 (2); 18906 (16);
19489 (9 & 28).

Gomez & Cittar 6409 (9); 6731 (9).

Gomez et al. 20423 (9).

Gonggrijp & Stahel 3300 (6).

Grant 10283 (21); 10436 (9); 10556 (3).

Granville 971 (9); 991 (9); 1143 (9);
1165 (9); 3674 (6); 3865 (28); 4012 (9):
B.4711 (9); 4841 (9); 5121 (6); 5219 (6).

Grayum & Sleeper 3448 (16).

Grijalva 300 (28).

Guppy 462 (9).

Gutiérrez & Barkley 170109 (28).

Haerchen 124 (20).

Hammel 3503 (2).

Handro 1229 (20); 2228 (20).

Harley et al. 10922 (10).

Harling et al. 7535 (28).

Hart 53 (9); 228 (1); 229 (28).

Hassler 11618 (10).

Hatch 90 (28).

Hatschbach 7424 (20); 8141 (20); 10749 (20);

25118 (10).
Haught 1325 (9); 1757 (5); 5498 (13).
Hayes 8 (9).
Henri-Stanislas 1709 (2); 1710 (28).
Herrera 1200 (6).
Hickey 801 (21).
Hioram 2495 (28).
Hioram & Clement 6424 (28).
Hodge 6592 (26); 6780 (2).
Holdridge 5153 (2).

133

134 ILLINOIS NATURAL HIstORY SURVEY

Holm & Iltis 200 (9 & 28).

Holm-Nielsen & Jeppsen 663 (11); 987 (7).

Holm-Nielsen et al. 4489 (9); 4495 (9);
22121 (11); 22484 (7).

Huber 2599 (9); 7235 (9).

Humboldt 459a,b (28).

Hunnewell 16514 (5); 18511 (20).

Idrobo & Schultes 1106 (30).

Idrobo et al. 10116 (28).

Iriarte 52 (1).

Irwin 55161 (9).

Irwin et al. 54669 (6); 54784 (6); 54896 (9).

Jameson 33 (26).

Jaramillo et al. 30783 (28).

Jermy 2137 (9); 2368 (28); 2846 (28);
3122 (9); 10826 (9); 11195 (28).

Jiménez M. 803 (16); 3200 (9); 3209 (2);
3210 (2); 3299 (28); 3304 (28).

Johnson 1221 (9).

Johnston 452 (9).

Jones & Facey 3246 (9).

Juncosa 1369 (17); 1467 (3).

Kalbreyer 1254 (2); 1798 (2); 1873 (17);
1877 (3).

Karsten 13 (32).

Kellerman 7354 (9).

Kennedy et al. 2074 (2).

Killip 2544 (9); 2845 (9); 2929 (9);
5163 (16); 5202 (16); 5386 (16);
5429 (16); 11343 (9); 11549 (15).

Killip & Cuatrecasas 38902 (28).

Killip & Lasser 37756 (28).

Killip & Smith 15341 (28); 20210 (21);
23876 (7); 23916 (11); 23991 (28);
24605 (7); 25452 (28); 25921 (13);
25972 (11); 26194 (11); 26543 (11);
26637 (9); 26955 (11); 28760 (11);
30690 (9).

Kirkbride 404 (5).

Kirkbride & Hayden 274 (2).

Klawe 1474 (28); 1504 (28); 1545 (2).

Klug 1166 (9); 1386 (7); 1390 (12);
3208 (28).

Knapp 4966 (9).

Knapp & Dressler 5461 (2).

Knapp & Mallet 5133 (9); 5402 (9).

Knapp et al. 4479 (9); 4542 (2).

Kramer 1954 (9).

Laegaard 51136 (11);51150 (11);51224 (11).

Lankester 653 (16).

Lasser & Vareschi 3926 (9).

Lechler 2156 (13); 2176 (13); 2321 (9);
2329 (9).

Vol. 34, Art. 1

Lehmann B.1T413 (13); SOOB (14); 2998 (9).

Lehnun 2562 (20).

Lellinger & delaSota 26 (9);39 (2);190 (2);
213 (28); 248 (3); 250 (28); 251 (18);
280 (15); 281 (13); 284 (15); 387 (18);
496 (9); 553 (28); 589 (9); 643 (9);

747 (15); 748 (15); 768a (18); 790 (9);
843 (18); 894 (3); 899 (18); 948 (18).

Lellinger & White 1244 (16); 1361 (16);
1436 (28).

Lent 299 (28); 1084 (27); 2642 (16);

3537 (27).

Leén 226 (6).

Liesner 820 (2); 16133 (28); 16293 (12).

Liesner & Judziewicz 14855 (16).

Liesner et al. 8359 (28).

Lindeman 4570 (9).

Lindeman & Teunissen 15291 (9).

Lindsay 262 (28).

Loefgren & Duvall 36217 (33).

Ludwig 262 (21).

Luederwaldt 21547 (33).

Luetzelburg 254 (20); 6858 (34).

Lugo S. 3293 (7).

Lundell 6416 (9).

Macbride 5602 (7).

Macedo 1447 (10); 1521 (10); 2682 (10).

Madison et al. 3356 (2); 4851 (26).

Maguire & Fanshawe 22855 (9).

Maguire et al. 46068-A (6); 54384 (6).

Maurel & Maurel 3816 (28).

Maurice 699 (16).

Maxon 412 (16); 639 (2); 2293 (28);

2470 (28); 4641 (9); 6895 (9):
8978 (28); 9366 (28); 9464 (28);
9522 (28).

Maxon & Harvey 8242 (16).

Maxon & Killip 148 (28); 756 (28).

Maxon etal. 6817 (9).

McAlpin & Kuhn 77-12 (14).

Mendez 75 (9).

Mexia 6246a (6).

Mickel 1944 (9); 2003 (16); 2624 (28):
2742 (2); 2803 (28); 2817 (28); 2958 (16);
3126 (16); 3180 (2); 3368 (9); 3426 (16);
3570 (9); 9472 (9).

Molina R. 1907 (9).

Mora 4160 (18); 4287 (13); 4430 (9).

Moran 2167 (28); 2170 (2); 2171 (2);

2173 (2); 2176 (2); 2178 (2); 2182 (9);

2185 (9); 2186 (9); 2237 (16); 2241 (16):
2325 (16); 2441 (27); 2442 (16); 3145 (5):
3160 (27); 3168 (16); 3214 (16); 3241 (27);

a ee

———

November 1987

3271 (27);3338 (27);3512 (24); 3527 (25);
3528 (22); 3530 (2); 3532 (16); 3534 (9);
3535 (7); 3536 (6); 3544 (26); 3546.5 (3);
3547 (28); 3559 (26); 3561 (3); 3562 (2);
3563 (4); 3564 (26); 3565 (3); 3569 (2);
3570 (16); 3585 (24); 3586 (25); 3588 (9);
3590 (2); 3592 (28); 3593 (2); 3600 (28);
3612 (28); 3615 (7); 3616 (7); 3617 (9);
3618 (28); 3640 (7); 3641 (7); 3642 (28);
3663 (9); 3671 (9); 3688 (26); 3709 (1);
3718 (28).

Morely & Whitefoord 655 (28); 675 (28);
695 (28).

Mori & Kallunki 3574 (2).

Moritz 277 (1); 278 (32).

Morton 7629 (2).

Mosen 3050 (20).

Muller 6039 (20).

Murillo 1457 (5); 1491 (5); 2580 (28).

Murillo et al. 289 (5).

Nee & Stockwell 11610 (2).

Nee et al. 8768 (2).

Nelson et al. 3296 (2).

Niell 2610 (9).

Niemeyer 44 (28).

Ocampo 727 (2); 877 (16).

Oldeman 1933 (9).

Ollgaard et al. 34634 (9); 34703 (11);
34748 (7); 34855 (7); 34894 (11);
34970 (7); 35124 (6); 35290 (9);
35350 (11); 35776 (24); 37702 (26);
37820 (26); 37823 (3); 37859 (3);
37867 (26); 38836 (7); 38845 (28);
38932 (9); 38894 (28); 38969 (11);
39039 (11); 39040 (11); 39084 (7);
39086 (28); 43798 (9); 53824 (25).

Ortega 636 (9).

Ortega & Grimann 2707 (5).

Ortega & Smith 2387 (28); 2491 (1);
2498 (32); 2510 (1); 2511 (32).

Pabst et al. 6772 (34).
Perkins 1163 (28).
Persaud 114 (13); 372 (9).

Pipoly 3542 (9);3817 (9);3824 (9); 3880 (9);
3901 (9); 5146 (28); 5299 (9).

Pittier 587 (19); 4492 (9); 9016 (28);
9076 (28); 10688 (9); 12416 (28);
16232 (2).

Plowman & Davis 4542 (13).

Plowman & Thomas 13678 (28).

Plowman et al. 4025 (7).

Poeppig 201 (9).

Porter et al. 4763 (9).

MONOGRAPH of POLYBOTRA 135

Prance & Silva 59677 (10).

Pranceetal. 10466 (5); 12069 (9); 12180 (7);
15332 (31).

Prestoe 1491 (1); 1492 (1).

Proctor 3969 (28); 16506 (28); 22269 (28).

Proctor et al. 27091 (9).

Purpus 6761 (2); 7245 (2).

Ratter et al. 2047 (10).

Reitz 151 (20); 2825 (20).

Reitz & Klein 1101 (20); 1102 (20);

2665 (20); 3466 (20); 4635 (20);
4682 (20); 5108 (20); 5549 (20).

Richards 803 (9).

Richardson 2037 (28).

Riedel 81 (35).

Rimbach 31 (13); 91 (26); 312 (26).

Rivero 513A (1).

Rivero et al. 1608 (1).

Rojas 10182 (10); 10431 (10); 10804 (10).

Rose & Russell 20790 (34).

Rossbach 3624 (9); 3625 (9); 3628 (28);
3710 (9).

Rovirosa 972 (2).

Rusby 442 (11); 443 (22).

Sandwith 1561 (9).

Scamman 5984 (28); 5985 (16); 5986 (16);
5987 (28);7152 (28);7153 (28);7155 (16);
7156 (9); 7482 (9); 7687 (28).

Scamman & Holdridge 7998 (9).

Schipp 273 (9); 8101 (28); 8108 (28).

Schmalz 78 (20); 158 (20).

Schnee 615 (1).

Schneus 3412 (20).

Schomburgk 1659 (9).

Schultes & Black 46-266 (7); 8467 (9).

Schultes & Cabrera 13963 (31); 16053 (11).

Schunke, C. 157 (7); 158 (7); 164 (28);
165 (6); 380 (9); A214 (6); 661 (7); 666
(6); 705 (28); 812 (7); 1341 (28); 1395
(28); 1396 (6); 1451 (28).

Schunke, J. 268 (7); 275 (28); 380 (28);
5785 (28); 5789 (11); 10173 (6);

10200 (26).

Seaverns 56 (9).

Sehnem 3092 (20).

Seifriz 24 (3).

Shafer 4458 (28); 8889 (28).

Shemluck 280 (7); 304 (16).

Sherring 13 (28).

Silva et al. BGS526 (5).

Skog et al. 5036 (2).

Skutch 3018 (28); 3236 (16); 4637 (28).

136 ILLINOIS NATURAL HISTORY SURVEY

Smith, A.C. 48/293 (16); 778 (16);
2984 (31); 61713 (13).

Smith, A.R. 1259 (1); 4055 (1).
Smith, A.R. et al. 922 (1); 1347 (28);
1388 (28).

Smith, H.H. 983 (3); 1050 (28); 1052 (5).

Smith & Brade 2286 (20).

Smith & Klein 7546 (20).

Smith & Reitz 6135 (20).

Sodiro 81 (28).

Soejarto 2805 (28).

Sota, de la 2343 (34); 5181 (28); 5252 (16).

Soukup 1065 (7).

Sperling 5914 (9).

Sperling & Bleiweiss 5034 (26).

Sperling et al. 5915 (28).

Spruce 2116 (9); 3880 (11); 4090 (6);
4337 (6); 4634 (9); 4744 (13); 4740 (11);
5685 (28).

Standley 9047 (9); 24195 (9 & 28);

25085 (9); 27597 (9); 33648 (16);
37096 (28); 53955 (2); 53983 (9);
67135 (2); 68203 (2).

Standley & Valerio 47127 (28); 48622 (9).

Starry 91 (9).

Steeves & Ray 504 (28).

Steinbach 3032 (9); 7499 (9).

Stern et al. 523 (9); 1034 (16).

Stevens & Krukoff 6518 (16); 8827 (9);
12056 (9); 12666 (9); 12741 (9);

13005 (9).

Stewart 241 (28).

Steyermark 37320 (2); 38229 (9); 38271 (9);
39197 (9); 39895 (9); 41698a (2);
41870 (2); 44675 (9); 52812 (26);
56114 (1); 61991 (1); 62015 (28);
89120 (9); 91756 (1); 94952 (28);
99185 (32); 104717 (1); 107148 (28).

Steyermark & Davidse 116499 (28).

Steyermark & Liesner 120634 (28).

Steyermark & Nevling 95930 (28).

Steyermark & Rabe 71756 (28); 96145 (28);

97133 (1).

Steyermark & Stoddart 118051 (1).
Steyermark et al. 95827a (5); 101552 (28);
106758 (1); 114331 (9); 114332 (9);
115021 (9); 115033 (9); 122814 (28);
124749 (1); 125039 (28); 126675 (1);

126680 (1).

Vol. 34, Art. 1

Stolze 1488 (16).

Stork & Horton 9509 (9).

Stiibel 875 (28); 913a (13); 914 (13).

Sugden 1151 (28).

Swingle et al. 70-02-05-2 (9).

Sydow 339 (26).

Tate 422 (9); 502 (6); 513 (6).

Taylor 447 (28); 1223 (9).

Tonduz 9006 (28); 9016 (28); 13337 (28);
14568 (28); 18879 (28).

Tryon & Kramer 5611 (9).

Tryon& Tryon 5204 (9);5221 (6);6591 (20).

Tschudi 162 (5); 166 (32); 167 (1).

Tuerckheim 148 (9); 358 (9); 7812 (9);
8040B (9); 8041 (9).

Tyson 2109 (9).

Underwood 1246 (28); 1250 (28); 1323 (28);
1612 (28).

Valerio 63 (16); 328 (28); 329 (2); 333 (28);
2356 (16); 33133 (2).

van Cotthem 1327 (1).

van der Werff 3486 (32).

van der Werff & Gonzalez 5202 (28).

van der Werff & Wingfield 3430 (1).

Vareschi 3147 (28); 7764 (1).

Vareschi & Gessner 1875 (28).

Vareschi & Pannier 1686 (28); 2636 (5):
2660 (28); 2715 (28).

Vargas 11280 (6); 17743 (28); 17800 (9).

von Sneidern 1590 (19).

Wacket 21556 (20).

Walker T10995 (28).

Watt 160 (28); 7276 (28).

Webb 26 (34).

Webster et al. 16467 (2).

Weddell 952 (35).

Wercklé 559 (16); 16770 (2).

Wetmore & Woodworth 130 (9).

Whitmore 752 (7).

Wiggins 11073 (26).

Williams 1188 (6); 1739 (9); 2137 (9);
3136 (11); 3999 (9); 4797 (11);
7620 (11); 7681 (9).

Wilson & Murray 564 (28).

Wilson & Webster 549 (28).

Wingfield 6914 (28).

Wright 786 (28).

Wurdack 1854 (6); 1933 (11); 2011 (9).

November 1987 MONOGRAPH of POLYBOTRA

Index to Taxonomic Names

Accepted names are set in roman type; new taxa
and combinations appear in bold type; all other
names are italicized. A number after a name refers
to the number of the accepted taxon. Abbreviations
refer to sections of this monograph: ex (excluded
names), unc (names of uncertain application), sub
(subdivision of the genus).

Acrostichum
botryoides 17
caenopteris 1

var. salicifolium 5
canaliculatum 32
caudatum 9

var. pubens 11
chrysolepis 26
cylindricum 20
fractiseriale 6
hackelianum 3

hartii |
incisum 20
insigne 4

juglandifolium 2
lechlerianum 13
osmundaceum 28
plumbicaule 6
polybotryoides 2
pubens 11
suberectum 3
Aspidium
scandens 33
Botryothallus see Polybotrya
kunzei 1
Dryopteris
guentheri 11
Olfersia
caudata 9
cervina ex
Polybotrya
sect. Arthrobotrya ex
sect. Egenolfia ex
sect. Lomagramma_ ex
sect. Teratophyllum ex
subg. Ectoneura ex
subg. Egenolfia ex
subg. Polybotrya sub
subg. Sorbifolia sub
subg. Soromanes sub
acrostichoides ex
acuminata unc.
var. villosa 9

aequatoriana 24
alata 23
alfredii 16
forma carpinterae
altescandens 26
andina 4
apiifolia ex
appendiculata ex
appressa 25
arfakensis ex
aristeguietae 28
articulata ex
aspidioides ex
asplenifolia ex
attenuata 14
aucuparia 2
aurita ex
bifurcata ex
blumeana_ ex

botryoides 17
canaliculata 32
caudata 9

cervina ex
cicutaria eX
coenopteris |
costaricensis 9
crassa |
crassirhizoma 7
crespiana 11
cyathifolia 29
cylindrica 20
decorata 11
duplicato-serrata ex
espiritosantensis 8
exaltata ex
filiculifolia ex
flabellata ex
fractiserialis 6
fraxinifolia ex
frondosa 20
fulvostrigosa unc
furcata ex
gaudichaudiana ex
glandulosa 12
gomezii 27
goyazensis 10
gracilis 16
hamiltoniana ex
hartii |
helferiana ex
hickeyi 21
incisa 20

16

137

138

ILLINOIS NATURAL HIisTORY SURVEY

intermedia ex
juglandifolia 2
var. lobata 2
kalbreyeri 2
kunzei 1
latisquamosa 30
lechleriana 13
litoralis 34
lomarioides (Blume) Kuhn ex
lomarioides Mettenius unc
lourteigiana 18
macbridei 7
macedoi 9
marattioides ex
marginata ex
montana ex
nana ex
neglecta ex
nieuwenhuisenii ex
nodiflora ex
nutans unc
orientalis ex
osmundacea 28
var. crispopaleacea 34
var. cylindrica 20
var. frondosa 20
var. incisa 20

peltata ex
pilosa 35
pittieri 19

plumbicaulis 6
plumieri ex
polybotryoides 2
polyphylla_ ex
prolifera ex
pteroides ex
pubens 11
puberulenta 22
quercifolia ex

rhizophylla ex
rosenstockiana 34
salicifolia 5
scandens Fée unc
scandens (Raddi) Christ 33
semipinnata 33
serratifolia 1
serrulata ex
sessilisora 31
sinensis eX
sorbifolia Kuhn 5
sorbifolia (L.) Keys. ex
speciosa 34
stenosemioides ex
stolzei 15
subelliptica 12
suberecta 3
subquinquefida ex
taccaefolia ex
tenuifolia ex
teysmanniana eX
tomentosa 34
trapezoides unc
trilobata ex
tripartita ex
vareschii 28
villosula 9
vivipara eX
wilkesiana ex
Polypodium
adiantoides 9
Psomiocarpa
acuminata unc
caudata 9
Soromanes see Polybotrya
coenopteris |
dentatum 1
integrifolium 1
serratifolium 1

Vol. 34, Art. 1

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Send three copies of manuscripts to be considered for publication to Office of the Chief, Illinois
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linois Natural History Survey
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Division of the Illinois Department of Energy and Natural Resources

' Say (ie
FEB 1 1990

DEPOSITORY ARY
DEC 0 * 1989

UNIVERSITY UF ILLINOIS
AT URBANA-CHAMPAIGN

Aster and Brachyactis in Illinois

Almut G. Jones

Illinois Natural History Survey Bulletin
Volume 34. Article 2

Aster and Brachyactis in Illinois

Almut G. Jones
Department of Plant Biology
University of Illinois at Urbana-Champaign

Illinois Natural History Survey Bulletin
Volume 34, Article 2
May 1989

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

A catalog of the publications of the Illinois Natural History Survey is available without charge
from the address below. A price list and an order blank are included with the catalog.

Illinois Natural History Survey
Distribution Center

Natural Resources Building
607 East Peabody Drive
Champaign, Illinois 61820

Printing costs for this publication were paid for in part by the Department of Plant Biology, University
of Illinois at Urbana-Champaign.

The cover drawing was executed by Aleta Holt.

Editor: Audrey S. Hodgins

Citation:

Jones, A.G. 1989. Aster and Brachyactis in Ilinois. Illinois Natural History Survey Bulletin
34 (2):139-194.

US ISSN 0073-4918

Printed by Authority of the State of Illinois
(72724-2M-5-89)

Contents

Introduction 139
Generic Description of Aster L. — 143
Conspectus of Classification of Illinois Aster Species 144
Key to Species of Aster and Brachyactis in Illinois 145
Descriptions of Aster Species 149
Aster anomalus Engelm. in Torrey & Gray 149
Aster borealis (Torrey & Gray) Prov. 150
Aster cordifolius L. 151
Aster ciliolatus Lindley in Hooker — 152
Aster drummondii Lindley in Hooker — 153
Aster dumosus L. var. strictior Torrey & Gray 154
Aster ericoides L. — 155
Aster falcatus Lindley in Hooker var. commutatus (Torrey & Gray) A.G. Jones
Aster fragilis Willd. = 157
Aster furcatus Burgess in Britton & Brown 158
Aster laevisL. 159
Aster lanceolatus Willd. 160
Aster lateriflorus (L.) Britton 162
Aster linariifolius L. 163
Aster macrophyllus L. 164
Aster novae-angliaeL. 165
Aster x amethystinus Nutt. 166
Aster oblongifolius Nutt. 166
Aster ontarionis Wieg. — 167
Aster oolentangiensis Riddell 168
Aster parviceps (Burgess in Britton & Brown) Mack. & Bush 169
Aster patens Aiton 170
Aster pilosus Willd. — 171
Aster praealtus Poir. 173
Aster prenanthoides Muhl. ex Willd. — 174
Aster puniceus L. — 175
Aster schreberi Nees — 177
Aster sericeus Vent. 178
Aster shortii Lindley in Hooker 179
Aster tataricus L.f. 180
Aster turbinellus Lindley in Hooker 181
Aster umbellatus Miller 182
Aster undulatus L. 183
Aster urophyllus Lindley in DC. — 184
Description of Brachyactis Species 186
Brachyactis ciliata (Ledeb.) Ledeb. 186

Glossary of Descriptive Terms 187

Literature Cited 190

Index to Scientific Species Names, Including Synonyms — 193
Index to Vernacular (Common) Names 194

156

Kane

endal

Whiteside
Rock Island LaSalle rund
Putnam
[ers Kankakee
Warren} Knox Peoria Woodford se Iroquois

Cook
DuPage

Henderson

2

Greene |Macoupin
Calhoun Montgomery
(
Effingha
Marion

Se Washington

Randolpn| Perry

Hancock

Douglas } Edgar

Clay awrence

County reference map to be used with species distribution maps.

Aster and Brachyactis in Ulinois

Introduction

The need for a comprehensive study and
taxonomic revision of asters in Illinois
becomes apparent when one looks at the
varied treatments of this genus in the principal
floristic literature of the state (G.N. Jones
1945, 1950, 1963; Jones and Fuller 1955;
Mohlenbrock 1975, 1986; Mohlenbrock and
Ladd 1978). Three factors contribute to the
differences found in these works: the number
of taxa recorded for Illinois, nomenclatural
considerations such as those dictated by the
principle of priority, and differences in
taxonomic concepts and interpretations.

In the first edition of Flora of Illinois
(G.N. Jones 1945), 33 species are treated
under Aster, not counting hybrids. Three
additional species appear in the third edition
(G.N. Jones 1963): A. chasei G.N. Jones in
Jones & Fuller, a species newly described
from Illinois; A. tataricus L.f., an occasion-
ally escaped cultivated species; and A.
parviceps (Burgess in Britton & Brown)
Mack. & Bush, probably inadvertently
omitted from the first two editions but
included in Jones and Fuller (1955). In the
most recent Guide to the Vascular Flora of
Illinois, Mohlenbrock (1986) treats 35
species. A notable change involves the
transfer of A. ptarmicoides (Nees) Torrey &
Gray to Solidago. In this revision, I recognize
31 species of Aster in the state of Illinois.

My research has concentrated on New
World and some Old World species of Aster
(sensu lato) for over 15 years (A.G. Jones
1974, 1977, 1978a, b, c, 1980a, b, 1982, 1983,
1984, 1987; Jones and Hiepko 1981; Jones
and Young 1983; Jones and Lowry 1986; and
others), and much additional information has
been introduced in this current study. My

Almut G. Jones

taxonomic concepts are summarized in the
conspectus of classification of Illinois
Aster species (p. 144).

A recent example of a name change
necessitated by application of the principle
of priority is that of Aster azureus Lindley
in Hooker. The name is placed in synon-
ymy under A. oolentangiensis Riddell
(A.G. Jones 1983). Publication of the
latter name preceded that of the former by
only seven months. Other examples of
name changes are A. junciformis Rydb. to
A. borealis (Torrey & Gray) Prov., and
A. pantotrichus S.F. Blake to A. ontarionis
Wieg. (Shinners 1949).

Most differences among various
accounts are attributable to disagreements
about taxonomic concepts and interpreta-
tions. Changes may be the result of the
union of two or more species under one
[e.g., Aster simplex Willd. = A. lanceola-
tus Willd. (Semple 1979; Semple and
Chmielewski 1987)], or they may involve
elevation to the rank of species of taxa
formerly considered at an inferior rank
[e.g., A. ericoides L. (var.) parviceps
Burgess in Britton & Brown = A. parv-
iceps|. Sometimes names have been
misapplied [e.g., A. ericoides for A.
pilosus Willd. (cf. Blake 1930)], and
occasionally a species is transferred to
another genus [e.g., A. pfarmicoides to
Solidago (Boivin 1972) and A. brachyactis
S.F. Blake to Brachyactis (A.G. Jones
1984)].

The transfer of Aster ptarmicoides to
Solidago is now generally accepted.
Characters of habit, phyllaries, and
achenes link this species [called Stuff

140 ILtinois NATURAL History SurveY BULLETIN

Aster by Swink (1974)] to the Oligoneuron
group of Solidago. The genetic alliance is
further supported by the fact that hybridiza-
tion between A. ptarmicoides and species of
the above group is not uncommon. One such
hybrid has been reported from Cook County
(Higley and Raddin 1891; Pepoon 1927;
Jones and Fuller 1955; Swink 1974): S. x
lutescens (Lindley in DC.) Boivin. It was
first described as a species: Diplopappus
lutescens Lindley in DC. [synonyms: A.
lutescens (Lindley in DC.) Torrey & Gray;
A. ptarmicoides var. lutescens (Lindley in
DC.) A. Gray], and S. riddellii Frank is
probably the other parent species involved in
the hybrid.

Following my survey of the tribe
Astereae (A.G. Jones 1984, 1985), I
proposed returning Brachyactis Ledeb. to its
originally published status as a separate
genus. I have, however, included the single
Illinois species Brachyactis ciliata (Ledeb.)
Ledeb. in this study because it is considered
a member of Aster in nearly all floristic
literature to the present and because recent
students of the taxon disagree with my
viewpoint (Houle and Brouillet 1985).

Other recent treatments of Aster that
espouse generic concepts different from
mine have been published by Semple and
Brouillet (1980a, b) and Reveal and Keener
(1981). These treatments, however, which
propose segregation from Aster of species of
A. subg. Virgulus (Raf.) A.G. Jones (v =5
chromosomes), have not been widely
adopted. Evidence of hybridization between
species of that assemblage and members of
A. subg. Symphyotrichum (Nees) A.G. Jones
(x = 8 chromosomes) supports a more
conservative generic concept (A.G. Jones
1982; Jones and Young 1983; Allen 1985).

The following chapters are organized in
the format used in most floristic manuals.
The generic description of Aster (sensu
stricto) is followed by a conspectus of
classification of the Illinois species and by
an indented key that includes both Aster and
Brachyactis. Next, the species descriptions
are given in alphabetic order by scientific

Vol. 34 An. 2

name, the last being that of Brachyactis
ciliata.

Within each entry, vernacular names
used in literature dealing with the Illinois
flora are listed after the scientific name.

A new vernacular name is proposed (with
some reluctance) in this work for Aster
fragilis Willd.: Brittle Aster, a translation
of the very appropriate specific epithet.

As much as possible, a uniform format is
maintained in the descriptions. The diagno-
sis is followed by a listing of the mitotic
(2n) chromosome number(s) and by those
synonyms of the specific name that are cited
for Illinois in other floristic works, including
the bibliographic references.

A separate paragraph deals with phenol-
ogy, ecology, and general geographic
distribution. Standard two-letter postal
abbreviations are used for names of the
states and of the Canadian provinces, for
example, AB (Alberta), NF (Newfound-
land), PE (Prince Edward Island), PQ
(Quebec), and YT (Yukon Territory).

Dot maps show the distribution of native
Illinois species at the county level (county
names are identified on the preceding map).
These maps were based entirely on speci-
mens I have examined, annotated, and
recorded, and I may well have missed some
counties for which valid records have been
published. Previously published maps and
lists, however, may include occasional
recordings based on misidentifications. In
addition to my own collections, I have
examined more than 10,000 herbarium
specimens for this study, including all
material on deposit in the principal Illinois
herbaria: DEK, EIU, F, ILL, ILLS, ISM,
KNOX, MOR, MWI, and SIU. Abbrevia-
tions correspond to those in /ndex Herbari-
orum (Holmgren et al. 1981). GH, MIN,
MO, NY, PH, US, and WIS are among the
larger herbaria outside the state that yielded
distribution data for the asters of Illinois.
Some personal collections were borrowed
from P. Shildneck and J. Schwegman.

Varieties are treated following the
description of the respective species, either

May 1989

in a short diagnostic statement or, in cases
where a clear distinction by a combination
of characteristics is possible, in the form of a
key. Varietal synonymies are listed only to
the extent that they have been cited for or
described from Illinois. One reason for the
inconsistency in the handling of varieties is
that keys are not very meaningful or
practical when a high degree of intergrada-
tion or overlap in characteristics exists.
Moreover, I am convinced that some
commonly recognized varieties have
acquired their distinctive characteristics via
gene flow from other species.

Hybrids are mentioned at the end of each
entry, and some herbarium collections are
listed as examples. Although only a few of
the putative hybrids observed in the wild
have been experimentally reproduced, the
fact that closely related Aster species have a
high degree of genetic homology and do
occasionally hybridize is now generally
accepted (Wiegand 1928, 1932; Cronquist
1947; Semple and Brammall 1982; and
others). The classic example of an experi-
mentally confirmed hybrid that may survive
for several seasons is A. X amethystinus
Nutt., pro sp. = A. ericoides x A. novae-
angliae L. (Wetmore and Delisle 1939; A.G.
Jones 1978c). At least some viable achenes
are commonly produced, and backcrosses to
plants of the parent species have been
successful. Because of the relatively
frequent occurrence of this hybrid, I have
included it in the key and given a compara-
tive description after that of one of its parent
species (A. novae-angliae). Herbarium
specimens of putative hybrid individuals,
however, are not entirely uniform, and some
are undoubtedly backcrosses to one or the
other parent. Intergradation between two
species is more evident in some cases than
in others, and the key to the species gener-
ally does not work well for hybrids or for
individuals that exhibit a high degree of
introgression from other species.

Aside from the hybrid Aster x amethys-
tinus, | have included in the key and
discussed under their closest relatives two

ASTER AND BRACHYACTIS IN ILLINOIS 141

adventive species that may or may not be
established in Illinois: A. falcatus Lindley in
Hooker (under A. ericoides) and A. ciliola-
tus Lindley in Hooker (under A. cordifolius
L.). | have also included the exotic species
A. tataricus because it is treated in current
manuals dealing with the flora of Illinois
(Gleason 1952; Gleason and Cronquist
1963; G.N. Jones 1963; Mohlenbrock 1975,
1986).

Three species of Aster are included in
the list of threatened species of Illinois
issued by the Department of Conservation
on March 14, 1989: A. furcatus Burgess in
Britton & Brown, A. schreberi Nees, and
A. undulatus L. In my opinion, A. parviceps
should be considered for inclusion in the
list. The latter species and A. furcatus are
true midwestern endemics, and they may be
threatened because the plants grow in
vulnerable habitats. Although the number of
locality records is considerable and some
populations are extensive, these two species
are probably not as common today as they
used to be. I have been unsuccessful in my
search in many places where they had been
previously collected. The other two asters
currently listed as threatened may be rare in
Illinois, but they are very common in some
other areas. Illinois populations represent
the western limit of range for these species.
Aster schreberi presents a special situation
because Illinois and Wisconsin populations
are disjunct from the nearest populations to
the east and have been named and described
as A. chasei (Jones and Fuller 1955). So far
I have been unable, however, to find bio-
logical evidence that would support
taxonomic recognition of these disjuncts as
distinct from typical A. schreberi, even at
the varietal rank. Other species that ap-
proach their limit of range in Illinois and are
much more common outside the state are A.
borealis, A. dumosus L., A. macrophyllus
L., and A. prenanthoides Muhl. ex Willd.

To help resolve some of the difficulties
of species delimitation that have haunted
earlier accounts of Aster, | have made my
keys and descriptions more detailed than

142 ILtinois NATURAL History SuRVEY BULLETIN

those generally found in floristic works.
Many structural features that provide
significant taxonomic information are often
given short treatment because they are
seasonal (e.g., those of mature fruits) or are
not always available in herbarium specimens
(e.g., rhizomes and basal rosettes). Because I
have grown plants of all the species de-
scribed here and observed them through the
seasonal cycle, I have been able to charac-
terize all structural elements. In addition to
providing more complete descriptions of the
species, this treatment will allow, at least in
some cases, the detection of hybrids and
populations showing intergradation.

The taxonomic method is an ongoing
endeavor, and revisions are needed as more
and more is learned about the genetics and
natural history of the taxa. No doubt, the
definitive account of Aster in Illinois will
continue to elude us.

Vol. 34 An. 2

Acknowledgments

The loan of specimens and help received
from the curators and collection managers of
the following herbaria are acknowledged:
DEK, EIU, F, GH, ILLS, ISM, KNOX,
MIN, MO, MOR, MWI, NY, PH, SIU, US,
and WIS. Thanks for testing the key and
checking some records goes to P. Shildneck
(Decatur) and A.C. Koelling (Illinois State
Museum, Springfield). I appreciate the
constructive criticisms of T.M. Barkley
(Kansas State University, Manhattan) and
K.R. Robertson (Illinois Natural History
Survey, Champaign), who refereed my
manuscript. I am especially grateful to W-F.
Lamboy (University of Illinois at Urbana-
Champaign) for many good suggestions
regarding this study and to my assistant
Laurel L. McKee for help with questions on
and proofing of the manuscript.

May 1989

Generic Description of Aster L.

Starwort, Wild Aster, Michaelmas Daisy,
Frost-flower.

Herbaceous or somewhat suffruticose perennials
with caudiciform or short to long-creeping
stoloniform rhizome systems but also including
some taprooted annuals (all native Illinois asters
are perennial). Stems erect or decumbent, terete
in cross section, or slightly ridged at the upper
nodes from decurrent leaf bases, single to few at
any point of emergence or cespitose, variously
branched, glabrous to densely pubescent,
sometimes glandular. Leaves simple, alternate,
usually polymorphic, those of the basal rosettes
and the lower stem often petiolate, those of the
middle or upper stem and of the branches (rameal
leaves) commonly subsessile or sessile; blades
variously shaped, glabrous to densely pubescent,
sometimes glandular, the apex acute, acuminate,
attenuate, or obtuse to rounded, the margins
entire, crenate, or serrate, never lobed, the base
cuneate to rounded or cordate, sometimes
sheathing or auriculate-clasping. Capitulescence
paniculiform, corymbiform, racemiform, or
rarely spiciform. Heads on short to long
peduncles (sometimes subsessile), often secund
from leaf axils of the terminal branchlets.
Involucre cylindric, turbinate (often an artifact in
pressed and dried specimens), campanulate, or
hemispheric, the bracts (phyllaries) in several
series. Phyllaries imbricated and usually gradu-
ated, occasionally subequal in length or the outer
ones longer than the inner, commonly with a
well-delimited apical green areole and a scarious,
often indurate or keeled basal portion, or
sometimes with green bands extending along the
midrib to the base; phyllaries of the outer series
often largely herbaceous. Receptacle flat or
slightly convex, alveolate with blunt or sharp
(rarely bristle-tipped) teeth or merely pitted in
older, fruiting heads. Ray florets several to
many, rarely absent or vestigial, mostly in | but
sometimes in 2 or more series, pistillate and
fertile, the strap-shaped or oblong ligule portion
of the corollas white, pink, lavender, purple, or
blue, never yellow, the tube portion yellowish
green when fresh. Disk florets few to many,
perfect and fertile, the corollas regular, cylindric
or funnelform, usually with well-delimited tube
and limb portions, the limb S5-lobed, cream-
colored or yellow, often appearing brighter

ASTER AND BRACHYACTIS 1N ILLINOIS 143

yellow before anthesis because of the presence
of yellow pollen, commonly aging to pink or
purple after the pollen is shed, the tube usually
slender, yellow or greenish. Style branches of
the disk florets with attenuate, papillate, sterile
appendages that are longer than the stigmatic
lines. Pappus often simple, composed of a single
series of relatively uniform, barbellate, capillary
bristles, or if more complex, either with 2 series
of capillary bristles more or less uniform in
length but with those of the inner series
clavellately expanded toward the apex, or with
an additional (third) outer series of distinct, very
short, attenuate bristles (a feature conventionally
referred to as a “double pappus”). Achenes
plump or somewhat compressed, 2—S times
longer than wide, gray or straw-colored to purple
or brown, glabrous to variously pubescent, with
a species-specific, definite number (3-10) of
ribs, these sometimes obscure. Basic chromo-
some numbers: x = 5, 8, and 9.

Excluded are Aster brachyactis S.F. Blake
{= Brachyactis ciliata (Ledeb.) Ledeb.] and A.
ptarmicoides (Nees) Torrey & Gray [= Solidago
ptarmicoides (Nees) Boivin].

Aster can be distinguished from closely
related genera (tribe Astereae) only by a
combination of characteristics: sterile append-
ages of the style branches attenuate, longer than
the stigmatic lines; phyllaries in 3 or more
usually graduated series; rays never yellow;
principal pappus bristles more or less uniform in
length, not or scarcely overtopping the disk
corollas; achenes usually with a definite number
of ribs, not densely striated; flowering period
typically late summer and fall.

Approximately 200 species, worldwide,
mostly in the northern hemisphere, with the
center of distribution in temperate North
America. Taxonomic boundaries well defined in
most species but blurred in some, largely as a
result of interspecific hybridization, which is
sometimes compounded by polyploidy.
Moreover, the species exhibit a high degree of
phenotypic variability, and similar morphologies
may have evolved from different biological
origins. Because many questions regarding the
taxonomy of Asfer cannot be answered at this
time, the species concepts adopted in this
treatment are relatively conservative.

144 ILtinois NatuRAL History SurvEY BULLETIN Vol. 34 Art. 2

Conspectus of Classification of Illinois Aster Species

Aster L. sect. Turbinelli (Rydb.) A.G. Jones,
subgenus Aster stat. and comb. nov.*
section Biotia DC. ex Torrey & Gray A. turbinellus Lindley in Hooker
A. furcatus Burgess in Britton & subg. Virgulus (Raf.) A.G. Jones
Brown sect. Concolores Torrey & Gray
A. macrophyllus L. A. sericeus Vent.
A. schreberi Nees sect. Ericoidei (Nees) Torrey & Gray
sect. Macrocephali (Kitamura) A.G. A. ericoides L.
Jones A. falcatus Lindley in Hooker
A. tataricus Lf. sect. Oblongifolii (Rydb.) A.G. Jones
subg. Doellingeria (Nees) A. Gray A. novae-angliae L.
sect. Triplopappus (Torrey & Gray) A. oblongifolius Nutt.
A.G. Jones sect. Patentes Torrey & Gray
A. umbellatus Miller A. patens Aiton

subg. /anthe (Torrey & Gray) A. Gray
A. linariifolius L.
subg. Symphyotrichum (Nees) A.G. Jones
sect. Concinni (Nees) Torrey & Gray
A. laevis L.
A. oolentangiensis Riddell
sect. Cordifolii G. Don in Loud.
A. anomalus Engelm. in Torrey &
Gray
. ciliolatus Lindley in Hooker
. cordifolius L.
. drummondii Lindley in Hooker
. shortii Lindley in Hooker
_ undulatus L.
_ urophyllus Lindley in DC.
sect. Dumosi Torrey & Gray
A. dumosus L.
A. fragilis Willd.
A. lateriflorus (L.) Britton
A. ontarionis Wieg.
sect. Porteriani (Rydb.) A.G. Jones
A. parviceps (Burgess in Britton
& Brown) Mack. & Bush
A. pilosus Willd.
sect. Salicifolii Torrey & Gray
A. borealis (Torrey & Gray)
Prov.
A. lanceolatus Willd.
A. praealtus Poir.
A. prenanthoides Muhl. ex Willd.
A. puniceus L.

>>> > Dd Db

* Based on Aster (species group) Turbinelli Rydb., Flora of the Prairies and Plains, 803, 804. 1932.
Syn.: A. sect. Eucephalus subsect. Turbinelli (Rydb.) A-G.Jones (1980a). Type species: A. turbinellus
Lindley in Hooker.

May 1989

ASTER AND BRACHYACTIS IN ILLINOIS 145

Key to Species of Aster and Brachyactis in Illinois

1. Plants annual with a taproot; rays absent or vestigial; pappus conspicuous at anthesis,

COMSINCEADLY OVELLODPIN, Ce COLOLAS\..5.cecces<er<<ecosecseccancaneccacenen-ecesdenceceestnezareavere Brachyactis ciliata
1. Plants perennial (Illinois species); rays typically conspicuous; pappus relatively inconspicuous

at anthesis, not or barely overtopping the disk corollas [Asrer}].

2. Basal and lower cauline leaves long-petiolate, the blades well delimited, with mostly

cordate to truncate or sometimes rounded bases.

Capitulescence corymbiform or cymiform; median phyllaries mostly obtuse to
rounded at the apex; disk corollas with the apparent tubular portion distinctly longer
than the flared or abruptly expanded portion of the limb (including the lobes);
achenes fusiform, with 7—10 (rarely more) ribs.

3h

4.
4.

Plants conspicuously glandular in the capitulescence ............... 13. A. macrophyllus

Plants typically not glandular (occasionally with a few glands on the peduncles

and phyllaries).

5. Rosettes of basal leaves prominent, often numerous forming a dense ground
cover; blades of basal and larger cauline leaves broadly ovate to cordate,
glabrous or with sparse indument, the petioles mostly much longer than
CGD ad eS ier ease cece cn cece erases rae seca enced ace raned Soren coer nee DA ASCARI EDENL

5. Rosettes of basal leaves not prominent, relatively few; blades of basal and
larger cauline leaves oblong-ovate or lanceolate, harshly scabrous
above and densely hirsute below, the petioles usually not longer than
the BLADES Foe sec acesnccvecocascusisseesaesstcseevessasciessisaseetesteats ssi sseegeven(eatecerss 8. A. furcatus

Capitulescence variously paniculiform; median phyllaries acute, acuminate, or
attenuate, or if obtuse, the apex with a sharp point, never rounded; disk corollas
with the tube portion shorter than the expanded limb portion; achens oblong-
obovoid, with 4 or 5(6) ribs.

6.

Phyllaries with strongly squarrose or reflexed tips; heads relatively large with
(22)25-35 (or more) rays, the involucre hemispherical .................4.. 1. A. anomalus
Phyllaries with appressed or only slightly spreading tips; heads relatively

smaller with ca 10-25 rays, the involucre campanulate or cylindrical

(or turbinate in pressed specimens).

7. Upper stem densely hirtellous or puberulent, the indument uniformly
distributed around the stem; principal leaves copiously pubescent on both
surfaces (or the upper surface glabrous in A. shortii).

8. Cauline leaves sessile and clasping or with auriculate-clasping
POUOIOS ec tererscceassecssctreceasthacesspcsctcscerscieotrecetasnecessusareiesr aces 30. A. undulatus
8. Cauline leaves not clasping, the petioles slender or winged, often
somewhat decurrent but never clasping.
9. Leaves of the midstem entire or subentire; rays mostly (12)15—25;
heads 1.5—2.5 cm in diameter when the rays are fully extended or
PYESSCG MG ds scsasevavsscescusssusasctncueth ontetconsetucssdiassvessssinetstegcens 26. A. shortii
9. Leaves of the midstem sharply serrate or crenate-serrate; rays
mostly (8)10—15; heads 1—1.5(1.8) cm in diameter when the
rays are fully extended or pressed ..............:.sc02++8 4. A. drummondii
1h peer stem glabrous or puberulent to villous in thin decurrent lines;
principal leaves glabrous or variously pubescent.
10. Phyllaries of the 2nd and 3rd series inward with usually broad,
short, rhombic green areoles; leaves of the midstem entire or subentire.
11. Leaves of the midstem auriculate-clasping, glabrous,
DIAUCOUS occ cxsessesnaptencasres once 9. A. laevis
11. Leaves of the midstem not clasping, scabrous above and hirsute
DELOW, MOL CIAUGOUS) cratsccvanssvssassessoreoseossensieeies 17. A. oolentangiensis

146

ILLinois NaTuRAL History SURVEY BULLETIN Vol. 34 An. 2

10. Phyllaries of the 2nd and 3rd series inward with obovate or usually narrower,

lance-rhombic or oblanceolate to linear green areoles; leaves of the midstem variously

toothed (sometimes subentire in A. urophyllus).

12. Rays typically white (occasionally faintly lavender); phyllaries with a long-attenuate
apex, largely scarious, often only the midrib light green, or the green
arcoleswinear-ODlanCe Ola te sesretenccracsessceenereenscaneceenees ateattateret meet pereeetasee 31. A. urophyllus

12. Rays typically purple or lavender; phyllaries with variously shaped apices, the
green areoles well developed, obovate or lance-rhombic to oblanceolate.

13. Heads I—1.5 cm in diameter when the rays are fully extended or pressed: total
number of florets 16—30 (Illinois plants); apex of the phyllaries acute or
SOMELIMES SNOL-ACUMIIMN ALE... -.-.-20n0-scrvcrsanseevensacsaveeerecsedssaseaseasraeeaeoa O= Pie CORAM OMS
13. Heads 1.5—2.5 cm in diameter when the rays are fully extended or pressed:
total number of florets 30-50; apex of the phyllaries attenuate or narrowly
AGUIMIMALE se vacoe cher esececsoncee-cvsceneenevarcesssvestcedeceeaderosncencsaducsecesdenteneees 3a. A. ciliolatus
Basal and lower cauline leaves sessile to subpetiolate or if apparently petiolate, the blades
with a cuneate base or gradually narrowed, not well delimited from the petiolar portion.
14. Pappus with an outer whorl of very short bristles in addition to 1 or 2 series of much

longer (principal) capillary bristles; leaves more or less uniform in contour except for

the lowermost, which are usually bracteiform; basal rosettes absent.

15. Leaves rigid, l-nerved, lacking secondary nerves, ca 1—3.5 cm long and 0.4 cm or

less wide; stems mostly less than 60 cm tall ..........e:ccceseseeseeseeeeeeeeneees 12. A. linariifolius
15. Leaves flexible, with well-developed secondary nerves, ca 4—12 cm long and
1 cm or more wide; stems mostly more than 60 cm tall ....................2.-- 29. A. umbellatus

14. Pappus simple, the bristles typically in a single series and more or less uniform in length;
leaves usually variable in contour but the lowermost not bracteiform; basal rosettes
present at least during part of the growing season.
16. Involucres and peduncles, often also the leaves and upper stems, glandular.
17. Rays (40)50—-100; outer and median phyllaries narrowly acuminate to
long-attenuate, strongly squarrose; cauline leaves auriculate-
CAS PUN 8 so eaves acc esecstesevcxsscennsecavcstsacuqnedesasyaseecescerdeuresesvereersontere 14. A. novae-angliae
17. Rays 12-35; outer and median phyllaries obtuse or acute to short-acuminate,
appressed or squarrose; cauline leaves variously inserted.
18. Cauline leaves auriculate-clasping: phyllaries strongly graduated,
appressed or with slightly recurved tips ..............:.sessesseeseeseeseeseess 19. A. patens
18. Cauline leaves not or only slightly clasping, rounded at the base, not
auriculate; phyllaries only slightly or not at all graduated, strongly
squarrose 15. A. oblongifolius
16. Involucres, peduncles, leaves, and stems not glandular.
19. Achenes with 7-10 ribs, purple or brown at maturity; phyllaries pubescent on
the outer (abaxial) surface; stem pubescence uniformly distributed around
the stem, never in lines.
20. Leaves and phyllaries appressed silvery-silky on both surfaces: ovaries
anid ACHENES | BL ADTOUS ees -2e ccc xc eseeceenece se cnc cae secusecrcncoencucoceiesverasnaxedeMOn AO WORTGCLS
20. Leaves and phyllaries strigillose, hirsute, or cinereous-puberulent; ovaries
and achenes pubescent.
21. Principal leaves auriculate-clasping; capitulescence an open,
divaricately branched panicle; heads mostly on long, stiff peduncles,
not crowded and not secund; rays purple .............0.cceseeeee 19. A. patens
21. Principal leaves rounded at the base, only slightly or not at all
clasping; capitulescence commonly a racemiform panicle; heads
subsessile or on relatively short peduncles, often crowded and secund;
rays white or lavender.
22. Rays lavender; involucre hemispherical; pappus tawny or
TOSCO BEM eo ccecccencorcterene cance rceatniegneeecennxceneenecs 14a. A. X amethystinus

May 1989

22.

ASTER AND BRACHYACTIS IN ILLINOIS 147

Rays typically white; involucre cylindrical to campanulate; pappus white or cream-colored.

23:

23.

Heads commonly secund; involucres usually less than 4.5 mm high; rays 18 or

ME WCKe eed eseeteneccccutcsctes tak coeptne dateccsevasstatsscunesagsionaasotegssetasbuvsdsdesveateeDeaorerevecte 6. A. ericoides
Heads commonly not secund; involucres 5 mm high or more; rays 20 or
RIL OM OMe ccece ces sees raevacr ascarvascisenopienstiesessaesdvidivcsvccwspeeestaneess 6a. A. falcatus var. commutatus

19. Achenes with 3—5(6) ribs, variously colored; phyllaries mostly glabrous on the outer surface
(rarely puberulent in a few species); stem pubescence variously distributed.
24. Leaves clasping, often strongly auriculate.

24.

2a:

Leaves glaucous; stems glabrous or nearly so; phyllaries strongly graduated, firm,
appressed when fresh, the apical green areole usually broad, rhombic, shorter than
PE SCAaLOUS| DaSall MOrtiON secscsssavschsacsevasesscaseeseevcssasosssecesccunctestavies coder ceeppecinss 9. A. laevis
Leaves not glaucous; stems variously pubescent; phyllaries not at all or only weakly
graduated, often flexible and leaflike, at least somewhat spreading or sometimes
reflexed, the apical green areole longer than the scarious basal portion, or often the
outer phyllaries largely herbaceous.
26. Principal cauline leaves serrate to subentire, gradually tapered toward

the base; stem variously hispidulous, as well as hirsute, either over the entire

length and sometimes coarsely hispid toward the base, or often with only

sparse hispidulous indument in the upper portion of the stem, the lower

glabrous or nearly so (depending on the variety); phyllaries long-acuminate or

attenuate sind stbnsvucesteecsecvsvessatenssarvarstesstsscsvpssesersvessts Doe, Ay PUNICEUS
26. Principal cauline leaves sharply serrate, abruptly contracted below the

middle into an entire-margined, sometimes apparently subpetiolar basal

portion; stem hirsute or villous, not hispidulous; phyllaries acute or

SHON =A CUM ALE Hoes teetece cneseescestencvestsearrcesensctaesesrceteoverporverearne.n D2> Ay prenanthoides

Leaves not clasping.

Alf

Involucre (7)8—12 mm high; phyllaries in 6-9 series, strongly graduated, blunt-tipped
and rounded on the back, largely scarious, the green areoles small, oblong, in the
APICAlh /S=/'s POLL OMS cate teseoccnscesstetespecatstonsstcseesteoseiepetsseeestesessusseesense LO. “Ac, tunbinellus
Involucre 3—8(10) mm high; phyllaries in 3—6 series, graduated to varying
degrees, acute to attenuate or if obtuse, with a sharply pointed tip, not rounded
on the back, the green areoles of various sizes and shapes.
28. Basal leaves 3-4 times larger than the principal cauline leaves, the blades
10-30 cm long and 5-10 cm wide, gradually tapered into winged petioles
10-40 cm long; phyllaries with a broad green band along the midrib extending
ROE ASC R eee elececte cee tece te act ucctcecee teescnscysvotsoespessbecutveesacheectectteseaedativee 27. A. tataricus
28. Basal leaves (often withered and deciduous at flowering time) mostly not
larger than the principal cauline leaves, often much smaller; phyllaries usually
with well-defined green areoles.
29. Phyllaries and rameal leaves with a spinulose bristle at the apex; rhizome
system caudiciform lacking stoloniform strands; new shoots initiated at
the base of old stems or as root sprouts; achenes gray, the ribs obscure.
30. Disk florets 20 or more; involucre somewhat urceolate, (4)5—8 mm

PAS SUR eerste cist cd dascustacsses sosscassosiantsactvccsacestravecuease Wesctescsssescciee DOs Al PHOSUS
30. Disk florets 6-12; involucre cylindric or turbinate, 3-4 mm
DU BIA Ns pines eteter ene ose seve avons vasessaevouaseesavscenviesiusdevivacttsoxsvecesssue 18. A. parviceps

29. Phyllaries and rameal leaves often with a sharp callus point at the apex
but lacking a spinulose bristle; rhizome system creeping or if short-branched,
the new shoots at least in part initiated at the ends of stoloniform rhizome
strands; achenes variously colored at maturity, the ribs mostly prominent.
31. Reticulate tertiary venation usually conspicuous on the lower leaf
surface, the veinlets often brownish, the enclosed green areolae
isodiametric; rays purple or lavender ................cc0. 21. A. praealtus

148

ILtinois NATURAL History SuRVEY BULLETIN Vol. 34 Art. 2

Reticulate tertiary venation inconspicuous on the lower leaf surface, the veinlets rarely
brownish, or the enclosed areolae oblong; rays variously colored.
32. Ovaries and achenes glabrous, purple or brown at maturity, median phyllaries with mostly
broad, short, rhombic (rarely lance-rhombic) green areoles; leaves scabrous above and
WUTSUte Delo Wie ree eee eee ere ace e rae u gn noosa se oan ee erent en Pan ESTES 17. A. oolentangiensis
Ovaries and achenes strigillose or puberulent, variously colored at maturity; median
phyllaries with linear, oblanceolate, or rhombic-obovate green areoles; leaves variously
pubescent or glabrous.
33. Lobes of the disk corollas longer than or as long as the fused portion of the limb
(not including the corolla tube); lower leaf surface at least somewhat pubescent;
achenes gray at maturity.
34. Lower leaf surface uniformly short-pubescent; heads not secund; rhizomes
distinctly long-creeping, stoloniform, the plants forming colonial stands,
DSHEVI Nab evs oo) Crfea Co) NY Leo ceca oreceecoccenn ceccer ceccec cence tear nee eceecet ce reser 16. A. ontarionis
34. Lower leaf surface villous to hirtellous along the midrib, otherwise usually
glabrous; heads secund; rhizomes short, sometimes tangled but not
long-creeping, the plants forming scattered individual clumps, mostly in
Upland (Sittia tions ee sesesies.ceseeess hasta cocoa tees eeseateee coon oes emeeentes 11. A. lateriflorus
Lobes of the disk corollas shorter than the fused portion of the limb; lower leaf
surface usually glabrous (rarely with a few trichomes along the midrib or slightly
scabrous toward the margins); achenes variously colored.
35. Flowering heads mostly 1.5—2.5 cm in diameter when the rays are fully
extended or pressed.
36. Larger cauline leaves 1—3.5 cm in width, no more than 12 times longer
than wide, usually sharply serrate; capitulescence a diffusely branched or
elongate panicle:;aCheMmes) Ora y weerececenceseeeennerereeeeceeeneeeeneeeneeees 10. A. lanceolatus
36. Larger cauline leaves mostly less than 1 cm in width, 12-15 (or more)
times longer than wide, shallowly and remotely serrate to subentire:
capitulescence a more or less dichotomously branched round- or flat-topped
panicle; achenes purple, or gray with purple streaks ................... 2. A. borealis
Flowering heads mostly 0.6—1.5 cm in diameter when the rays are fully
extended or pressed.
37. Median phyllaries (3rd or 4th series inward) ca twice as wide as those of
the outer series, typically obtuse, the green areoles rhombic-obovate to
broadly oblanceolate; rameal leaves relatively uniform in size; peduncles
usually at least | cm long (often much longer) with many bracteiform
leaves; subsessile heads relatively few .............::sscsseseseeeeseeeee J. A. dumosus
37. Median phyllaries slender, less than twice as wide as those of the
outer series, acute or attenuate, the green areoles linear to narrowly
oblanceolate; rameal leaves notably unequal in size; peduncles variable
in length but the bracteiform leaves typically few; subsessile heads often
MUMELOUSeccececeerconveaceccecueeereeceseoreees 7. A. fragilis

Ww
i)

Ww
ies)

es)
nn

May 1989

Description of Aster Species

1. Aster anomalus Engelm. in Torrey & Gray
Many-rayed Aster, Blue
Aster

Herbaceous, with stout
branched-caudiciform
rhizomes that turn
woody with age. New
shoots arising at or near the
base of old stems but also
sometimes initiated as root
sprouts, the plants forming
scattered individual clumps. Stems
l-several, erect or ascending, stout,
50-120 cm tall, with ascending or spreading
branches mostly from above the middle,
uniformly hirtellous or hirsute. Leaves polymor-
phic, the basal and larger cauline ones petiolate,
those higher up on the stem gradually reduced in
size, changing from subpetiolate to sessile; first
leaves of new shoots and of the vernal rosettes
with oblong-ovate to lanceolate blades 14 cm
long and 1—2 cm wide, usually purple at least on
the lower surface, densely hirtellous, acute or
often with obtuse to rounded tips, the margins
few-toothed or subentire; principal cauline leaves
with ovate to broadly lanceolate blades 4-9 cm
long and (1.5)2.5—5 cm wide, with several pairs
of usually arching and anastomosing secondary
nerves, hirtellous or hirsute on both surfaces, the
apex acute or acuminate with a short callus point,
the margins mosily entire to subentire (rarely
crenate-serrate), the base cordate, usually with
broad sinuses (rarely truncate), the petioles
(2)4-8 cm long, slender on the lower leaves,
narrowly winged on those of the midstem,
densely hirsute to pilose; upper leaves of the
main stem lanceolate, entire; rameal leaves
abruptly reduced, those of the peduncles
bracteiform, 2-6 mm long, mucronulate,
intergrading with the phyllaries. Capitulescence
a usually ample diffuse panicle, the head-bearing
branchlets sometimes racemiform. Flowering

ASTER AND BRACHYACTIS IN ILLINOIS 149

heads 2—3 cm in diameter when the rays are
fully extended, on peduncles 0.3—5(10) cm long
or rarely subsessile. Involucre hemispherical,
5-10 mm high, the phyllaries imbricated in
4-6(7) series. Phyllaries graduated, slender,
linear-lanceolate, strongly reflexed or squarrose,
grayish green, both surfaces typically hirtellous,
the apex long-attenuate, often with purple tips
and with a conspicuous bristly mucro, the
margins scabrous; outer phyllaries 3-4 mm long,
ca half as long as the inner, commonly herba-
ceous to near the base; median and inner
phyllaries green in the apical '/>—’/; portion, the
basal portion scarious. Receptacle alveolate with
sharp or bluntish teeth. Ray florets (22)25—35
(or more), the corollas 10—15(18) mm long,
purple or blue (rarely white), glabrous. Disk
florets 20-35 (or more), the corollas (4)4.5—5.5
mm long, glabrous or with a few trichomes, the
limb only slightly expanded, cream-colored or
light yellow turning purple after anthesis, the
lobe/limb fraction 0.2—0.25, the tube shorter than
the limb. Pappus simple, the bristles as long as
the disk corollas or slightly shorter, tawny or
rose-tinged, slender, attenuate. Achenes oblong-
obovoid, somewhat compressed, 2.5—3.5(3.8)
mm long and 0.8—1.2 mm across, deep purple, or
brown with purple speckles, glabrous, with 5 or
6 often irregularly spaced straw-colored ribs. 2
= 16; reported chromosome counts of 2 = 18
are probably in error (A.G. Jones 1977, 1980b).

Late August—October. Sandy or loamy clay
soils underlain by calcareous rocks or sandstone:
rocky open woods, woody hilltops, and dry
ridges. Occasional near and along major rivers in
the western half of the state, north to Peoria and
Woodford counties. The range for the species
includes MO, AR, s.e. KS, and e. OK.

There is good evidence in Illinois of
intergradation between this species and Aster
shortii [e.g., Winterringer 11997 (ISM) from
Alexander Co., Rexroat 8084 (ISM) from
Menard Co., and A.G. Jones 6817 (ILL) from
Union Co.}.

150 ILLinois NATURAL History SurvEY BULLETIN

2. Aster borealis (Torrey & Gray) Prov.
Rush Aster

Herbaceous, with very
slender, creeping,
stoloniform rhizomes.
New shoots mostly
initiated from the rhizomes,
the plants forming clonal
stands usually interspersed
with other vegetation. Stems
1—few at any point of emergence,
slender, erect, 30—80(100) cm tall,
often somewhat zigzag above,
branched only in the capitulescence, sometimes
red-tinged to slightly glaucous, nearly glabrous or
sparsely puberulent in thin decurrent lines (rarely
with uniformly distributed indument on the
peduncles). Leaves polymorphic, the basal and
lower cauline ones commonly withered and
deciduous at flowering time; basal rosette leaves
subpetiolate, the blades variously linear-
oblanceolate to ovate or suborbiculate, 0.5—1.5
cm long and 2-6 mm wide, the secondary
venation obscure, both surfaces glabrous or
minutely puberulent toward the margins, the
lower surface also usually purple-tinged, the apex
obtuse or rounded, the margins shallowly crenate,
abruptly narrowed to a slightly winged subpetio-
lar portion of variable length, the base dilated and
sheathing; principal cauline leaves very slender,
linear to linear-oblanceolate, (4)6—15 cm long
and 2—6(8) mm wide, with usually only the
midrib strongly expressed, glabrous or the upper
surface minutely strigillose toward the margins
and the lower with a few trichomes along the
midrib, the apex attenuate and callus-pointed, the
margins usually inrolled, entire or remotely and
shallowly serrate, smooth or scabrous, the base
sessile and somewhat decurrent; rameal leaves
relatively few, ascending or spreading, firm or
flexible, entire, similar to the cauline leaves in
contour and vestiture, the base somewhat
sheathing and decurrent; leaves of the peduncles
flexible, not phyllarylike, linear-lanceolate, 4-12
mm long, often red-tinged, and with a slightly
dilated or rounded base. Capitulescence an open,
round- or flat-topped, more or less dichotomously
branched, terminal panicle, rarely also with
ascending lateral branches initiated in leaf axils

Vol. 34 An. 2

of the midstem. Flowering heads (1.5)2—2.5 cm
in diameter when the rays are fully extended, not
crowded and not secund, typically on slender
puberulent or glabrous peduncles 1—5 cm in
length. Involucre broadly campanulate, (5.5)6—7
mm high, the phyllaries imbricated in 4-6 series.
Phyllaries appressed when fresh or rarely the
outermost slightly recurved, subequal or
somewhat graduated; outer phyllaries oblong-
lanceolate, 3—5 mm long, the innermost slender.
linear, 5-6 mm long, the green areoles changing
from slenderly oblanceolate to nearly linear, the
surfaces glabrous or nearly so, the apex acute,
callus-pointed in the outer phyllaries, often red-
tipped in the median and inner ones, the margins
irregularly ciliolate (rarely villous), or erose-
hyaline and with a scarious or reddish rim in the
median and inner phyllaries, the basal '/2—/:
portion scarious. Receptacle shallowly alveolate
with sharp teeth. Ray florets 20—30 (or more),
the corollas 10-12 mm long, mostly white,
sometimes pink or bluish, glabrous or with a few
trichomes at the throat and on the tube. Disk
florets 25—30 (or more), the corollas funnelform,
5-6 mm long, glabrous or nearly so, the limb
abruptly expanded at the throat, cream-colored or
pale yellow turning pink after anthesis, the
lobe/limb fraction 0.2, the tube shorter than the
limb. Pappus simple, the bristles approximately
as long as the disk corolla, whitish, soft, slender,
and attenuate. Achenes oblong or oblanceolate in
contour or often somewhat falcate, plump or
slightly compressed, (2.2)2.5—3 mm long, 4-5
times longer than wide, purple, or gray with
purple streaks, thinly strigillose, with 4 or 5
straw-colored ribs. 2n = 16, 32 (mostly), 48.

[A. junceus misapplied, not of Aiton—Brendel
1887; Pepoon 1927; Deam 1940. A. junciformis
Rydb.—G.N. Jones 1945, 1950, 1963; Fernald
1950; Gleason 1952; Jones and Fuller 1955;
Gleason and Cronquist 1963; Swink 1974;
Mohlenbrock 1975; Swink and Wilhelm 1979.}

Late August—October. Wet ground: tamarack
bogs, sedge meadows, grassy lakeshores, and
calcareous fens. Occasional in the counties
including and surrounding Chicago, now
probably rare because of destruction of the
habitat. The species range is still to be accurately
mapped. I have been unable, for example, to
locate the specimen cited for Winnebago County
by Fell (1955: 144) under the name Aster

Junciformis; the descriptive statement suggests

that the plant (found “in boggy places in Coon

May 1989

Creek bottom”) may belong here. According to
the floristic literature, the species has been
recorded from Anticosti Island, NB, and PQ,
westward through ON, MB, SK, s. NT, and s. YT
to BC and s. AK, southward to areas surrounding
the Great Lakes in NY, PA, OH, and IN, and also
including stations inn. IA, ND, SD, n. NE (?),
and MT. Excluded are some areas listed in the
literature for which the recorded specimens, in
my judgment, belong in other species.

The plants may be confused with those of
some other species, e.g., Aster dumosus var.
strictior Torrey & Gray, but the latter taxon is
distinct by having smaller heads and a strict,
thyrsiform capitulescence. The characteristics of
A. borealis suggest a close relationship in the
eastern half of the continent to A. /anceolatus and
A. longifolius Lam., and in the western half to
A. eatonii (A. Gray) Howell and A. occidentalis
(Nutt.) Torrey & Gray. Of these four species only
A. lanceolatus is an element of the Illinois flora,
and good evidence exists of intergradation
between that species and A. borealis [e.g., Benke
5278 (F) from McHenry Co.].

3. Aster cordifolius L.
Heart-leaved Aster, Blue
Wood Aster

Herbaceous, with a
branched-caudiciform or
creeping horizontal
rhizome system (depend-
ing on the variety). New
shoots originating at the base
of old stems or from rhizome
strands, the plants forming
scattered individual clumps or
small colonial patches. Stems
l-several, erect, (40)60—120 cm tall, often
reddish or maroon-colored, commonly somewhat
zigzag above, with ascending or divaricate
branches from above the middle, typically
puberulent in decurrent lines from the upper
nodes, in the leaf axils, and on the peduncles,
glabrous in the lower stem portion (sometimes
entirely glabrous). Leaves polymorphic, the basal
and principal cauline ones petiolate, those higher
up on the stem gradually reduced in size,

Pe

mae
ee cor
ST felelet © |

ASTER AND BRACHYACTIS IN ILLINOIS 151

changing to subsessile or sessile in the branches;
leaves of new shoots and of the vernal rosettes
with ovate-cordate to suborbiculate blades 1-8
cm (or more) long and 1—3 cm (or more) wide,
similar to the cauline leaves in venation and
vestiture, purplish below at least during the cool
season, the apex acuminate or acute (sometimes
obtuse or rounded), the margins coarsely and
often irregularly crenate-serrate, the base cordate
(rarely rounded), the petioles 1—3 times as long
as the blades, ciliate, slender or often winged,
dilated toward the sheathing base; blades of
principal cauline leaves ovate in contour,
4-10(13) cm long and 2—4(6) cm wide, with
several pairs of arching and anastomosing
secondary nerves, and a weakly expressed
reticulum of tertiary veins with irregular areolae,
the upper surface sparsely scabrous or glabrous,
the lower thinly villous or hirtellous along the
major nerves (rarely over the entire surface) or
nearly glabrous, the apex acuminate and
callus-pointed, the margins sharply serrate with
acuminate callus-pointed teeth, the base deeply
cordate or truncate to rounded, the petioles
progressively shorter and more broadly winged
upward on the stem; rameal leaves subsessile to
sessile, ovate-lanceolate, nearly glabrous or
minutely puberulent above, acuminate and
callus-pointed, the margins entire or commonly
somewhat toothed and ciliolate; leaves of the
peduncles densely spaced, 1.5—3 mm long, the
lower ones flexible, the upper bracteiform and
intergrading with the phyllaries. Capitulescence
a leafy, divaricately much-branched, usually
round-topped panicle in the upper '/:—'/> portion
of the stem, the head-bearing branchlets rarely
racemiform. Flowering heads |—1.5 cm in
diameter when the rays are fully extended, often
crowded, usually not secund, on densely
bracteate peduncles 0.3—2 cm long. Involucre
narrowly campanulate (sometimes turbinate
when pressed and dried), 3.5—5(6) mm high, the
phyllaries imbricated in (3)4 or 5 series.
Phyllaries strongly graduated, the outer ones
lanceolate, (1)1.5—2 mm long, the inner slender,
linear-oblanceolate, 3.5—4.5(5) mm long,
appressed when fresh, glabrous, the green
areoles rhombic to obovate and well delimited in
the apical '/:—'/» portion, commonly red-tipped,
the apex typically acute or obtuse (rarely
acuminate) in the outer and median, acuminate to
short-attenuate in the inner phyllaries, the

152 ILtinois NATURAL History SURVEY BULLETIN

margins erose-hyaline, irregularly ciliolate or
thinly villous toward the apex, the basal portion
scarious, often somewhat rounded on the back
and indurate. Receptacle alveolate with sharp
teeth. Ray florets (8)10—15, the corollas 7-10
mm long, typically purple (rarely pink), glabrous
or nearly so. Disk florets (8)10—12, the corollas
4-4.5(5) mm long, funnelform, glabrous or with a
few trichomes at the throat, the limb abruptly
dilated, cream-colored or light yellow turning
purple after anthesis, the lobe/limb fraction
().2-0.25, the tube slightly shorter than the limb.
Pappus simple, the bristles as long as or slightly
shorter than the disk corolla, whitish or faintly
rose-tinged, soft, slender, and attenuate. Achenes
oblong-obovoid or often slightly falcate,
somewhat compressed, 2—2.5 mm long and
0.6-0.8 mm across, dull purple or light brown,
glabrous, with 4 or 5 ribs. 2n = 16, 32 (often with
1 or 2 B-chromosomes); reported chromosome
counts of 2n = 36 are probably in error (A.G.
Jones 1977). Including A. sagittifolius Wedem.
ex Willd. (sensu stricto—A.G. Jones 1980b,
1987; Jones and Hiepko 1981).

September—October. Loamy or rocky soils in
mesic, mostly wooded habitats: open-wooded
slopes and bluffs, woodland edges, also in
somewhat disturbed ground at road cuts and
along ditches. Occasional throughout the state but
more common in the northern part. The range of
this variable species extends from PE, PQ, NB,
NS, and ME south to GA and AL, westward to
MN, IA, e. NE, MO, and AR.

Two varieties can be distinguished in Illinois,
typical var. cordifolius and Aster cordifolius var.
sagittifolius (Wedem. ex Willd.) A.G. Jones [A.
sagittifolius of authors only in part (see also A.
drummondii and A. urophyllus), A. cordifolius
subsp. sagittifolius (Wedem. ex Willd.) A.G.
Jones (1980b)]:

1. Rhizomes notably creeping, with slender
stoloniform strands, the plants colonial;
leaves of the midstem with slender or
narrowly winged petioles, the blades with a
deeply cordate base ............... var. cordifolius

Vol. 34 Art. 2

1. Rhizomes short, branched-caudiciform,
the plants forming scattered individual
clumps; leaves of the midstem with
broadly winged petioles, the blades
with a truncate or shallowly cordate
Loy Ore ero Tas ceo cre cere recor var. sagittifolius

Plants of var. sagittifolius exhibit traits that
may have been acquired via gene flow from other
species, such as Aster ciliolatus and A. drum-
mondii. If underground parts are missing,
specimens of this taxon can be distinguished only
with difficulty from those of typical var.
cordifolius. There is evidence of intergradation
between A. cordifolius and A. drummondii [e.g.,
Henry 2941, 2943 (MWI) from Adams Co. and
Chase 14906 (ILL) from Peoria Co.], A. laevis
[e.g., Chase 3728 (ILL) from Tazewell Co.], and
A. urophyllus [e.g., Evers 58628 (ILLS) from

from Union Co.].

3a. Aster ciliolatus Lindley in Hooker
Lindley’s Aster

I have included this species in the key on the
strength of three herbarium specimens collected
in Cook County and located at F [R. Bebb 1570,
Beverly Hills; F. Gates 43, Glencoe; Raddin s.n.,
n. Evanston]. The plants more or less exhibit the
traits attributed to Aster ciliolatus: stems glabrous
or nearly so; principal leaves glabrous or sparsely
ciliate below along the major nerves; peduncles
with no or few bracteiform rameal leaves: heads
relatively large, the involucres 6-7 mm high:
phyllaries lanceolate, long-attenuate, with lance-
rhombic green areoles and indurate, scarious or
somewhat discolored bases. In fact, the Gates
collection was annotated by Sherff as A.
lindleyanus Torrey & Gray, a synonym of

A. ciliolatus. Field study is needed to ascertain
whether this transcontinental boreal species is
really an established element of the Illinois flora.
The plants may be waifs, or they may be part of a
hybrid swarm involving A. cordifolius and A.
ciliolatus (or some other species, e.g., A. laevis).

May 1989

4. Aster drummondii Lindley in Hooker
Drummond’s Aster

Herbaceous, with stout
branched-caudiciform
rhizomes that often turn
woody with age. New
shoots originating at or near
the base of old stems, the
plants forming scattered
individual clumps. Stems
I-several, erect, stout, 60-120 cm
tall, with ascending or divaricate
branches mostly from above the middle,
uniformly soft-hirtellous in the branches, coarsely
hirsute on the main stem, the indument often in
lines, sometimes glabrescent in the lower portion.
Leaves polymorphic, the basal and lower cauline
ones petiolate, those higher up on the stem
gradually or abruptly reduced in size; leaves of
new shoots and vernal rosettes with ovate-oblong
to suborbiculate blades 1—-6(10) cm long and
1—3(5) cm wide, purplish below at least during
the cool season, copiously soft-pubescent on both
surfaces, the apex acute, obtuse, or rounded, the
margins crenate-serrate; principal cauline leaves
with broadly ovate to ovate-lanceolate blades
6—12(15) cm long and 3—S(6) cm wide, with 5—12
pairs of arching and anastomosing secondary
nerves, thinly soft-pubescent to scabrous or rarely
glabrous above, copiously hirtellous to hirsute
below, the apex acuminate to attenuate and
usually callus-pointed, the margins crenate-
serrate to sharply (sometimes doubly) serrate, the
base cordate or truncate, sometimes oblique;
petioles (2)4-8 cm long, usually conspicuously
winged and with ciliate margins; upper leaves of
the main stem lanceolate, subsessile; rameal
leaves oblong-lanceolate, sessile, typically
pubescent, callus-pointed, shallowly toothed to
entire and ciliolate; leaves of the peduncles
bracteiform, 2-5 mm long, intergrading with the
phyllaries. Capitulescence a usually ample
panicle, distinctly overtopping the leafy stem
portion, the terminal branchlets often
racemiform. Flowering heads |—1.5(1.8) cm in
diameter when the rays are fully extended, on
densely bracteate peduncles 0.24 cm long, or

ASTER AND BRACHYACTIS IN ILLINOIS 153

sometimes subsessile and secund. Involucre
campanulate (or turbinate when pressed and
dried), (3.5)4-6 mm high, the phyllaries
imbricated in (3)4 or 5 series. Phyllaries
strongly graduated, the outer ones lanceolate

or subulate, 1.5—2 mm long, the inner
linear-lanceolate, ca three times as long as the
outer, appressed or slightly spreading, typically
glabrous but occasionally puberulent on the
abaxial surface and thinly puberulent on the
adaxial surface, the apical green areoles rhombic
to oblong or oblanceolate, the apex acuminate or
attenuate, often red-tipped, the margins irregu-
larly ciliolate toward the apex, and with a
scarious rim extending to the tapered portion of
the tip, the basal '/; —'/2 portion scarious except
for the often green midrib. Receptacle shallowly
alveolate with sharp or blunt teeth. Ray florets
(8)10—15 (or more), the corollas 7-12 mm long,
purple or lavender (sometimes white), glabrous.
Disk florets (10)13—15 (or more), the corollas
3.5-4.5(5) mm long, funnelform, glabrous, the
limb abruptly dilated at the throat, cream-colored
or light yellow turning deep purple after anthesis,
the lobe/limb fraction 0.18—0.25, the tube
slender, slightly shorter than the limb. Pappus
simple, the bristles subequal, slightly shorter
than the disk corolla, whitish or rose-tinged, soft,
slender, and attenuate. Achenes oblong-obovoid,
often slightly falcate, somewhat compressed,
(1.5)2—3 mm long and 0.6—1 mm across, dull
purple or brown, glabrous or often sparsely
puberulent in the top portion, with 4 or 5 ribs.

2n = 16, 32 (mostly); reported chromosome
counts of 2” = 36 are probably in error (A.G.
Jones 1977). [A. sagittifolius Wedem. ex Willd.
var. drummondii (Lindley in Hooker) Shinners—
Fernald 1950; Kibbe 1952; Dobbs 1963; Swink
1974; Mohlenbrock 1975; Swink and Wilhelm
1979. A. undulatus misapplied, not of L.—Mead
1846; Higley and Raddin 1891; Pepoon 1927;
Kibbe 1952; Dobbs 1963.]

August—October. Loamy or rocky soils in
mesic, mostly wooded habitats: stream banks,
open-wooded slopes, thickets and edges of
swamps; also sometimes in disturbed ground
along roads and ditches. Common throughout
the state. The range of this variable species
extends from s. MN east to c. OH, south to w.
MS, n. LA, and e. TX, and west to IA, e. NE, e.
KS, and e. OK.

154 ILtinois NaturAL History Survey BULLETIN

Plants in Illinois belong mostly in typical var.
drummondii, including Aster drummondii var.
rhodactis Benke, described from Cook County
[Benke 4530 (F)| and characterized by
rose-colored rays and “rufescent” leaves and
stems. Individuals exhibiting phyllaries that are
puberulent on the back may fall within the limits
of A. finkti Rydb., a species (or more likely a
hybrid) described from Iowa and recognized by
Shinners (1941) as an endemic of the driftless
area of s.w. WI. I have recorded collections of
these plants from 16 counties throughout the state
and did not detect geographic or ecological
patterns or any consistent correlation of this trait
with other characters. Occasional hybridization
with and gene flow from A. shortii or A. undula-
tus (or both) may in part account for the occur-
rence of puberulent phyllaries in A. drummondii.
I find it impossible, therefore, to give taxonomic
recognition to this morphological variant. There
is also strong evidence of intergradation between
A. drummondii and A. urophyllus |e.g., Evers
28089 from Jo Daviess Co., 28300 from Pope
Co., 71937 from Adams Co., and 77383 from Lee
Co. (all at ILLS)].

5. Aster dumosus L. var. strictior Torrey & Gray
Bushy Aster, Rice-button
Aster

Herbaceous, with a
creeping stoloniform
rhizome system. New
shoots mostly initiated
from the rhizomes or
sometimes at the base of old
stems, the plants forming
scattered individual clumps or
small colonial patches. Stems
l-few at any point of emergence,
slender, erect, 30—70(100) cm tall, terete, the
branches commonly ascending (this variety),
sometimes divaricate, slightly ridged from
decurrent leaf bases, sparsely puberulent in lines
or nearly glabrous, the indument more copious on
the peduncles and sometimes uniformly
distributed. Leaves somewhat polymorphic, the
basal and larger ones mostly withered and
deciduous at flowering time; basal rosette leaves
spatulate, 1-5 cm long, glabrous or minutely
scabrous above, usually purple-tinged below,

Vol. 34 Ant. 2

crenate-serrate, abruptly narrowed to a coarsely
ciliate subpetiolar portion, dilated and sheathing
at the base; principal cauline leaves linear to
linear-oblanceolate, 3—10 cm long and 2—3(5)
mm wide, with (usually) only the midrib strongly
expressed, scabrous or glabrescent above,
glabrous below, the apex acute with a sharp
callus point, the margins usually inrolled,
scabrous, shallowly serrulate or rarely entire, the
base sessile and often somewhat decurrent but
not clasping: rameal leaves numerous, bractei-
form, linear-oblong, rather uniform in size at
each branch level and with relatively few clusters
of smaller leaves in the axils, ascending or
spreading, scabrous-margined, and callus-
pointed, those of the peduncles densely spaced,
1-3 mm long, intergrading with the phyllaries.
Capitulescence a narrow panicle with stiffly
ascending racemiform branches above the
middle, or sometimes more broadly and diffusely
branched from the lower nodes. Flowering heads
0.8—1.5 cm in diameter when the rays are fully
extended, typically on slender, puberulent or
glabrous peduncles 1—5 cm (or more) in length
(rarely subsessile). Involucre 3—5 mm high,
narrowly campanulate (or turbinate when pressed
and dried), the phyllaries imbricated in 4 or 5
series. Phyllaries strongly graduated, appressed
or slightly spreading, glabrous on the abaxial
surface but often with a few trichomes on the
adaxial surface, the green areoles well delimited,
thombic-obovate to broadly oblanceolate, the
apex obtuse or acute, the margins irregularly
ciliolate, hyaline, and also with a scarious rim,
the basal */: portion scarious except for the often
slightly keeled green or brown midrib; outer
phyllaries 1—1.5 mm long, oblong-lanceolate,
those largest in surface area (3rd or 4th series
inward) more than twice as long and ca twice as
wide as the outer ones, linear or somewhat
expanded toward the apex; innermost phyllaries
very slender. Receptacle shallowly alveolate
with sharp teeth. Ray florets 15—25 (or more),
the corollas 4—6(8) mm long, typically pink or
lavender (sometimes white), glabrous. Disk
florets 15—20 (or more), the corollas narrowly
funnelform, 3.54.5 mm long, glabrous, the limb
cream-colored or light yellow turning pink after
anthesis, the lobe/limb fraction 0.25—0.35, the
tube shorter than the limb. Pappus simple, the
bristles approximately as long as the disk corolla,
whitish, soft, slender, and attenuate. Achenes
oblong-obovoid, plump or slightly compressed,
1.5—2.5 mm long and 0.5-0.7 mm across, pink, or

May 1989

straw-colored with pink streaks, strigillose and
sometimes minutely puncticulate, with 3—5
prominent ribs, the sculpturing conspicuous even
in immature ovaries. 2” = 16, 32, sometimes with
additional B-chromosomes.

Late August—October. Moist or wet ground:
bogs and sandy or calcareous flats. Not common
in Illinois; apparently concentrated in the
northeastern part of the state, south to Champaign
County and west to Lee County. Collections
recorded from Hancock and Jo Daviess counties
may belong in or near Aster fragilis rather than A.
dumosus. Our records mark the northwestern
limit for this extremely variable species, the
range of which extends from ME to s. ON, MI,
and s. WI, south to s. FL and LA, and southwest
to MS, AR, the s.e. corner of OK, and e. TX.

Good evidence suggests intergradation
between this taxon and the often sympatric
populations of Aster fragilis [e.g., AG. Jones
3303 (ILL) from Iroquois Co.] and A. lanceolatus
var. interior (Wieg.) Semple & Chmielewski
[e.g., A.G. Jones 3304 (ILL) from Iroquois Co.
and E.J. Hill 100, 1884 (ILL) from Cook Co.]. In
fact, all characteristics that, according to Torrey
and Gray (1841) and Wiegand (1928), distinguish
A. dumosus var. strictior from typical var.
dumosus may be traceable to similar characteris-
tics in one or both of the above-mentioned taxa,
e.g., the “strict” branching habit, the often
racemiform and relatively short-peduncled
exposition of heads, and the comparative paucity
of indument. These species are closely related
and share a basic chromosome number of x = 8.

6. Aster ericoides L.
White Prairie Aster, Heath
Aster, Wreath Aster

aS

Herbaceous, with a
strongly creeping
(Illinois plants) or short’
caudiciform rhizome
system, depending on the
variety. New shoots initiated
at the base of old stems or from \
stoloniform rhizome strands, the
plants forming colonies or scattered
individual clumps. Stems |—few at
any point of emergence, erect, the branches

Bigna
Boaiperoin
smeccoccueliry

ASTER AND BRACHYACTIS IN ILLINOIS 155

ascending or often somewhat fastigiate and
arching, 30—80(100) cm tall, uniformly and often
harshly pubescent, the trichomes appressed-
strigose, ascending, or spreading to reflexed
hirsute (depending on the variety), the lower
stem portion sometimes glabrescent. Leaves
numerous, firm, sessile, entire, variously
pubescent on both surfaces, often with clusters of
smaller leaves at the nodes and in the axils of the
larger ones, commonly all except the rameal
leaves withered and deciduous at flowering time;
basal rosette leaves spatulate or oblanceolate,
2-6 cm long and | cm or less wide, triple-
nerved, i.e., with 1(sometimes 2) pair(s) of
secondary nerves running from the base more or
less parallel to the midrib, the surfaces sparsely
pubescent or often glabrous, the apex rounded or
obtuse and with a clear spinule, the base
sheathing; principal cauline leaves linear to
linear-lanceolate, (1)3—6 cm long and 2-7 mm
wide, with only the midrib prominent or triple-
nerved as in the basal leaves, grayish green, the
indument strigose or hirsute (depending on the
variety), the apex acute or obtuse and spinulose,
the base rounded or slightly clasping; rameal
leaves similar in contour, color, and vestiture,
abruptly reduced in size, those of the peduncles
bracteiform, oblong-ovate, 1.5—5(6) mm long,
usually squarrose or sometimes appressed to
ascending, intergrading with the phyllaries.
Capitulescence a somewhat fastigiately
branched panicle or often pyramidal at the top,
the head-bearing branchlets arching, racemiform.
Flowering heads numerous, small, mostly less
than | cm in diameter when the rays are fully
extended, crowded and secund, subsessile arising
from the leaf axils, or on densely bracteate
peduncles 0.5—1(2) cm long. Involucre cylindric
or narrowly campanulate, ca 34 mm high, the
phyllaries imbricated in 3 or 4 series. Phyllaries
strongly graduated, firm, squarrose, with a well-
delimited oblanceolate or spatulate green areole
in the apical portion, the basal '/:—'/> portion
scarious and somewhat indurate; outer phyllaries
oblanceolate to spatulate, 1.5—2 mm long, at least
somewhat pubescent on both surfaces, often
conspicuously scabrous-hirsute abaxially, the
apex acute to obtuse and with a prominent
spinule, the margins scabrous; inner phyllaries
linear-lanceolate, more slender than the outer
and median, 3-4 mm long, usually glabrous,
acuminate or attenuate. Receptacle alveolate
with sharp teeth. Ray florets 8—18(20), the
corollas 4—6(7) mm long, white (rarely pink or

156 ILtinois NatuRAL History SURVEY BULLETIN

bluish), glabrous or with a few trichomes on the
tube. Disk florets 8—12(15), the corollas
narrowly funnelform, 2.5—4 mm long, glabrous or
with a few trichomes at the throat, the limb
gradually expanded, light yellow turning purple
after anthesis, the lobe/limb fraction 0.3—0.35, the
tube ca half as long as the limb. Pappus simple,
the bristles about as long as the disk corolla,
whitish, soft, slender, and attenuate. Achenes
plump, oblong-obovoid, often somewhat falcate,
1.2-2 mm long and 0.40.6 mm across, deep
purple turning brown when weathered, sericeous
or densely strigillose, and with 7-9 ribs. 2n = 10
(mostly), 20. Including A. exiguus Rydb. [A.
multiflorus Aiton—Mead 1846; Brendel 1887;
Higley and Raddin 1891; Pepoon 1927; Benke
1928.]

September—October. Unshaded habitats:
prairie remnants, dunes, gravelly exposed slopes,
and open woods; also in somewhat disturbed
ground that still supports native prairie vegeta-
tion, e.g., roadsides and railroad rights-of-way,
but usually not in severely weedy situations.
Widely distributed throughout the state. The
species range extends over much of the North
American continent, in the East from s. ME to s.
VA, in the Midwest from s. ON and s.e. MB to
AR, and in the West from SK, AB, and e. BC
through e. WA, n.e. OR, ID, and the Great Plains
to NM, TX, e. AZ, and n. Mexico.

Plants of Illinois belong in subsp. ericoides
(A.G. Jones 1978b, c), comprising var. ericoides
and var. prostratus (Kuntze) S.F. Blake [Aster
exiguus—G.N. Jones 1945, 1950, 1963; Fell
1955; Jones and Fuller 1955; Mohlenbrock and
Voigt 1959]. Although these two varieties are
widely sympatric and plants with intermediate
characteristics have been recorded, the identity
and integrity of the taxa seem to be largely
maintained as follows:

1. Pubescence of stems, leaves, and phyllaries
often relatively sparse and soft, appressed-
strigose, or trichomes on the stems some-
timeS aSCENdINg...............000.2---- var. ericoides

1. Pubescence of stems, leaves, and phyllaries
copiously and harshly hirsute, the trichomes
on the stems spreading or somewhat reflexed

aaehee lech aecheseveeceeatuctocerenerenectear var. prostratus

Vol. 34 Art. 2

In addition, two color forms originally
described from Illinois by Benke (1928, 1932b)
are occasionally observed: Aster ericoides f.
caeruleus (Benke) S.F. Blake (plants with blue to
purplish rays), and A. ericoides f. gramsii Benke
(plants with “rose-red” rays). Also included here
are Illinois collections cited as belonging in the
hybrid A. ericoides var. prostratus x A. pilosus
(Mohlenbrock 1986). The hybrid formula was
originally published as A. exiguus x pilosus and
attributed to H.E. Ahles (Jones and Fuller 1955:
468). These plants are not hybrids but are
somewhat atypical individuals of var. prostratus,
probably the result of injury, e.g., from mowing.
Hybridization is well documented and rather
common between A. ericoides and A. novae-
angliae. The plants are known under the binomial
A. x amethystinus Nutt., pro sp. (see discussion
under 14a).

6a. Aster falcatus Lindley in Hooker var.
commutatus (Torrey & Gray) A.G. Jones
Western Heath Aster [A. commutatus (Torrey
& Gray) A. Gray—Fernald 1950]

A few specimens collected along a railroad track
in Kane County [Sherff 1798 (F, ILL)] seem to
approach the characteristics of and perhaps
belong in this taxon, which is of common
occurrence from the Dakotas and Nebraska
westward, especially in the Great Plains. The
plants exhibit a somewhat decumbent sprawling
habit, heads that are not secund and are some-
what larger than is typical for Aster ericoides,
and an abundance of larger cauline leaves not
usually present at flowering time in plants of

A. ericoides. | have included this western taxon in
the key, but if the Illinois plants really belong
here, rather than representing aberrant forms of
A. ericoides that resulted from some kind of
injury or other environmental influence, they are
almost certainly waifs. I do not share the view of
Jones and Fuller (1955: 468), who suggested that
collections of “Aster commutatus sensu Fernald
(1950) 1433, quoad pl. IIl—Non (Torr. & Gray)
A. Gray (1884)” are hybrids between A. ericoides
(as A. exiguus) and A, pilosus.

May 1989

7. Aster fragilis Willd.
Brittle Aster (my suggested
vernacular name), Small
White Aster

Herbaceous, with
creeping (sometimes
short) horizontal
rhizomes. New shoots
arising mostly from slender
stoloniform rhizomes, rarely
at the base of old stems, the
plants forming colonial patches
or scattered individual clumps.
Stems slender, erect, |—-few at any
point of emergence, 30—80(100) cm tall,
commonly branched from below the middle, the
branches divaricate or ascending, brittle, often
slightly ridged from decurrent leaf bases,
minutely puberulent in thin decurrent lines, the
main stem usually glabrescent to glabrous.
Leaves somewhat polymorphic, with clusters of
smaller leaves in axils of the larger ones at most
nodes, the principal leaves commonly withered
and deciduous at flowering time; basal rosette
leaves subpetiolate, spatulate or oblanceolate,

1-4 cm long and less than 1 cm wide, usually
purple-tinged below, with several weakly
expressed and anastomosing secondary nerves,
glabrous, the apex acute or obtuse, the margins
crenate, the subpetiolar portion sparsely ciliate,
winged, and somewhat dilated toward the
sheathing base; principal cauline leaves linear to
linear-oblanceolate, 2—6 cm (or more) long and
(1)2—3 mm wide, usually with only the midrib
strongly expressed, both surfaces glabrous or the
upper minutely puberulent, the apex acute to
attenuate with a sharp callus point, the margins
often inrolled, scabrous, entire or shallowly
serrulate, the base sessile, usually slightly
sheathing and decurrent; rameal leaves numerous,
linear or oblong, ascending or spreading, notably
unequal in size, the smaller bracteiform, those of
the peduncles often aciculiform, 1—2 mm long,
glabrous, intergrading with the phyllaries.
Capitulescence a diffuse panicle, the head-
bearing branchlets typically racemiform and
often arching. Flowering heads 0.6—1 cm in
diameter when the rays are fully extended, often

ASTER AND BRACHYACTIS IN ILLINOIS 157

crowded but commonly not secund, subsessile,
or on slender peduncles 0.2—3 cm (or more) in
length. Involucre 2.54 mm high, narrowly
campanulate or turbinate, the phyllaries imbri-
cated in 4 or 5 series. Phyllaries strongly
graduated, appressed or the outer ones spreading,
glabrous, the green areoles linear to linear-
oblanceolate, usually extending over the entire
length of the midrib, the apex acute, the margins
sparsely ciliolate and with a scarious rim
extending to near the apex; outer phyllaries
slender, linear, 0.8—1 mm long; median and inner
phyllaries linear-oblanceolate, the innermost
2.5—3.5 mm long, those largest in surface area
(3rd or 4th series inward) typically less than
twice as wide as the outer ones. Receptacle
alveolate with sharp teeth. Ray florets
(12)16—20, the corollas 3-4(6) mm long, white
(rarely pink), glabrous. Disk florets 16—20 (or
more), the corollas narrowly funnelform, 2.5—3.5
mm long, glabrous, the limb abruptly dilated at
the throat, cream-colored or light yellow turning
pink after anthesis, the lobe/limb fraction
0.35-0.45, the lobes notably recurved, the tube
slightly shorter than the limb. Pappus simple,
the bristles approximately as long as the disk
corolla, whitish, soft, slender, sometimes slightly
flattened at the acute apex. Achenes oblong-
obovoid, plump or slightly compressed, 1.0—1.8
mm long and ca 0.5 mm across, gray, thinly
strigillose, with 4 or 5 often obscure ribs. 21 =
16, 32. [A. vimineus misapplied, not of Lam. (cf.
Jones and Hiepko 1981; Jones and Lowry 1986),
including A. vimineus var. subdumosus Wieg.—
Deam 1940; G.N. Jones 1950, 1963; Jones and
Fuller 1955; Mohlenbrock and Voigt 1959;
Gleason and Cronquist 1963; Steyermark 1963;
Mohlenbrock 1975, 1986; Swink and Wilhelm
1979; and others. |

Late August—October. Moist or wet ground:
bogs, meadows, lakeshores, streamsides, and
open bottomlands. Occasional throughout much
of the state. The range of this variable species
extends from ME south to FL, and at its western
limit from s. WI southward through MO and AR
to LA and e. TX.

Most if not all Illinois populations belong in
Aster fragilis var. subdumosus (Wieg.) A.G.
Jones, a taxon typified by a collection [Ridgway

158 ILLinois NaTuRAL History Survey BULLETIN

68 (GH)] from Olney, Richland County
(Wiegand 1928: 171; A.G. Jones 1984). These
plants differ from typical var. fragilis mainly by
having relatively long peduncles, 1.e., by
somewhat resembling plants of A. dumosus.
Good evidence suggests occasional hybridization
and gene flow between A. fragilis and several
other taxa that share the basic chromosome
number of x = 8, such as A. dumosus, A.
lateriflorus [e.g., A.G. Jones 3108 (ILL) from
Clark Co. and Jelinek 22/ (ILL) from Pope Co.],
A. lanceolatus var. interior [e.g., Mead s.n. (ILL)
from Hancock Co.], and A. ontarionis [e.g.,
Shildneck 15860 (ILL) from Cass Co. and Neill
16357 (ISM) from St. Clair Co.].

8. Aster furcatus Burgess in Britton & Brown
Forked Aster

Herbaceous, with a
creeping, horizontal
rhizome system
producing fleshy,
stoloniform rhizome
strands. New shoots arising at
the base of old stems or from
the rhizomes, the plants forming
colonial stands or scattered
individual clumps, the few small
rosettes produced later in the season typically
vanishing shortly after the first killing frost.
Stems |-several, erect, often somewhat zigzag in
the upper portion, (30)50—120 cm tall, the
branches typically confined to the capitulescence,
ascending, more or less dichotomously forked,
somewhat ridged from decurrent leaf bases, the
indument uniformly distributed around the stem,
hirtellous or puberulent on the upper stem and in
the branches, the lower portion of the stem
glabrescent. Leaves polymorphic, those of the
basal rosettes and the main stem petiolate; leaves
of new shoots and of the autumnal rosettes with
oblong to ovate-lanceolate blades 4-10 cm long
and 2—5 cm wide, rugose, with an often reddish
midrib and several pairs of pinnate secondary
nerves, minutely scabrous above and hirsute
below, the apex acute or obtuse, the margins
serrate, the base rounded or obliquely subcordate,
the petioles short, winged, sheathing at the base:

Vol. 34 An. 2

cauline leaves gradually reduced in size upward
on the stem, those of the midstem with ovate to
lance-ovate blades 10—12(15) cm long and (3)6-8
cm wide, rugose, with (6)8—12 pairs of pinnate
secondary nerves, and with a weakly expressed
reticulum of tertiary veins with more or less
isodiametric areolae, harshly scabrous on the
upper surface, densely hirsute over the entire
lower surface resulting in a grayish green color,
the apex acuminate with a long mucro, the
margins sharply serrate with conspicuously
mucronate teeth, the base shallowly cordate or
truncate to rounded; petioles slender, mostly
shorter than the blades, glabrous or irregularly
ciliate, the base slightly dilated, sheathing, and
somewhat decurrent; rameal leaves relatively
few, similar in contour but abruptly reduced in
size, subpetiolate to sessile, the base slightly
clasping or sheathing, the midrib strongly keeled
and decurrent; leaves of the peduncles 0—3, often
bracteiform and phyllarylike, depending on the
position, sometimes closely subtending the heads.
Capitulescence variable, broad-topped, compara-
tively few-headed and cymiform in the natural
habitat but often ample, much-branched, and
compound corymbiform in disturbed situations.
Flowering heads 2.5—3.5 cm in diameter when
the rays are fully extended, not crowded and not
secund, the peduncles 0.5—3(5) cm long, often
leafless or nearly so, densely hirtellous or villous.
Involucre campanulate (rarely hemispherical
when pressed and dried), 6-8(10) mm high, the
phyllaries imbricated in 5—7 series. Phyllaries
appressed when fresh, strongly graduated, the
outer ones ovate, 1.5—3 mm long, the innermost
linear-lanceolate, 5.5—7.5 mm long, those of the
outer 2 or 3 series twice as wide as the inner,
puberulent on both surfaces, the apex obtuse or
rounded, the margins lanate, erose-hyaline, and
also often (median and inner phyllaries) with a
red rim, the chlorophyllous areoles poorly
delimited, with fuzzy edges, broadly obovate to
rhombic or oblong, in the apical /:—/s portion or
rarely extending to the base in the form of bands,
the basal '/:—/; portion scarious, indurate, and
keeled or rounded on the back. Receptacle
shallowly alveolate with blunt teeth. Ray florets
(12)15—20, the corollas 12—18 mm long, white
(sometimes aging to pink), glabrous. Disk florets
25-35 (or more), the corollas funnelform, 6—-7(8)
mm long, glabrous, the limb partway slenderly
tubular, abruptly dilated 1—1.5 mm above the
point of insertion of the filaments, cream-colored

May 1989

or light yellow turning purple after anthesis, the
fraction of lobe/expanded limb portion 0.4—0.5,
the apparent tube (i.e., including the tubular limb
portion) distinctly longer than the expanded limb
portion. Pappus slightly shorter than the disk
corolla, whitish or somewhat tawny, the bristles
in 2 series, those of the inner series firm, with a
clavellately expanded apex, those of the outer
somewhat shorter, more slender, and attenuate.
Achenes fusiform, plump, (2.5)3—3.5(4) mm long
and 1—1.2 mm across, dull purple or brown,
thinly puberulent in lines or glabrescent, with
8-10(12) densely spaced, often straw-colored
ribs. 2n = 18, with 0O-several B-chromosomes.
[A. divaricatus misapplied, not of L—Mead
1846; Kibbe 1952.]

(Late July) August—October. Low springy or
marshy ground in woody areas, usually underlain
by sandstone or limestone: lower (mostly north-
facing) slopes and edges on the cutting (eroding)
sides of streams and rivers. Rare and possibly
threatened, apparently less common now than is
indicated by the collection records, although
occasional stands are extensive. The populations
are restricted to the northern one-third of the
state, extending south along the major rivers to
Tazewell, Fulton, and Hancock counties. The
species range includes scattered stations in s. MI,
w. IN, s.e. WI, e. IA, and s.e. MO.

9. Aster laevis L.
Smooth Aster, Smooth
Blue Aster

Herbaceous, with
branched caudiciform or
tangled rhizomes that
turn woody with age but
also producing stoloniform
rhizome strands. New shoots
arising at the base of old stems
or from rhizome branches, the
plants forming scattered individual
clumps or small patches. Stems
1-several, erect, (30)50—120 cm tall, with stiff,
typically ascending, and commonly somewhat
ridged branches above the middle, often reddish
in the lower portion, glaucous and essentially
glabrous except for sparse hirsute indument on

ASTER AND BRACHYACTIS IN ILLINOIS 159

the peduncles and in the leaf axils. Leaves
polymorphic, somewhat fleshy, glabrous and
glaucous, the basal and larger cauline ones
sometimes withered at flowering time; basal
rosette leaves commonly with long, winged, and
basally dilated petioles, the blades oblong or
ovate to lanceolate, 3-10 cm (or more) long and
ca 1-3 cm wide, the apex acute or obtuse to
rounded, the margins subentire or crenate-
serrate, the base rounded or truncate; principal
cauline leaves subsessile or mostly sessile,
extremely variable in size and shape, oblong-
ovate or lanceolate to linear, sometimes abruptly
constricted above the base, (4)8—15 cm long and
(1)2—4.5 cm wide, with 1—several pairs of
ascending secondary nerves, and a weakly
expressed reticulum of tertiary veins with more
or less isodiametric areolae, the apex acute or
obtuse, usually callus-pointed, the margins entire
or shallowly crenate-serrate, and minutely
scabrous, the base often strongly auriculate-
clasping; rameal leaves similar in contour but
abruptly reduced in size, with a clasping or
sheathing and somewhat decurrent base; leaves
of the peduncles usually densely spaced,
bracteiform, lance-subulate, 3-6 mm long, with
an often purple callus point, intergrading with
the phyllaries. Capitulescence a narrow or often
somewhat flat-topped panicle in the upper '/: —'/»
of the stem, the branches commonly stiffly
ascending (rarely arching). Flowering heads
1.5—2.5(3) cm in diameter when the rays are
fully extended, terminal on somewhat ridged and
often sparsely puberulent peduncles varying
from 0.2 to 6 cm (or more) in length. Involucre
campanulate, 4.5—7(8) mm high, the phyllaries
imbricated in 4-6 series. Phyllaries strongly
graduated, firm, appressed when fresh, the outer
ones subulate or lanceolate, 1.5—2 mm long, the
median and inner ones three times as long, linear
or often slightly expanded toward the apex, the
surfaces glabrous, the apical green areoles well
defined, rhombic to lance-rhombic, glaucous,
commonly smaller in area than the scarious basal
portion, the apex obtuse, acute, or acuminate,
with a sharp and often red or purple callus point,
the basal portion of outer and median phyllaries
usually keeled and indurate. Receptacle
alveolate with sharp and typically bristle-tipped
teeth. Ray florets 15—25(30), the corollas
(10)12—15 mm long, blue or purple (rarely
white), glabrous. Disk florets 15—25 (or more),
the corollas narrowly funnelform, (4.5)5-6 mm

160 ILLinois NATURAL History SURVEY BULLETIN

long, glabrous, the limb slightly dilated at the
throat, yellow turning purple after anthesis, the
lobe/limb fraction ca 0.2, the tube slightly shorter
than the limb. Pappus simple, the bristles usually
somewhat shorter than the disk corolla, tawny or
rose-tinged, soft, slender, and attenuate. Achenes
oblong-obovoid, plump or somewhat com-
pressed, 2-3 mm long and 0.8—1.3 mm across,
deep purple at maturity, or brown when weath-
ered, glabrous, with 4 or 5 often irregularly
spaced ribs. 2 = 48.

August—October. Mesic, mostly open
habitats: prairie remnants, meadows, lakeshores,
stream banks, open woods, and exposed slopes;
also in somewhat disturbed ground along roads
and railroad rights-of-way. Fairly common
throughout most of the state. The range of the
species extends from PQ south to GA and AL,
northwest to AB, BC, and s. YT, and in the
Rocky Mts. and Great Plains southward through
c. CO to NM and the Guadalupe Mts. in w. TX.

Although somewhat variable, Illinois plants
are assigned to typical var. /aevis. There is
evidence of occasional hybridization and
intergradation with other species that share the
basic chromosome number of x = 8, such as Aster
lanceolatus [e.g., Benke 5280 (F)—cf. Benke
1932b], A. puniceus [e.g., Fuller 13293a (F) from
McHenry Co.], A. oolentangiensis [e.g.,
Heitmann & Heitmann 1027 (F) from Fulton
Co.], and A. Jateriflorus |e.g., Benke 2927 (F)
from Kane Co.}.

10. Aster lanceolatus Willd.
Panicled Aster

Herbaceous, with a
strongly creeping
horizontal rhizome
system. New shoots
initiated mostly at the tips of
stoloniform rhizomes, the plants
extensively clonal but the
connecting strands short-lived
and often disintegrating after one or
two seasons. Stems mostly single at
any point of emergence, often stout, erect,

Vol. 34 An. 2

40—120(150) cm tall, pubescent in thin decurrent
lines from the upper nodes, usually glabrescent or
glabrous below the middle, often branching from
near the base or producing in midsummer
secondary shoots composed of clusters of smaller
leaves plus a few heads at most of the nodes and
in the leaf axils. Leaves polymorphic, the basal
and larger cauline ones often withered or
deciduous at flowering time; basal rosette

leaves petiolate or subpetiolate, the blades
elliptic-oblanceolate or obovate to suborbiculate,
1-8 cm long and 0.5—2 cm wide, often purplish
below, with several pairs of anastomosing
secondary nerves, glabrous, the apex acute,
obtuse, or rounded, the margins coarsely crenate,
the petioles winged and with a few long marginal
cilia, the base dilated and sheathing; principal
cauline leaves sessile or subsessile, elliptic or
oblanceolate to linear-lanceolate, (4)6—-15 cm
long and (0.5)1—2(3.5) cm wide, gradually
reduced in size up the stem, with relatively
weakly expressed secondary nerves and a
reticulum of tertiary veins with oblong areolae,
glabrous (rarely scabrellous above), the apex
acute to attenuate and callus-pointed, the margins
sharply serrate, the base cuneate and somewhat
decurrent; rameal leaves similar in contour and
texture but mostly entire, those of the peduncles
relatively few, flexible, 2-6(10) mm long, rarely
somewhat intergrading with the phyllaries.
Capitulescence an ample, leafy, diffuse or often
narrowly elongate panicle. Flowering heads
1—2(2.5) cm in diameter when the rays are fully
extended, rarely secund, the peduncles 0.5—5 cm
in length. Involucre campanulate, 3.5—6(7) mm
high, the phyllaries imbricated in 3—5(6) series.
Phyllaries somewhat or often strongly graduated,
comparatively slender, appressed or slightly
spreading, the outer ones |.5—3 mm long, the
inner 3—6 mm; outer and median phyllaries
linear-oblanceolate, glabrous on the abaxial
surface, sparsely puberulent on the adaxial
surface, the green areoles linear-oblanceolate, the
apex acute to acuminate and callus-pointed, the
margins somewhat ciliolate, erose-hyaline, and
with a narrow scarious rim extending to the tip,
the basal '/:—'/> portion scarious and slightly
dilated; innermost phyllaries linear, very slender,
acuminate to attenuate. Receptacle alveolate with
sharp teeth. Ray florets 20—35 (or more), the
corollas 7—10(12) mm long, white (rarely pink),

May 1989

glabrous. Disk florets (15)20—30 (or more), the
corollas funnelform, 3—5(6) mm long, glabrous,
the limb cream-colored turning purple after
anthesis, dilated at the throat, with comparatively
long recurved lobes, the lobe/limb fraction
0.35—0.45(0.5), the slender tube somewhat
shorter than the limb. Pappus simple, the bristles
as long as the disk corolla or often slightly
longer, whitish, soft, slender, and attenuate.
Achenes oblong-obovoid, somewhat compressed,
1.5—2 mm long and 0.5—0.8 mm across, gray,
thinly strigillose, 4- or S-ribbed. 21 = 32, 64
(mostly), rarely 40 or 48. Including A. simplex
Willd. and A. interior Wieg. (Semple 1979;
Semple and Chmielewski 1987). [A. paniculatus
Lam. non Miller; A. tradescantii misapplied,

not of L.]

August—October. Low moist or swampy
ground: woods, pastures, bottomlands of rivers
and streams, ditch margins, and lakeshores. Very
common. Under the broad taxonomic concept
adopted by Semple and Chmielewski (1987), the
species has a transcontinental range that includes
localities in nearly every state and province of
North America.

Three varieties can be distinguished in
Illinois: 1) var. lanceolatus [Aster paniculatus—
Brendel 1887; Higley and Raddin 1891; Pepoon
1927; Deam 1940; G.N. Jones 1945, 1950; Fell
1955. A. simplex var. ramosissimus (Torrey &
Gray) Cronq.—Fernald 1950; Gleason 1952;
Dobbs 1963; Gleason and Cronquist 1963;
Steyermark 1963], 2) var. simplex (Willd.) A.G.
Jones [A. simplex (sensu stricto)—Mead 1846;
Fernald 1950; Gleason 1952; Kibbe 1952; Jones
and Fuller 1955; Mohlenbrock and Voigt 1959;
G.N. Jones 1963; Gleason and Cronquist 1963;
Swink 1974; Mohlenbrock 1975, 1986; Swink
and Wilhelm 1979. A. paniculatus var. simplex
(Willd.) Burgess in Britton & Brown—Wiegand
1933; Deam 1940_A. lanceolatus subsp. simplex
(Willd.) A.G. Jones—1984], and 3) var. interior
(Wieg.) Semple & Chmielewski [A. tradescantii
sensu auct. non L.—Mead 1846; Brendel 1887;
Higley and Raddin 1891; Pepoon 1927; G.N.
Jones 1950; Fell 1955. A. vimineus sensu auct.,
pro parte, non Lam.—Higley and Raddin 1891;
Kibbe 1952. A. interior—Deam 1940; G.N. Jones
1945; Jones and Fuller 1955; Mohlenbrock and

ASTER AND BRACHYACTIS IN ILLINOIS 161

Voigt 1959. A. simplex var. interior (Wieg.)
Crong.—Fernald 1950; Gleason 1952; Gleason
and Cronquist 1963; Swink 1974; Swink and
Wilhelm 1979. A. x interior Wieg., pro sp.—
A.G. Jones 1980a, b. A. lanceolatus subsp.
interior (Wieg.) A.G. Jones—1984]. Plants of
these three varieties show a great deal of
intergradation. For the most part, however, they
may be distinguished as follows:

1. Flowering heads 1—1.5 cm in diameter when
the rays are fully extended; involucres 34
mm high; disk corollas 3—3.5 mm long;
branchlets often racemiform ..... var. interior

1. Flowering heads 1.8—2.5 cm in diameter
when the rays are fully extended;
involucres (4)4.5—7 mm high; disk
corollas typically (4)4.5—5.5 mm long:
branchlets usually not racemiform.

2. Leaves of the midstem linear-lanceolate,

12 times (or more) longer than wide ......

spesutenctcvstecersnespbiseteesteeed var. lanceolatus

Leaves of the midstem elliptic-

lanceolate to oblanceolate, mostly less

than 11 times longer than wide...............

Bewasduaihe totes tveg suest ieee venidseews var. simplex

tN

For a different interpretation see Semple and
Chmielewski (1987). These authors do not
regard Aster simplex as sufficiently distinct from
typical A. lanceolatus to merit taxonomic
recognition. Collections of var. simplex from the
central and southern parts of the Midwest,
however, consistently seem to differ in several
ways from the more northern populations of var.
lanceolatus. Aside from the key characters
mentioned and the somewhat differential
geographic distribution, plants of var. simplex
occur more frequently in shady bottomland
woods, whereas plants of both var. lanceolatus
and var. inferior are more common in the
unshaded habitats of bogs, meadows, lakeshores,
and ditch margins.

Good evidence suggests that this extremely
variable species occasionally hybridizes and
intergrades with other species of the x = 8
chromosome number assemblage, e.g., Aster
borealis, A. fragilis, A. laevis, A. lateriflorus, A.
puniceus, and others that are not elements of the
flora of Illinois.

162 ILLiInois NATURAL History SURVEY BULLETIN

11. Aster lateriflorus (L.) Britton
Starved Aster, Calico
Aster, Side-flowered
Aster, White Woodland
Aster

Herbaceous, with a short,
somewhat tangled
rhizome system. New
shoots initiated at the base
of old stems or from short
ascending rhizome branches,
the plants forming scattered
individual clumps. Stems
l—several, 30—80(120) cm tall,
comparatively slender, brittle, with
ascending or often divaricate branches from the
lower nodes resulting in a sprawling or bushy
habit, the indument villous or hirtellous, usually
in decurrent lines from the leaf bases, sometimes
uniformly distributed in the upper branches, the
main stem often glabrescent or glabrous. Leaves
polymorphic, the larger cauline ones at least in
part persistent throughout the flowering period;
basal rosette leaves subpetiolate, the blades
spatulate or obovate to suborbiculate, the upper
surface minutely puberulent or glabrous, the
lower surface villous along the midrib (rarely
glabrous) and sometimes purplish, the margins
crenate-serrate, abruptly narrowed to the winged,
ciliate, somewhat sheathing subpetiolar base;
principal cauline leaves sessile or subsessile,
mostly elliptic-oblanceolate, rarely linear,
(3)5—10(15) cm long and (0.2)1—2(3.5) cm wide,
gradually reduced in size up the stem, usually
with several pairs of anastomosing secondary
nerves, and a weakly expressed reticulum of
tertiary veins with oblong areolae, the upper
surface typically scabrellous, the lower surface
villous or hirtellous along the midrib but
otherwise usually glabrous, the apex acuminate to
attenuate with a sharp callus point, the margins
serrate and minutely scabrous, the base cuneate
and somewhat decurrent; rameal leaves similar in
contour, more or less glabrous, entire, those
subtending the head 1—-few, oblong-lanceolate,
1.5—3 mm long, flexible, not phyllarylike.
Capitulescence an often ample diffuse panicle,
commonly branched from below the middle, the
head-bearing branchlets slender and wiry,
arching, racemiform. Flowering heads small,
0.8—1.3 cm in diameter when the rays are fully
extended, often crowded, secund, sessile or
subsessile, the peduncles rarely more than | cm

Vol. 34 Art. 2

in length. Involucre campanulate (turbinate when
pressed and dried), 3.5-4.5(5) mm high, the
phyllaries imbricated in (3)4 or 5 series.
Phyllaries strongly graduated, appressed or
slightly spreading, glabrous on the abaxial
surface, the apical green areoles obovate-
oblanceolate, the basal '/;—/: portion scarious
except for the green midrib; outer phyllaries
linear-lanceolate, 1-2 mm long, less than half as
long as the inner, acute and callus-pointed:
median phyllaries linear or often slightly
expanded toward the acute to acuminate apex,
sparsely puberulent on the adaxial surface, the
margins irregularly ciliolate, erose-hyaline, and
with a narrow scarious (sometimes reddish) rim
extending to the tip; innermost phyllaries very
slender, linear, 3.54.5 mm long. Receptacle
alveolate with sharp or bluntish teeth. Ray florets
10—15(20), the corollas (4)5—8 mm long, white,
glabrous. Disk florets 10—15(20), the corollas
funnelform, (2.5)3.5-4.5 mm long, glabrous or
with a few trichomes, the limb abruptly dilated

at the throat, cream-colored or light yellow
before anthesis, soon turning deep magenta, the
lobe/limb fraction 0.5—0.75, the lobes strongly
reflexed, the slender tube slightly shorter than the
limb. Pappus simple, the bristles about as long as
the disk corolla, white, soft, slender, and
attenuate. Achenes oblong-obovoid, plump or
slightly compressed, (1.3)1.8—2.2 mm long and
0.5—0.7 mm across, gray, sparsely strigillose,
obscurely 3—5 ribbed. 27 = 16, 32 (mostly), 48.
Including A. pendulus Aiton, A. horizontalis
Desf., and A. hirsuticaulis Lindley in DC.

[A. vimineus Lam. (see A.G. Jones 1984: 379;
Jones and Lowry 1986). A. diffusus Aiton—
Brendel 1887; Higley and Raddin 1891. A. miser
sensu Aiton non L.—Mead 1846.]

Late August—October. Wooded slopes, high
banks of streams and rivers, and edges of bogs:
Illinois plants more common in well-drained or
upland situations than in wet ground habitats.
Common throughout the state. The range of the
species extends in the East from NB, ME, PQ,
and s. ON south to FL, and to the west from MN
and e. IA through e. KS to LA and s.e. TX.

Varietal names can be assigned to collections
from Illinois only with the greatest difficulty
because of a great deal of intergradation, not only
between the varieties that have been described
(Wiegand 1928) but also involving gene flow
from other species with a basic chromosome
number of x = 8, e.g., Aster dumosus, A. fragilis,

May 1989

A. lanceolatus, and A. ontarionis. The majority of
specimens more or less fit the circumscription
and concept of typical var. lateriflorus. Plants of
var. horizontalis (Desf.) Farw. [A. lateriflorus
var. pendulus sensu Wieg. non (Aiton) Burgess in
Britton & Brown] are not uncommon, especially
in the southern half of the state. The characteris-
tics that weakly distinguish this variety (very
small heads combined with a sprawling habit and
a tendency toward relatively moist habitats) may
have been introduced in part through gene flow
from A. fragilis. Illinois collections labelled

A. lateriflorus var. hirsuticaulis (Lindley) Porter
{sensu Pepoon 1927] are partly referable to A.
ontarionis and partly to typical var. lateriflorus.
The range of var. hirsuticaulis [including var.
tenuipes Wiegand (1928)] probably does not
extend to Illinois; these plants are common in the
northeastern states and in Canada west to s. ON.

12. Aster linariifolius L.
Flax-leaved Aster,
Savory-leaved Aster

Tufted, often suffruti-
cose, with a tangled or
cormoid-caudiciform
rhizome system that turns
woody with age. New
shoots arising at the base of
old stems but also at the end of
ascending, densely bracteate,
rhizome strands; basal rosettes
absent. Stems cespitose, several to
many, slender and wiry, (10)20-40(60) cm tall,
simple, or few-branched only in the capitules-
cence (unless injured through grazing or
mowing), cinereous, puberulent, sometimes
glabrescent toward the base. Leaves numerous,
firm, densely spaced, sessile, ascending or
spreading to squarrose, more or less uniform in
contour, linear or nearly so, entire, those of the
main stem 1.5—2.5(3.5).cm long and 1.5—2.5(4)
mm wide, with a keeled midrib, the secondary
venation not evident, glabrous except for the
scabrous margins and the midrib below, the
epidermal cells of both surfaces strikingly
bulliform (hand lens needed to see this), the apex
acute with a sharp callus point; lowermost
cauline leaves bracteiform; rameal leaves
abruptly reduced in size, those of the peduncles

ASTER AND BRACHYACTIS IN ILLINOIS 163

intergrading with the phyllaries. Capitulescence
racemiform or corymbiform in the upper '/»—'/s
of the stem. Flowering heads relatively few per
stem, commonly fewer than 30, 1.5—3 cm in
diameter when the rays are fully extended, on
leafy peduncles 0.5—S(15) cm in length.
Involucre turbinate or campanulate, (6)8—10(12)
mm high, the phyllaries imbricated in 5-8 series.
Phyllaries strongly graduated, firm, appressed,
often largely scarious on the abaxial surface, the
green areoles not well defined, centered in the
apical '/:—'/> portion or forming two bands along
the whitish midrib, sometimes extending to the
base; phyllaries of the outer 2 or 3 series 2.5-4
mm long, lanceolate, notably keeled, the apex
acute with a red callus point, the margins ciliate
to fimbriate, or lanate toward the apex, those of
the Sth or 6th series inward largest in surface
area, (6)7—8(10) mm long, linear-lanceolate,
keeled or rounded on the back, often with a red
rim, the apex obtuse or rounded, the margins
conspicuously lanate over the entire length.
Receptacle alveolate with sharp teeth. Ray
florets 13-17, the corollas (10)12—15(18) mm
long, violet or reddish purple (rarely white),
glabrous. Disk florets 20—30 (or more), the
corollas narrowly funnelform, 5—6.5(8) mm long,
glabrous, the limb gradually expanded, light
yellow turning reddish after anthesis, the
lobe/limb fraction ca 0.2, the tube slightly
shorter than the limb. Pappus tawny, “double”
(according to literature references) but often
composed of 3 series of bristles, the principal
bristles firm, in 1 or 2 series, somewhat unequal,
4-7 mm long, as long as the disk corolla or
slightly shorter, with at least the innermost
clavellately expanded and flattened toward the
apex; bristles of the outermost whorl distinctly
shorter, 1 mm long or less, and attenuate.
Achenes obconic, plump, 2.5—3(3.5) mm long,
ca 1 mm across, dull purple or brown, 3—5
ribbed, the color and sculpturing of the pericarp
often obscured by densely sericeous indument.
2n = 18. [Diplopappus linariifolius (L.)
Hooker—Mead 1846.]

Late August—October. Dry sandy, loamy, or
rocky soils: exposed slopes and ridges, open
woods, dunes, and barrens. Most prominent in
the northern half of the state but extending south
to St. Clair County. The range of the species
extends from s. PQ and ME south to FL, and to
the west from MN through s. MO, s.e. KS, and e.
OK toe. TX.

164 ILtinois NATURAL History SURVEY BULLETIN

13. Aster macrophyllus L.
Large-leaved Aster,
Big-leaved Aster

Herbaceous, with a
strongly creeping
horizontal rhizome
system. New shoots
arising mostly at the ends of
stoloniform rhizome
branches, the plants typically
forming extensive colonies, the
autumnal basal rosettes numerous,
prominent, often forming a dense
ground cover. Stems 1—few at any point of emer-
gence, erect, sometimes slightly zigzag in the
upper portion, 30—70(90) cm tall, terete or
somewhat angular above from decurrent leaf
bases, often maroon-colored, the branches
confined to the capitulescence, ascending, the
upper stem and branches pubescent with white,
multicellular, antrorsely curved or spreading
trichomes and also invested with sessile or
stipitate glands, the lower stem glabrescent or
glabrous. Leaves polymorphic, those of the basal
rosettes and the lower half of the stem petiolate;
basal rosette leaves largest, the blades broadly
ovate to cordate, (8)15—25 cm long and
(6)8—-12(15) cm wide, with 6-12 pairs of pinnate
secondary nerves, and a reticulum of tertiary
veins with more or less isodiametric areolae, the
upper surface minutely scabrous or glabrous, the
lower thinly puberulent, especially along the
nerves, the apex acuminate with a sharp mucro,
the margins crenate-serrate, each tooth
mucronate, the base deeply cordate and some-
times slightly oblique, the petioles 1—2 times as
long as the blades, with slightly winged and
ciliate margins, dilated and sheathing at the base;
cauline leaves gradually reduced in size up the
stem, blades of the larger ones nearly as wide as
long, similar to the basal leaves in contour and
vestiture except for sparse glandularity along the
midribs below, the petioles shorter and progres-
sively more broadly winged upward on the stem;
upper stem leaves subpetiolate or often sessile
and clasping, ovate or obovate to oblanceolate;
rameal leaves relatively few, abruptly reduced
but variable in size, ovate or oval, the upper
surface scabrous and stipitate-glandular,
especially along the midrib and the margins, the
lower surface more sparsely so, the apex acute or

Vol. 34 An. 2

acuminate, the margins crenate-serrate to entire,
the base rounded or cuneate, the 0—-few leaves of
the peduncles ovate-lanceolate, 2-5 mm long,
thinly to densely glandular, attenuate, not
intergrading with the phyllaries. Capitulescence
a dichotomously branched corymbiform panicle
in the upper '/:—'/; portion of the stem, the
branchlets short and firm. Flowering heads
variable, ca 2—3 cm in diameter when the rays are
fully extended, sometimes crowded in the
individual cymules, the peduncles 0-2.5 cm long,
often densely glandular-hirsute. Involucre
campanulate or hemispherical, (6)8—10 mm high,
the phyllaries imbricated in (4)5—7 series.
Phyllaries appressed or somewhat spreading,
strongly graduated, those of the outer 2(or 3)
series ovate, 2—3 mm long and ca half as wide,
acute to obtuse, the green areoles oblong-
obovate, in the apical '/:—'/2 portion (rarely
extending to the base in the outer phyllaries), the
basal portion mostly scarious, indurate, keeled, or
rounded on the back, glandular-puberulent over
the entire abaxial surface (outermost phyllaries)
or only in the chlorophyllous zone and along the
midrib (median phyllaries), the adaxial surface
more or less glabrous, the margins lanate and
glandular toward the apex, hyaline toward the
base, and with a scarious or often reddish rim:
phyllaries of the innermost series slenderly
linear-lanceolate, (6)7—9 mm long, ca | mm
wide, acute, largely scarious or sometimes red-
rimmed. Receptacle alveolate with sharp teeth.
Ray florets 10—15, the corollas 10-13 mm long,
varying from white to deep lavender, glabrous.
Disk florets 20-30 (or more), the corollas
funnelform, 7-8 mm long, glabrous, the limb
partway slenderly tubular, more or less abruptly
flared ca 1.5—2 mm above the point of insertion
of the filaments, cream-colored or light yellow
aging to purple, the lobes reflexed, the fraction of
lobe/expanded limb portion 0.6—0.7, the apparent
tube (i.e., including the tubular limb portion)
much longer than the expanded limb portion.
Pappus tawny, in 2 series of somewhat unequal
bristles, those of the inner series about as long as
the disk corolla, firm and with a clavellately
expanded apex, those of the outer slightly shorter,
more slender, and attenuate. Achenes slenderly
fusiform or cylindric, 3.5-4(4.5) mm long and
1-1.2 mm across, dull purple or brown, glabrous
or with a few scattered trichomes near the top,
with 8-10 thick, densely spaced (sometimes
double-stranded), glossy, golden-brown ribs.

2n= 12.

May 1989

August—October. Moist or swampy ground
(sometimes dry soil) in wooded areas. Local, rare
in this state, recorded only from Cook and Lake
counties, much more common to the north and
east of Illinois. The range of this variable species
extends in the East from NS, PE, NB, and PQ
south to GA, in the Midwest from ON south to
s.e. IN, and at the western limit from e. MB
through e. MN to n.e. IA.

14. Aster novae-angliae L.
New England Aster

Herbaceous, with a
tangled or sometimes
caudiciform rhizome
system. New shoots
arising at the base of old
stems or from short, fleshy,
horizontal rhizomes, the plants
forming scattered individual
clumps or small patches. Stems
1-several, SO—150 cm tall, stout, with
ascending or divaricate branches from above the
middle, densely hispidulous-hirsute or pilose,
stipitate-glandular in the upper portion, the lower
portion less pubescent and usually brown in
color. Leaves numerous, polymorphic, the basal
and lower cauline ones often withered and
deciduous at flowering time; basal rosette leaves
spatulate, ca 2-6 cm long and 0).5—1.5 cm wide,
with 1—3 pairs of anastomosing secondary nerves
more or less aligned with the midrib or margins,
sparsely hirsute on both surfaces, the margins
ciliolate and occasionally with a few shallow
teeth; principal cauline leaves sessile and entire,
oblong or lanceolate, with several pairs of weakly
expressed secondary nerves, and a reticulum of
tertiary veins with more or less isodiametric
areolae; leaves of the midstem ca 5—10 cm long
and 5—15(20) mm wide, thinly strigose above,
hirsute or hispidulous below, and invested with
both sessile and stipitate glands (more copiously
so in the upper leaves), the apex acute and
mucronulate, the margins pustulate-scabrous, the
base auriculate-clasping; rameal leaves similar in
contour and vestiture, gradually reduced in size,
the 1-4 small leaves of the peduncles not
phyllarylike. Capitulescence a leafy, broad,
round-topped or somewhat corymbiform panicle

ASTER AND BRACHYACTIS IN ILLINOIS 165

in the terminal (‘/.0)'/.—'/s portion of the stem.
Flowering heads 2—4.5(5) cm in diameter when
the rays are fully extended, often crowded but
not secund, the peduncles 0.34 cm long,
glandular and hispidulous, commonly somewhat
dilated just below the head. Involucre broadly
hemispherical, 5-15 mm high, the numerous
phyllaries imbricated in 4-6 series. Phyllaries
slender, linear-lanceolate, long-acuminate or
attenuate, commonly strongly squarrose or
reflexed, mostly subequal in length, (5)6—12(15)
mm long or the outer ones slightly shorter (rarely
longer) than the inner; outer phyllaries largely
herbaceous, dark green and often purple-tinged,
densely invested with both stipitate and sessile
glands; median and inner phyllaries scarious in
the basal '/;—'/> portion. Receptacle shallowly
alveolate or merely pitted. Ray florets numer-
ous, ca 50-100, the corollas 10—20(25) mm long,
rose-colored to deep purple (rarely white), with a
few trichomes near the throat. Disk 5-15 mm in
diameter. Disk florets numerous, 50 or more, the
corollas narrowly funnelform, (4)5—7 mm long,
thinly puberulent in the throat region, the limb
only slightly expanded, light yellow turning
purple after anthesis at least in the lobes, the
lobe/limb fraction 0.2—0.25, the tube ca half as
long as the limb. Pappus simple, the bristles
somewhat unequal and often slightly shorter than
the disk corolla, tawny or rose-tinged, soft,
slender, and attenuate. Achenes oblong to
obconic, plump, 1.8—2.5(3) mm long, ca 0.6—1
mm across, dull purple or brown, obscurely
glandular and densely sericeous, with 7—9 ribs.
2n = 10, 20, occasionally with supernumerary
chromosomes.

August—October. Unshaded mesic situations:
prairie remnants, open woods, streamsides, fens,
also in disturbed soils of pastures, old fields,
roadsides, and railroad rights-of-way. Common
throughout the state; also widely cultivated and
often escaped, a fact that partly accounts for the
various color forms observed. The range extends
from s. PQ, s. ON, and ME south to AL, MS,
and AR, west to s. MB, c. ND, c. NE, and e. KS,
with scattered stations in WY, CO, OK, and NM.

Including f. roseus (Desf.) Britton [var.
roseus (Desf.) DC.—Higley and Raddin 1891;
Pepoon 1927; Benke 1928], plants with rose-
colored rays; and f. geneseensis House (Benke
1932a), a white-rayed form. A short-lived, large-
headed, white-rayed individual found in
Champaign County (A.G. Jones 2604), to which

166 ILLinois NATURAL History SuRveEY BULLETIN

I have applied the latter name, possessed an extra
set (genome) of chromosomes (A.G. Jones
1980b). When cross-pollinated from a typical
individual, a few viable achenes were produced
that grew into typical purple-rayed plants.
Hybrids between plants of Aster novae-angliae
and A. ericoides [A. x amethystinus]| are
occasionally found in Illinois (see 14a).

14a. Aster x amethystinus Nutt., pro sp.
[A. ericoides x A. novae-angliae}

This hybrid is readily identifiable (Benke 1930)
and has also been produced experimentally
(Wetmore and Delisle 1939; A.G. Jones 1978c).
The plants occur with sufficient frequency in
Illinois to warrant inclusion in the key and a
comparative diagnosis: Habit similar to that of
Aster novae-angliae, the plants usually with a
tangled rhizome system and short stoloniform
strands forming small patches. Stems erect, aging
to a light brown color, densely hispidulous-
hirsute but not glandular. Leaves numerous,
sessile, at least some of the larger cauline ones
persistent throughout the flowering period and
with clusters of smaller leaves produced in the
axils; principal cauline leaves linear or oblong to
elliptic-oblanceolate, 4—6 cm long and 3-6 mm
wide, copiously and harshly hirsute on both
surfaces but not glandular, the apex acute or
obtuse, mucronulate, the margins entire and
scabrous, the base rounded or slightly clasping
but not auriculate; rameal leaves similar in
contour and vestiture, those of the peduncles few,
oblong-lanceolate, 3—5 mm long, not intergrading
with the phyllaries. Capitulescence an ample
leafy panicle with ascending branches and a
pyramidal or round top. Flowering heads
intermediate to the two parent species in most
characteristics, 1.3—2 cm in diameter when the
rays are fully extended, often crowded, some-
times secund. Involucre hemispherical, 4-6 mm
high. Phyllaries strongly reflexed, somewhat
graduated or often subequal, scabrous-puberulent
but not glandular, mostly with a conspicuous,
scarious basal portion. Ray florets 20—30 (or
more), the corollas 5-10 mm long, lavender. Disk
4-7(8) mm in diameter. Disk florets 20-30 (or
more), the corollas 3—4 mm long. Pappus tawny
or rose-tinged. Achenes fusiform or obconic,
1.5—2 mm long, dull purple or brown, densely
sericeous but not glandular, 7—9 ribbed. 2” = 10.

Vol. 34 Art. 2

Collections of this hybrid have been recorded
in habitats shared with the parental species from
Champaign, Cook, De Witt, Du Page, Fulton,
McHenry, Menard, Peoria, Piatt, Richland,
Vermilion, and Winnebago counties. The
specimens show considerable variability and are
probably not all first generation hybrids.

15. Aster oblongifolius Nutt.
Aromatic Aster

Herbaceous, with a
tangled, often stoloni-
form or sometimes cau-
diciform rhizome system.
New shoots usually arising
from rhizome strands (rarely
at the base of old stems), the
plants forming scattered
sprawling clumps or small colonial
stands. Stems |-several, erect or de-
cumbent, brittle, 15—70(100) cm tall, with
ascending or divaricate branches from below the
middle, variably hispidulous-hirsute or hirtellous
and also glandular, more copiously so on the
branches, the lower stem usually lacking glands.
Leaves numerous, at least some of the larger
cauline ones persistent throughout the flowering
period; basal rosette leaves oblanceolate or
spatulate, 2—5(7) cm long and 0.5—1.5 cm wide,
with I—3 pairs of secondary nerves aligned with
the midrib or the margins, hirsute, often also
glandular (rarely glabrous or nearly so), the apex
rounded and with a short mucro, the margins
entire and coarsely ciliate (sometimes remotely
toothed); principal leaves sessile, oblong or
linear-lanceolate, (1.5)3—10 cm long and
0.5—1.5(2) cm wide, usually with 1-3 pairs of
weakly expressed and anastomosing secondary
nerves, and a reticulum of tertiary veins with
more or less isodiametric areolae, the upper
surface usually scabrous and often glandular, the
lower copiously hirsute, the apex acute or obtuse,
spinulose-mucronulate, the margins entire and
scabrous-ciliolate, the base rounded or slightly
clasping but not auriculate, often with axillary
clusters of smaller leaves; upper cauline and
rameal leaves gradually reduced in size, similar
in contour, copiously invested with both sessile

May 1989

and stipitate glands; leaves of the peduncles
bracteiform, densely spaced, several to many,
linear or oblong, 3-8 mm long, spinulose, usually
spreading or squarrose, rarely ascending, more or
less intergrading with the phyllaries. Capitules-
cence an often ample, leafy, diffuse panicle
commonly branching from the lower nodes.
Flowering heads |.5—3 cm in diameter when the
rays are fully extended, usually not secund, on
glandular-hispidulous peduncles 0.5—5 cm in
length. Involucre campanulate (or hemispherical
when pressed and dried), 5—7(8) mm high,
imbricated in 4-6 series. Phyllaries not strongly
graduated, sometimes subequal, 3.5—7(8) mm
long, densely glandular on both surfaces and
along the margins, those of the outer 2 or 3 series
with strongly squarrose tips; outermost phyllaries
sometimes largely herbaceous, oblanceolate,

acute; median phyllaries green in the apical '/:—'/2

portion, linear-lanceolate, long-acuminate or
attenuate, the margins erose-hyaline or ciliolate,
the basal '/2—7/: portion scarious; innermost
phyllaries very slender. Receptacle alveolate
with sharp teeth. Ray florets (20)25—35, the
corollas lavender to deep purple (rarely white),
10-15 mm long, glabrous or nearly so. Disk
florets 30-40 (or more), the corollas narrowly
funnelform to nearly tubular, 4.5—6 mm long,
glabrous or nearly so, the limb weakly delimited,
deep yellow before anthesis aging to reddish
purple, the lobe/limb fraction 0.18—0.2, the tube
ca half as long as the limb. Pappus simple, the
bristles somewhat shorter than the disk corolla,
tawny or commonly rose-tinged, soft, slender,
and attenuate. Achenes fusiform or obovoid,
often slightly falcate, 2-2.5 mm long, 0.5—0.8
mm across, dull purple at maturity or brown
when weathered, strigillose or sericeous, with
7-10(11) straw-colored ribs. 27 = 10, 20.

Late August—October. Dry sandy, loamy, or
rocky soils in mostly open habitats: dunes, hill
prairies, open-wooded bluffs, and sandstone or
calcareous cliffs. Locally abundant in uplands
associated with larger rivers. The range of the
species extends from PA south to AL, west to w.
ND, s.c. CO, n.e. NM, and n.c. TX.

Most if not all the plants in Illinois belong in
typical var. oblongifolius [including Aster
oblongifolius var. angustatus Shinners (1941 )—
Gleason 1952; Gleason and Cronquist 1963;
Steyermark 1963]. Occasional plants seem to
verge toward the southwestern var. rigidulus A.
Gray [A. Kumleini (sic) Fries ex A. Gray (1884),

ASTER AND BRACHYACTIS IN ILLINOIS 167

pro syn.], which is distinct by being compara-
tively low-stemmed with numerous firm,
relatively small leaves and by having a notably
fastigiate branching habit.

16. Aster ontarionis Wieg.
Ontario Aster

Herbaceous, with a
creeping horizontal
rhizome system. New
shoots initiated at the tips
of stoloniform rhizomes, the
plants forming colonies but
the connecting strands short-
lived and mostly disintegrating
after one or two seasons. Stems
single (rarely 2 or 3) at any point of
emergence, 40—120 cm tall, erect, with ascend-
ing or divaricate branches from near or below the
middle and also in midsummer with short
secondary shoots composed of smaller leaves
plus a few heads arising from the leaf axils, the
indument villous or hirsute, uniformly distrib-
uted on the branches, often in decurrent lines in
the middle portion of the stem, the lower portion
glabrescent. Leaves polymorphic; vernal rosette
leaves with spatulate to oblanceolate-obovate
blades, sometimes purplish below, minutely
puberulent or rarely glabrous on both surfaces,
the apex acute to rounded, the margins crenate-
serrate, abruptly narrowed to a winged, ciliate,
somewhat sheathing subpetiolar base; principal
cauline leaves usually persistent throughout the
flowering period, sessile or subsessile, oblanceo-
late or elliptic-lanceolate, 2—8(12) cm long and
0.53.5 cm wide, gradually reduced in size
upward on the stem, with several pairs of
anastomosing secondary nerves, and with a
weakly expressed reticulum of tertiary veins with
oblong areolae, scabrellous above, softly (often
minutely) puberulent over the entire lower
surface, and sometimes densely villous along the
midrib below, the apex acute or acuminate to
short-attenuate, usually callus-pointed, the
margins crenate-serrate and ciliolate, the base
cuneate and somewhat decurrent; rameal leaves
similar in contour and vestiture but entire and
notably unequal in size, those subtending the

168 ILLinois NATURAL History SURVEY BULLETIN

head 1—few, linear-lanceolate, |-3 mm long,
flexible, not phyllarylike. Capitulescence an
often ample, diffuse or elongate panicle above the
middle of the stem, the head-bearing branchlets
ascending or spreading but not strongly arching
and typically not racemiform. Flowering heads
0.8—1.5 cm in diameter when the rays are fully
extended, often crowded but typically not secund,
subsessile or on peduncles 0.2—1(2) cm in length.
Involucre campanulate (turbinate when pressed
and dried), 3-4.5(5) mm high, the phyllaries
imbricated in 3—S series. Phyllaries appressed or
somewhat spreading, strongly graduated, the
outer ones 1—2 mm long, less than half as long as
the inner, slenderly linear, the median ones
somewhat expanded toward the tip, the green
areoles linear-oblanceolate, often extending to the
base, both surfaces slightly puberulent, the apex
acute or acuminate and callus-pointed, the
margins irregularly ciliolate, hyaline, and with a
scarious rim extending to the tip; innermost
phyllaries glabrous, very slender and attenuate.
Receptacle alveolate with sharp teeth. Ray
florets 15—25, the corollas 4-8 mm long, white,
glabrous or nearly so. Disk florets 12—20 (or
more), the corollas funnelform, (2.5)3—4.5 mm
long, glabrous, the limb strongly flared, cream-
colored or light yellow before anthesis turning
magenta or purple, the lobes long and reflexed,
the lobe/limb fraction 0.5—0.6, the slender tube
slightly shorter than the limb. Pappus simple,
the bristles about as long as the disk corolla
whitish, soft, slender, and attenuate. Achenes
oblong-obovoid, plump or slightly compressed,
1.2-1.8(2) mm long, 0.4—0.6 mm across, gray,
strigillose, and often puncticulate (caused by the
pustulate trichome bases), with 3—5 ribs. 2n = 32.
[A. missouriensis Britton in Britton & Brown non
(Nutt.) Kuntze, including var. thyrsoides (A.
Gray) Wieg.—Deam 1940. A. lateriflorus var.
hirsuticaulis misapplied, not of (Lindley)
Porter—Pepoon 1927, at least in part. A. panto-
trichus S.F. Blake (see Shinners 1949)—G.N.
Jones 1945.]

September—October. Moist ground habitats:
river bottoms, creek margins, bogs, and marshes,
frequently in wooded areas. Common throughout
the state. The species range extends in the East
from s.w. PQ, s.w. ON, and NY south to AL, and
at the western limit from MN and n.e. SD
through e. NE and s.e. KS toe. TX and n. LA.

Some of the specimens examined suggest
intergradation with other species that share the

Vol. 34 Art. 2

basic chromosome number of x = 8, e.g., Aster
fragilis, A. lanceolatus, and A. lateriflorus. A
considerable number of herbarium specimens
have been misidentified as A. /ateriflorus. If
rhizomes are lacking, the two species are
sometimes not readily distinguishable.

17. Aster oolentangiensis Riddell
Sky-blue Aster, Azure
Aster

Herbaceous, with short
horizontal rhizomes or a
branched-caudiciform
system. New shoots :
arising at the base of old (le
stems or from rhizome branches, /|
the plants forming scattered .
individual clumps. Stems
l-several, erect, (30)60—100 cm tall, \#
typically with stiff, ascending NY,
branches above the middle, nearly glabrous

or with hispidulous lines in the leaf axils and
along decurrent leaf bases, the head-bearing
branchlets slightly ridged, often uniformly
scabrous-puberulent. Leaves polymorphic, the
basal and lower cauline ones petiolate, commonly
persistent throughout the flowering period; blades
of basal rosette and larger cauline leaves ovate to
ovate-lanceolate, 4-15(18) cm long and 1+4(6)
cm wide, with 2—several pairs of anastomosing
secondary nerves, and a reticulum of tertiary
veins with more or less isodiametric areolae, the
upper surface scabrous, the lower softly hirsute,
the apex acute to acuminate, the margins
subentire to shallowly serrate, the base rounded,
cordate, or truncate, the petioles twice as long

(or more) as the blades, slightly winged, ciliate,
dilated toward the sheathing base; leaves of the
midstem subpetiolate or sessile, ovate-lanceolate
(in Illinois plants), gradually or somewhat
abruptly reduced, acute or attenuate, and callus-
pointed, the margins entire and scabrous, the base
cuneate and somewhat decurrent; rameal leaves
abruptly reduced in size, linear-lanceolate or
subulate, appressed or closely ascending, those of

the peduncles bracteiform, densely spaced,
1.5—3(4) mm long, scabrous-margined, inter-
grading with the phyllaries. Capitulescence
typically a narrow thyrsiform panicle in the upper

May 1989

'/:/: portion of the stem, the branches ascend-
ing, sometimes racemiform. Flowering heads
1.5—2 cm in diameter when the rays are fully
extended, sometimes secund, the peduncles
variable in length to 6 cm, densely bracteate.
Involucre cylindric to campanulate (often
turbinate when pressed and dried), 5—7 mm high,
the phyllaries imbricated in 4 or 5(6) series.
Phyllaries strongly graduated, appressed or those
of the outer series slightly spreading, glabrous,
the outer ones |.5—2 mm long, less than half as
long as the inner, linear-lanceolate to subulate,
the median and inner ones linear or often slightly
expanded toward the apex, the apical green
areoles rhombic, usually shorter than the
indurate, scarious basal portion of the phyllaries,
the apex sharply acute, acuminate, or obtuse, and
mucronulate, the margins erose-hyaline and
somewhat ciliolate. Receptacle alveolate with
sharp teeth. Ray florets (12)15—20, the corollas
8-12(14) mm long, commonly blue or violet-
purple (rarely white), glabrous. Disk florets
15-25 (or more), the corollas narrowly fun-
nelform, 44.5 mm long, glabrous, the limb light
yellow turning deep purple after anthesis, the
lobe/limb fraction 0.2—0.25, the tube considerably
shorter than the limb. Pappus simple, the bristles
about as long as the disk corolla, cream-colored
or slightly rose-tinged, soft, slender, and
attenuate. Achenes oblong-obovoid, somewhat
compressed, |.8—2 mm long and less than | mm
across, dull purple, or straw-colored with purple
streaks, glabrous or nearly so, with 4 or 5 ribs.

2n = 32; reported chromosome counts of 2” = 36
(under the name A. azureus) are probably in error
(A. G. Jones 1977). [A. azureus Lindley in
Hooker—most authors prior to this publication
(see A.G. Jones 1983).]

August—October. Dry sandy, loamy, or rocky
soils in relatively open or unshaded situations:
prairie remnants, open woods, bluffs, dunes, and
barrens. Locally common throughout the state.
The species range extends from s. ON and w. NY
west to MN and e. SD and south through the
central states to n.c. FL, LA, and n. TX.

Plants in Illinois belong in typical var.
oolentangiensis (cf. A.G. Jones 1983). There is
some suggestion of intergradation with and gene
flow from other species that share the basic
chromosome number of x = 8, e.g., Aster
drummondii and A. laevis.

ASTER AND BracHyAactis tN ILLINOIS 169

18. Aster parviceps (Burgess in Britton &
Brown) Mack. & Bush
Small-headed Aster

Short-lived herbaceous
perennial with a
caudiciform rhizome.
New shoots originating
at the base of old stems or
often from root sprouts
produced in autumn of the
previous season, the plants
forming small scattered individual
clumps. Stems |—few, slender, erect,
20-70(90) cm tall, with ascending branches in
the upper '/;—'/> portion, variably pilose to
hirsute, the indument uniformly distributed or
sometimes in decurrent lines from the leaf bases,
the lower stem glabrescent. Leaves polymorphic,
the basal and larger ones commonly withered
and deciduous at flowering time; basal rosette
leaves subpetiolate, oblanceolate or spatulate,
1+ cm long and 3—7 mm wide, with weakly
expressed anastomosing secondary nerves,
glabrous or sparsely pubescent, and often
purplish on the lower surface, the apex obtuse
and bristle-tipped, the base sheathing; cauline
leaves sessile, often with clusters of smaller
leaves in the axils, those of the midstem linear-
oblanceolate to lanceolate, 4-8 cm long and 2—5
mm wide, thinly pilose above, puberulent to
hirsute below or sometimes nearly glabrous, the
apex acute to attenuate and usually armed with a
hyaline bristle, the margins entire or shallowly
serrate, ciliolate; rameal leaves copious, mostly
subulate and spinulose, those of the peduncles
bracteiform, appressed or ascending, 2—4 mm
long, scabrous-margined, intergrading with the
phyllaries. Capitulescence a narrow panicle,
with a pyramidal or flat top, in the upper '/:—'/2
portion of the stem, the branches ascending or
sometimes arching, typically racemiform.
Flowering heads small, 7—10 mm in diameter
when the rays are fully extended, secund, the
peduncles densely bracteate, 0.3—2(4) cm long.
Involucre cylindric to narrowly campanulate
(turbinate when pressed and dried), 34.5 mm
high, the phyllaries imbricated in 3—S series.
Phyllaries graduated, appressed when fresh,
glabrous, the apical green areoles lance-rhombic;
outer phyllaries subulate, 1.5—2 mm long, ca half

170 ILtinots NATURAL History SURVEY BULLETIN

as long as the inner; median and inner phyllaries
linear-lanceolate, attenuate or acuminate with
marginally inrolled spinulose tips, the margins
erose-hyaline and commonly slightly ciliolate,
the basal '/:—'/: portion scarious. Receptacle
alveolate with sharp teeth. Ray florets
10—16(18), the corollas ca 5 mm long, white,
glabrous. Disk florets 8—10(15), the corollas
narrowly funnelform, (2)2.5—3 mm long,
glabrous, the limb light yellow turning purple
after anthesis, the lobe/limb fraction 0.25, the
tube shorter than the limb. Pappus simple, the
bristles about as long as the disk corolla, white,
soft, slender, and attenuate. Achenes oblong,
plump, 0.8-1.5 mm long and 0.4—0.6 mm across,
whitish or gray, puberulent, obscurely 24
ribbed. 2n = 16, 32. [A. tenuifolius misapplied,
not of L—Mead 1846. A. pilosus Willd. subsp.
parviceps (Burgess in Britton & Brown) A.G.
Jones (1984).]

August-October. Sandy or loamy soils in
mostly unshaded habitats: barrens, hill prairies,
rarely in somewhat disturbed ground that still
supports prairie vegetation. Not common; mostly
occurring in the western half of the state but also
recorded from Cook and Will counties. As
delimited herein [i.e., not including Aster
depauperatus (Porter) Fern.], the species range
extends from IA and IL south to MO, n. AR, and
s.e. KS, with one station recorded from n.e. OK.

Aster parviceps is very closely related to A.
pilosus and was originally described as a variety
of that species (under the name A. ericoides
parviceps Burgess in Britton & Brown). For the
time being and pending further study, I have
reversed my decision (1984) to combine the two
species, mainly because their habitats and
reproductive vigor are strikingly different.
Furthermore, there seems to be a difference in the
predominant ploidy levels. Based on x = 8
chromosomes, plants of A. parviceps are recorded
as being mostly diploid or sometimes tetraploid
(Semple and Chmielewski 1985), whereas those
of A. pilosus are generally hexaploid, at least in
my experience.

Vol. 34 Art. 2

19. Aster patens Aiton
Spreading Aster. Late
Purple Aster

Herbaceous, with a
branched-caudiciform,
often tangled or some-
times cormoid rhizome
system, but also producing
stoloniform rhizome strands.
New shoots initiated at the
base of old stems or from the
rhizomes, resulting in scattered
l—several stemmed individuals or
small patches. Stems erect. brittle, often stout,
50-100(120) cm tall, with stiff divaricate or
ascending branches above the middle, the
indument uniformly distributed around the stem,
variously scabrous-hirsute to cinereous-pube-
rulent, or villous on the branches and peduncles.
Leaves polymorphic, the basal and lower cauline
ones withered and deciduous at flowering time:
basal rosette leaves spatulate, rugose, with 1+
pairs of arching and anastomosing secondary
nerves, variously scabrous-hirsute, the apex acute
to rounded, the margins entire or often with a few
teeth, the narrowed subpetiolar base winged and
sheathing; principal cauline leaves ovate-
lanceolate to oblanceolate, rarely spatulate, often
constricted above the strongly auriculate-clasping
base, 2—6(10) cm long and 1—2(2.5) cm wide,
grayish green, rugulose on both surfaces, with
usually pinnate and anastomosing secondary
nerves, and a reticulum of tertiary veins with
isodiametric areolae, variously pubescent, more
densely so along the midrib below, the apex acute
or obtuse and with a short mucro, the margins
entire or appearing minutely denticulate from the
pulvinate bases of coarse scabrous cilia; rameal
leaves abruptly reduced in size, those of the
peduncles bracteiform, 1—3(5) mm long,
appressed or sometimes squarrose, intergrading
with the phyllaries. Capitulescence an open,
divaricately branched panicle. Flowering heads
2-3.5 cm in diameter when the rays are fully
extended, not crowded and not secund, typically
at the ends of stiffly ascending or spreading,
densely bracteate peduncles 2—10(15) cm in
length. Involucre campanulate to turbinate,
5-8(10) mm high, the phyllaries imbricated in

May 1989

5-8 series. Phyllaries strongly graduated, linear
to ovate-lanceolate, appressed or often slightly
recurved-spreading, strigillose or cinereous-
puberulent on the abaxial surface and near the tip
on the adaxial surface, the apical portion also
frequently invested with both sessile and short-
stipitate glands; outer phyllaries 1.5—2.5 mm, the
inner (4)6—7 mm long; median phyllaries acute or
acuminate to obtuse, the chlorophyllous areoles
grayish green, rhombic-oblanceolate, poorly
delimited in the apical ('/:)'/:—'/2 portion, mostly
shorter than the scarious basal portion. Recep-
tacle shallowly alveolate with sharp or rounded
teeth. Ray florets (12)15—25 (or more), the
corollas typically blue or deep purple (rarely
white), 10—15(17) mm long, glabrous or with a
few scattered trichomes. Disk florets 20—S0, the
corollas narrowly funnelform, 4.5—6 mm long,
glabrous or nearly so, the limb light yellow
turning purple after anthesis, the lobe/limb
fraction 0.18—0.2, the tube shorter than the limb.
Pappus simple, the bristles about as long as the
disk corolla or slightly shorter, tawny, soft, and
attenuate. Achenes oblong-obovoid or fusiform,
plump, 2.5-3.5 mm long and 0.8—1.2 mm across,
dull purple or brown, sericeous or strigillose,
with 7-10 ribs. 2 = 10, 20 (Illinois plants).
Including A. patentissimus Lindley in Hooker
(see R.L. Jones 1983).

September—October. Dry sandy or rocky
soils: edges of Ozarkian forest lands, open-
wooded bluffs, sandstone and limestone ridges;
also in disturbed soils of clearings and old fields.
Occasional in the southern one-third of the state,
north to Jersey, Montgomery, and Crawford
counties. The range of the species extends from
New England to n. FL, west to s. IL, s. and c.
MO, s.e. KS, and the eastern half of OK and TX.

Two sympatric varieties have been recorded
from Illinois with approximately equal frequency,
var. patens and var. patentissimus (Lindley in
Hooker) Torrey & Gray. The taxa intergrade
freely but may be distinguished as follows:

1. Involucre 8-10 mm high, broadly campanu-
late or turbinate; median phyllaries ovate-
lanceolate, 1.2—1.5:mm in width, obtuse,
appressed, copiously cinereous-strigillose on
the back, not conspicuously glandular, or
toward the apex with minute sessile glands
that are obscured by the indument.................-

= cep EUROS CEL GSE PRCEEEOSE CELE CLAPPED" var. patentissimus

ASTER AND BRACHYACTIS IN ILLINOIS 171

1. Involucre usually less than 8 mm high,
slenderly campanulate or turbinate; median
phyllaries linear-lanceolate, mostly less than
1.2 mm in width, acute or acuminate,
somewhat recurved-spreading, sparsely
strigillose on the back mainly along the
midrib, conspicuously glandular with both
sessile and short-stipitate glands ...................

var. patens

The var. patentissimus was originally
described (as Aster patentissimus) from a
Missouri collection and is, according to Steyer-
mark (1963), the more common form of
A. patens in the Ozarks. Plants of typical var.
patens are more prevalent to the east. Occasional
herbarium collections seem to verge morpho-
logically toward the small-leaved and slender-
stemmed southern var. gracilis Hooker.
However, plants of the latter variety are largely
diploid [2n (= 2x) = 10], and that ploidy level has
not been recorded for Illinois collections (A.G.
Jones 1980b; R.L. Jones 1983). Specimens
forming the basis for records from the Chicago
region (Higley and Raddin 1891; Pepoon 1927),
including A. patens |var.] phlogifolius sensu
Higley &Raddin and Pepoon non (Muhl. ex
Willd.) Nees, do not belong in this species.

20. Aster pilosus Willd.
Hairy Aster, Frost-weed
Aster

Herbaceous, with a
caudiciform rhizome.
New shoots arising at
the base of old stems or
from root sprouts produced
in the previous season, the
plants forming scattered
individual clumps. Stems
l-several, stout, erect, 30-120 cm
tall, with ascending or divaricate
branches from near or below the middle,
variously pilose to hirsute, the indument in
decurrent lines from the nodes or often uni-
formly distributed, sometimes the stems glabrous
or nearly so. Leaves polymorphic, the basal and
larger cauline ones commonly withered and
deciduous at flowering time; rosette leaves

172 ILLinois NATURAL History SuRVEY BULLETIN

spatulate, the blade 1-6 cm long and 0.5—1 cm
wide, abruptly narrowed to a winged subpetiolar
portion, with 1 or 2 pairs of secondary nerves
arching toward the apex, glabrous, or the lower
surface thinly pilose along the midrib and often
purplish, the apex rounded or obtuse, the margins
softly ciliate or scabrous and often shallowly
serrate, the base sheathing; cauline leaves sessile
or subsessile, often with clusters of smaller
leaves in the axils, the principal ones elliptic-
oblanceolate to linear-lanceolate, 4-12 cm long
and 0.5—2.5 cm wide, gradually reduced in size
upward on the stem, with a prominent midrib and
1-several pairs of anastomosing secondary
nerves, glabrous to variously pilose or hirsute
(more so along the nerves below), the apex
attenuate and usually armed with a hyaline
spinule, the margins serrate or often entire, softly
ciliate, the base cuneate, somewhat sheathing and
decurrent; rameal leaves copious, subulate, those
of the peduncles bracteiform, often densely
spaced, appressed to ascending or sometimes
spreading, 3-6(10) mm long, ciliolate, more or
less intergrading with the phyllaries. Capitules-
cence an ample, leafy, usually diffuse panicle, the
branches commonly racemiform. Flowering
heads 1.5—2(2.5) cm in diameter when the rays
are fully extended, often crowded and secund,
subsessile, or on densely bracteate peduncles
(0.5—3(5) cm in length. Involucre campanulate or
broadly urceolate (i.e., somewhat constricted
below the slightly recurved phyllary tips),
3.5-5(6) mm high, the phyllaries imbricated in 4
or 5 series. Phyllaries commonly at least
somewhat graduated (rarely subequal in length),
appressed or slightly spreading, glabrous except
for a few marginal cilia near the apex; outer
phyllaries subulate, 2—3(4) mm long, sometimes
largely herbaceous, the median and inner ones
linear-oblanceolate, 3—5(5.5) mm long, with
broadly rhombic to lance-rhombic green areoles
in the apical '/;—'/: portion, the scarious basal
portion somewhat indurate and rounded on the
back, the apex with sharply acute or marginally
inrolled acuminate tips, spinulose, the margins
erose-hyaline. Receptacle shallowly alveolate
with sharp teeth. Ray florets 15—30, the corollas
7-9(10) mm long, white (rarely pink), glabrous.
Disk florets 30-40 (or more), the corollas
narrowly funnelform, (3)3.5—4.5 mm long,
glabrous or nearly so, the limb cream-colored or
light yellow turning lavender or reddish purple
after anthesis, the lobe/limb fraction 0.25—0.3, the

Vol. 34 Art. 2

tube much shorter than the limb. Pappus simple,
the bristles about as long as the disk corolla,
white, soft, slender, and attenuate. Achenes
oblong, plump, 1-1.5 mm long and 0.5—0.7 mm
across, whitish or gray, minutely puberulent, and
obscurely 2-4 ribbed. 2n = 32, 40 (rarely), 48
(mostly). Including A. pringlei (A. Gray) Britton
in Britton & Brown. [A. ericoides misapplied, not
of L._—Brendel 1887; Higley and Raddin 1891;
Pepoon 1927; Benke 1928; Kibbe 1952. in part.
A. villosus Michaux non Thunb. A. ericoides var.
villosus (Michaux) Torrey & Gray—Higley and
Raddin 1891; Pepoon 1927; Benke 1928. A.
polyphyllus Willd. non Moench—Higley and
Raddin 1891; Pepoon 1927.]

August—October. Disturbed ground in mostly
unshaded habitats: roadsides, railroad rights-of-
way, open woods, and pastures. Very common,
weedy, recorded from every county. The range
extends from s. PQ and s. ON south to GA and
n. FL and at the western limit from e. SD through
e. NE, e. KS, and e. OK to LA.

Several varieties of Aster pilosus recognized
in widely circulated floristic manuals (e.g., Deam
1940; Fernald 1950; Gleason 1952; Gleason and
Cronquist 1963; Steyermark 1963; and Cronquist
1980) occur in Illinois: 1) var. pilosus, 2) var.
platyphyllus (Torrey & Gray) S.F. Blake [A.
ericoides var. platyphyllus Torrey & Gray—
Pepoon 1927], 3) var. demotus S.F. Blake, and 4)
var. pringlei (A. Gray) S.F. Blake [A. ericoides
var. pringlei A. Gray (1884). A. pringlei—Jones
and Fuller 1955; G.N. Jones 1963; Mohlenbrock
1986]. The type collection for A. pilosus (sensu
stricto, i.e., for var. pilosus) is from Illinois
(Jones and Hiepko 1981); it was originally named
by Michaux and is also the type for A. villosus
(Jones and Lowry 1986). The relatively broad-
leaved, densely villous to pilose plants of var.
platyphyllus (accepted by Deam 1940; Dobbs
1963; Steyermark 1963) can be, in my opinion,
produced under the influence of certain environ-
mental conditions. For the most part, they fit
quite well in typical var. pilosus, a view also
taken by others (A. Gray I884; Gleason 1952:
Gleason and Cronquist 1963; Cronquist 1980).
Recent studies suggest that vars. pringlei and
demotus perhaps should be combined under the
former name (Semple and Chmielewski 1985; see
also Mohlenbrock 1975). My own observations
tend to support this concept (including here also
A. polyphyllus). The variety 1s characterized as
being nearly glabrous and having relatively

May 1989

narrow leaves; these plants are fairly common
throughout the state. When taken in the narrow
sense, however, var. pringlei is restricted in
Illinois to Lake County (Swink 1974; Mohlen-
brock and Ladd 1978; Swink and Wilhelm 1979).
A great deal of intergradation between the above
varieties can be found in herbarium specimens
and observed in the field. I have not attempted,
therefore, to elaborate on keys in the literature
dealing with varieties of this variable species
(see Deam 1940; Fernald 1950; Gleason 1952;
Gleason and Cronquist 1963; Steyermark 1963;
and Cronquist 1980).

21. Aster praealtus Poir.
Willow-leaved Aster,
Willow Aster

Herbaceous, with a
strongly creeping
rhizome system, the
connecting strands often
persistent for several
seasons. New shoots mostly
originating at the tips of
fleshy stoloniform rhizome
strands, the plants forming
extensive colonies. Stems
commonly single at any point of
emergence, stout, erect, sometimes slightly
reddish or glaucous, (40)60—150 cm tall, with
ascending branches above the middle, variously
hispidulous to hirsute, the indument in decurrent
lines from the nodes or sometimes uniformly
distributed around the stem, especially on the
branchlets, the lower stem portion glabrescent.
Leaves polymorphic, the basal and lower cauline
ones commonly withered and deciduous at
flowering time; basal rosette leaves fleshy,
spatulate, 4-7 cm long and 1—2.5 cm wide, deep
green above and often purplish below, with
several pairs of anastomosing secondary nerves,
minutely scabrous above, glabrous below, the
apex rounded, the margins entire or shallowly
serrate, ciliate on the narrowed subpetiolar
portion, the base dilated and sheathing; principal
cauline leaves firm, sessile, elliptic to linear-
lanceolate, 4—12(15) cm long and 0.3—1.5 cm
wide, gradually reduced in size upward on the
stem, rugulose, with indistinct secondary nerves
but a conspicuous reticulum of brownish tertiary

ASTER AND BRACHYACTIS IN ILLINOIS 173

veins (notable on the lower surface), enclosing
isodiametric areolae, the upper surface com-
monly scabrous at least near the margins,
sometimes strongly so, rarely glabrous, the lower
surface mostly glabrous (in Illinois plants) or
sometimes puberulent, the apex acute to
attenuate with a sharp callus point, the margins
frequently inrolled, entire or shallowly serrate,
and scabrous, the base gradually tapered, often
slightly rounded, decurrent; leaves of the upper
stem and branches usually with clusters of
smaller leaves (plus a few heads) in the axils,
therefore numerous and notably unequal in size,
oval to linear-lanceolate, acute or obtuse, callus-
pointed, scabrous-margined; leaves of the
peduncles relatively few, similar in contour,
ascending or recurved-spreading, (2)4—-10 mm
long, flexible, not intergrading with the phyllar-
ies but often closely subtending the head.
Capitulescence a mostly ample, leafy, slenderly
thyrsiform or broadly diffuse panicle in the
terminal '/;—'/> stem portion, the head-bearing
branchlets rarely racemiform. Flowering heads
1.5—2(2.5) cm in diameter when the rays are
fully extended, usually not secund, on peduncles
0.3—2 cm (or more) in length. Involucre
campanulate, (4)5—7(8) mm high, the phyllaries
imbricated in 4 or 5(6) series. Phyllaries
appressed or with slightly recurved tips, at least
somewhat or often strongly graduated, the outer
ones 2—3(3.5) mm, the inner (4)5—6.5 mm long:
outer and median phyllaries oblanceolate,
somewhat constricted above a slightly dilated
base, the green areoles oblanceolate or lance-
rhombic, or sometimes the outer phyllaries
largely herbaceous, the abaxial surface glabrous,
the adaxial surface sparsely puberulent, the apex
acute to acuminate and often with a reddish
callus point, the margins ciliolate, erose-hyaline,
and with a narrow scarious rim extending nearly
to the tip, the basal '/2—'/: portion scarious;
innermost phyllaries very slender, linear, with
linear-oblanceolate, light green areoles, the apex
acuminate or attenuate. Receptacle alveolate
with sharp teeth. Ray florets (15)20—30 (or
more), the corollas 7—10(12) mm long, typically
lavender to rose-purple (rarely white), glabrous.
Disk florets 25—35 (or more), the corollas
tubular or narrowly funnelform, 4—-6.5 mm long,
the limb cream-colored or light yellow turning
purple after anthesis, the lobe/limb fraction
0.18—0.2, the tube much shorter than the limb.
Pappus simple, the bristles often slightly longer

174 ILtinois NATURAL History Survey BULLETIN

than the disk corolla, whitish, soft, slender, and
attenuate. Achenes oblong-obovoid, plump or
slightly compressed, 1.5—2 mm long and 0.5—0.8
mm across, purple, or straw-colored with purple
streaks, thinly strigillose, with 4 or 5 ribs. 2n =
32. Including A. subasper Lindley in Hooker (see
Wiegand 1933). [A. salicifolius Aiton and sensu
auct. non Lam.—Brendel 1887; Higley and
Raddin 1891; Pepoon 1927; G.N. Jones 1945,
1950; Fell 1955. A. carneus misapplied, not of
Nees—Mead 1846, and many herbarium sheets
so labelled.]

September—October. Low moist or swampy
ground: woods, thickets, meadows, banks of
streams and rivers, ditch margins, and lakeshores.
Common throughout the state. The range of the
species extends in the eastern half from ON, MI,
and PA to n. FL and in the western half from s.
MB through ND, SD, NE, and OK to TX and n.
Mexico.

The plants are variable, but attempts to
distinguish the varieties that have been described
(Wiegand 1933) are not very successful. The
reason is a high degree of morphological
intergradation and an apparent lack of convincing
geographic range separation. The majority of
Illinois collections can be identified with typical
var. praealtus [including var. angustior Wieg.,
pro parte (and in the sense of some authors and
collectors)—Deam 1940; Steyermark 1963]. The
var. subasper (Lindley in Hooker) Wieg.,
originally described (as Aster subasper) from the
St. Louis area, has been recognized by several
authors (Deam 1940; Steyermark 1963; Swink
1974; Swink and Wilhelm 1979). Plants of this
variety are fairly common; they are more readily
identifiable by their copiously scabrous leaves
and stems than by the characters used to define
the taxon in the keys of Deam (1940) and
Steyermark (1963), namely cauline leaves that
are relatively short and elliptic, and rameal leaves
that are mostly oval and obtuse. Good evidence
exists of occasional hybridization and intergrada-
tion with A. lanceolatus [e.g., V. H. Chase 137
(F) from Stark Co., Evers 109660 (ILLS) from
Johnson Co., A.G. Jones 4616 (ILL) from De Witt
Co.] and with A. puniceus var. firmus (Nees)
Torrey & Gray [e.g., Winterringer 1626] (ISM)
from Cook Co., Wolf /70 (F) from Fulton Co.,
Wade & Wade 2383 (F) from Ogle Co., and
others].

Vol. 34 Art. 2

22. Aster prenanthoides Muhl. ex Willd.
Crooked-stemmed Aster,
Crooked Aster

Herbaceous, with a
creeping rhizome
system. New shoots
produced at the tips of
stoloniform rhizome
strands, the plants typically
colonial. Stems mostly single
at any point of emergence,
erect, (40)60—120 cm tall, the
older ones often dark purple,
branched near or commonly above the middle,
the branches zigzag, somewhat angled in cross
section from decurrent leaf bases, villous or
hirsute in decurrent lines, the indument some-
times uniformly distributed on the peduncles, the
lower portion of the stem glabrescent or glabrous.
Leaves polymorphic, those of the basal rosettes
subpetiolate, the blades obovate to oblanceolate,
1.5—7 cm long and 1-2 cm wide, with several
pairs of anastomosing secondary nerves,
scabrellous above or glabrous on both surfaces,
acute or obtuse, crenate to crenate-serrate,
abruptly narrowed to a slender or slightly winged
petiolelike portion, with long marginal cilia and
an often reddish, dilated, sheathing base:
principal cauline leaves mostly persistent
throughout the flowering period, subsessile or
sessile, oblanceolate in outline but abruptly and
strongly contracted near or below the middle,
8—16(20) cm long and 1.5—5.5 cm wide,
gradually reduced in size upward on the stem, the
blades with 6-10 (or more) pairs of anastomosing
secondary nerves; nerves in the wider, apical
portion pinnate, those in the narrower, basal
portion running parallel to the midrib, the
reticulum of tertiary veins weakly expressed with
irregular areolae, the upper surface scabrellous,
the lower minutely strigillose or glabrous but
commonly hispidulous or villous along the
midrib, the apex long-acuminate with a callus
point, the margins sharply serrate, with callus-
pointed teeth on the wider blade portion but
commonly entire on the narrowed portion, the
base dilated and strongly auriculate-clasping, the
midrib extended on the stem as a decurrent ridge;

May 1989

rameal leaves similar in most aspects but often
only slightly contracted near the middle, glabrous
or nearly so, the margins less sharply toothed or
often entire, the base sessile and clasping; leaves
of the peduncles relatively few, lanceolate, 3—12
mm long, not intergrading with the phyllaries.
Capitulescence a broad, flat- or round-topped,
dichotomously branched panicle in the upper
'/:—/2 portion of the stem, the branches slender,
sometimes purplish. Flowering heads (1.8)2—2.5
cm in diameter when the rays are fully extended,
not crowded and not secund, on slender
peduncles 14 cm in length. Involucre campanu-
late or hemispheric, S—6 mm high, the phyllaries
imbricated in 4 or 5(6) series. Phyllaries often
only slightly graduated, flexible, with recurved or
sometimes reflexed tips, the outer ones mostly
lanceolate and slightly constricted near the
middle, 2.54 mm long, the inner and median
ones linear-oblanceolate, 4S mm long, the green
areoles linear-oblanceolate, or sometimes the
outer phyllaries largely herbaceous, the abaxial
surface glabrous, the adaxial surface slightly
puberulent, the apex acute and callus-pointed, the
margins irregularly ciliolate, erose-hyaline, and
with a narrow scarious rim extending nearly to
the tip, the basal '/;—'/> portion scarious. Recep-
tacle alveolate with blunt teeth. Ray florets
18-25 (or more), the corollas 10-15 mm long,
typically lavender or bluish (rarely white),
glabrous. Disk florets 20—25 (or more), the
corollas funnelform, (3.5)4—5 mm long, often
thinly puberulent near the throat, the limb cream-
colored or light yellow turning purple after
anthesis at least in the lobes, the lobe/limb
fraction 0.2—0.25, the tube slightly shorter than
the limb. Pappus simple, the bristles as long as
or slightly shorter than the disk corolla, some-
what discolored, soft, slender, and attenuate.
Achenes cylindric-oblanceolate or obovoid,
slightly compressed, 2—3(3.5) mm long and
0.5—0.8 mm across, dull purple or straw-colored,
thinly strigillose, 4- or S-ribbed. 2 = 32
(mostly), 48.

Late August—October. Moist or swampy
ground: woods, thickets, meadows, seeps, and
stream banks. Occasional, recorded by me from
only seven counties in Illinois, much more
common to the east. The range of the species
extends from s. ON, NY, and PA south to NC
and TN and west to MN and IA.

ASTER AND BRACHYACTIS IN ILLINOIS 175

23. Aster puniceus L.
Purple-stemmed Aster,
Red-stemmed Aster,
Swamp Aster

ia
a
ia
Ole

Herbaceous, with a
horizontal, short or often
strongly creeping
rhizome system, the con-
necting strands persistent for
several seasons. New shoots
mostly originating at the tips of
fleshy stoloniform rhizome
strands, the plants sometimes
forming extensive colonies. Stems
mostly single at any point of emergence, stout,
erect, commonly purplish or maroon-colored,
50—150(180) cm tall, with ascending or divari-
cate branches in the upper half, variously hirsute
to hispidulous (or both) in decurrent lines from
the nodes, or the indument nearly uniformly
distributed around the stem, the basal portion
sometimes coarsely hispid. Leaves polymorphic,
the basal and lower cauline ones commonly
withered and deciduous at flowering time, but
the head-bearing branchlets often very leafy
(var. firmus); basal rosette leaves spatulate or
oblanceolate, 3-10 cm (or more) long and

0.3—2 cm (or more) wide, dark green and
somewhat rugulose above, often purplish below,
the venation and vestiture similar to that of the
principal leaves, the apex acute to rounded, the
margins remotely crenate-serrate to subentire,
the lower subpetiolar portion of the blade
winged, dilated, and sheathing at the base;
principal cauline leaves variable, firm, sessile,
(6)10—15(20) cm long and (1)2—5 cm wide,
gradually reduced in size upward on the stem,
with several pairs of pinnate and anastomosing
secondary nerves, and a reticulum of tertiary
veins with isodiametric or oblong areolae, the
upper surface scabrous or glabrous, often glossy
(resulting from the presence of bulliform
epidermal cells), the lower surface glabrous or
minutely strigillose and often with scabrous,
hispidulous, or villous indument along the
midrib, the apex acute, acuminate, or attenuate,
with a callus point, the margins shallowly serrate
to subentire, the base typically clasping, often
strongly auricled in the lower leaves, sometimes
merely rounded and decurrent in those higher up

*]
ae
a
peat fc
are

es eal

om
is
Hy

176 ILtinots NATURAL History SurveY BULLETIN

on the stem; rameal leaves often numerous,
elliptic-lanceolate to linear-lanceolate, gradually
reduced, acute to attenuate, callus-pointed, the
vestiture similar to that of the larger leaves, the
margins subentire to entire and often inrolled, the
midrib and larger nerves commonly extending
downward on the stem as decurrent or almost
winged ridges; leaves of the peduncles relatively
few, 0.5—2 cm long, flexible, somewhat con-
stricted above a dilated base, often closely
subtending the head and appearing to be a part of
the involucre, but rarely bracteiform.
Capitulescence an often ample, leafy, broad,
round- or flat-topped, more or less dichotomously
branched panicle in the upper '/:—'/2 stem portion,
the branches ascending or divaricate. Flowering
heads 1.5—3.5(4) cm in diameter when the rays
are fully extended, typically not secund,
subsessile and overtopped by the subtending
rameal leaves, or often on densely hirsute to
villous peduncles 0.2—3 cm (or more) in length.
Involucre campanulate, (6)8—12(15) mm high,
the phyllaries imbricated in 4-6 series. Phyllar-
ies typically not or only somewhat graduated,
6-10(15) mm long, flexible, leaflike, the outer
ones often appreciably longer than the inner and
largely herbaceous; phyllaries of the 2nd and 3rd
series inward lanceolate to linear-oblanceolate,
often spreading or reflexed, somewhat constricted
above a slightly dilated base, glabrous (rarely
slightly scabrous) on the abaxial surface and
thinly puberulent on the adaxial surface, th> 2pex
attenuate or acute to obtuse (depending on the
variety), and callus-pointed, the margins
irregularly ciliolate or lanate, the scarious basal
portion typically much smaller than the green
portion and often indurate or slightly keeled:
innermost phyllaries very slender, linear, with
linear-oblanceolate, light green areoles in the
apical '/> —/; portion, the margins erose-hyaline
and with a scarious rim. Receptacle alveolate
with sharp teeth. Ray florets 20-40 (or more),
the corollas 12—18(20) mm long, lavender to deep
bluish purple (rarely white), glabrous or slightly
puberulent on the throat and tube. Disk florets
(20)30—S0 (or more), the corollas narrowly
funnelform, abruptly dilated at the throat,
(4.5)5—6 mm long, glabrous or with a few
trichomes, the limb cream-colored or yellow
turning pink or purple after anthesis, the
lobe/limb fraction 0.2—0.3, the tube shorter than
the limb. Pappus simple, the bristles about as
long as the disk corolla, whitish or slightly

Vol. 34 Art. 2

discolored, soft, slender, and attenuate. Achenes
oblong or oblanceolate in contour, often
somewhat falcate, plump or slightly compressed,
(2)2.5—3.5(4) mm long and | mm or less across,
purple at maturity or brown when weathered,
thinly puberulent or glabrescent, 4- or 5-ribbed.
2n = 16 (typically), 32. Including A. firmus Nees
and A. lucidulus (A. Gray) Wieg. (see A.G. Jones
1984, 1987). [A. novi-belgii misapplied, not of
L.—Brendel 1887; Higley and Raddin 1891;
Mohlenbrock 1975, 1986. A. longifolius
misapplied, not of Lam.—Pepoon 1927; Fell
1955; Jones and Fuller 1955. A. patens misap-
plied, not of Aiton—Higley and Raddin 1891;
Pepoon 1927.]

(Late August) September—October. Low
moist or swampy ground: woods, thickets,
meadows, banks of streams and rivers, ditch
margins, and lakeshores. Common in the northem
two-thirds of the state, extending south to St.
Clair, Bond, Fayette, and Lawrence counties. The
range of the species (sensu /ato) extends in the
East from NF and LB south to FL, and in the
West from SK and AB to ND, with a few
scattered records from SD, e. NE. IA, MO, and
from Van Zandt and Smith counties in n.e. TX.

The plants are extremely variable, but two
morphologically well-delimited varieties can be
distinguished in Illinois: var. puniceus {including
f. demissus (Lindley) Fern.—Dobbs 1963] and
var. firmus (Nees) Torrey & Gray [Aster firmus—
Mohlenbrock 1986. A. puniceus var. lucidulus
A. Gray—Higley and Raddin 1891; Pepoon
1927; Mohlenbrock 1975. A. /ucidulus—Deam
1940; G.N. Jones 1945, 1950, 1963: Gleason
1952; Jones and Fuller 1955; Gleason and
Cronquist 1963; and others. A. puniceus subsp.
firmus (Nees) A.G. Jones (1984)]:

1. Stems nearly glabrous or glabrescent in the
lower half, hirsute or sparsely hispidulous in
lines in the upper portion; lower leaf surface
glabrous or sparsely scabrous along the
midrib; phyllary tips short-attenuate or often
acute to obtuse; rhizomes with strongly
creeping stoloniform strands ........ var. firmus

1. Stems hispidulous over the entire length, the
indument at least in part uniformly distrib-
uted around the stem; lower leaf surface
minutely strigillose, the midrib hispidulous
or densely hirsute to villous; phyllary tips
generally long-attenuate; rhizomes with
short, thick, fleshy strands........ var. puniceus

May 1989

The ranges of these two varieties in Illinois
are overlapping, but plants of var. firmus seem to
be more common than those of var. puniceus.
Attempts to separate the two taxa at the level of
species have been frustrated because of a high
degree of intergradation and inconstancy in the
character states. There is also evidence of
intergradation between Aster puniceus and such
other members of the x = 8 chromosome number
assemblage as A. /anceolatus [e.g., Bebb s.n. (F
17355) from Winnebago Co. and Wade & Wade
1744 (ISM) from Ogle Co.], A. lateriflorus (see
Steyermark 1963), and A. praealtus [e.g., Smith
667 and 683 (F) from Cook or Du Page Co. and
Shildneck 11617 (ILL) from Fayette Co.]. Illinois
specimens that have been labelled A. longifolius
belong mostly in A. puniceus var. firmus.
Although G.N. Jones (1950, 1963) in his keys
treated A. /ongifolius in accordance with
Lamarck’s type and circumscription, the species
probably does not get into Illinois. The collec-
tions cited by Pepoon (1927) for A. patens
{Raddin (F) and Umbach (F)] belong in A.
puniceus var. firmus, not in A. laevis as suggested
by Swink and Wilhelm (1979).

24. Aster schreberi Nees
Schreber’s Aster

Herbaceous, with a
strongly creeping
horizontal rhizome
system. New shoots
arising mostly at the ends of
stoloniform rhizome
branches, the plants forming
extensive colonies; basal
rosettes typically numerous, even
in midseason far outnumbering the
flowering stems and often forming a dense
ground cover. Flowering stems single at any
point of emergence, erect, 30-80 cm tall, terete
or, at the upper nodes, somewhat angular in cross
section from decurrent nerves, the ascending or
divaricate branches mostly confined to the

capitulescence, the main stem sometimes reddish,

glabrous, the branches sparsely scabrous or
puberulent, sometimes in decurrent lines, but the
indument usually uniformly distributed on the

ASTER AND BRACHYACTIS IN ILLINOIS 177

peduncles. Leaves polymorphic, those of the
basal rosettes and the lower half of the stem
petiolate; basal rosette leaves largest, the blades
broadly ovate to cordate, (8)10—25 cm long and
(6)8—16 cm wide, with 5-8 pairs of pinnate
secondary nerves, and a weakly expressed
reticulum of tertiary veins with mostly isodia-
metric (rarely oblong) areolae, the upper surface
glabrous or thinly scabrous, the lower with a few
trichomes along the nerves, the apex acuminate
with a sharp mucro, the margins coarsely and
often irregularly crenate-serrate, each tooth with
a clear or green to purplish mucro, the base
deeply cordate, the lobes sometimes overlapping
or the sinuses often broad, the petioles |—2 times
as long as the blades, with slightly winged and
glabrous or sparsely ciliate margins, somewhat
dilated and sheathing at the base; cauline leaves
gradually reduced in size upward on the stem,
blades of the larger ones often nearly as wide as
long, similar to the basal leaves in contour and
indument except for the progressively shorter
and gradually more broadly winged petioles:
upper cauline leaves subpetiolate or sessile,
elliptic or obovate, with a truncate or cuneate
base; rameal leaves relatively few, abruptly
reduced, variable in size, ovate or oval,
subglabrous or the indument similar to that of the
larger leaves, the apex acute or acuminate, the
margins finely and sharply serrate and ciliate, the
base rounded or cuneate; leaves of the peduncles
0—few, ovate-lanceolate, 2-3 mm long, flexible,
attenuate, entire, ciliate, not intergrading with the
phyllaries. Capitulescence a terminal corymbi-
form panicle, but often also with lateral
flowering branches from the upper nodes of the
main stem, the cymule-bearing branchlets stiffly
divaricate or dichotomous, the peduncles (if
developed) usually ascending at a wide angle,
(0).3—2 cm long, slender, typically not glandular
(rarely with a few stipitate glands). Flowering
heads variable, 2—2.5 cm in diameter when the
rays are fully extended, sometimes crowded and
sessile in the individual cymules. Involucre
campanulate (turbinate when pressed), 5—7 mm
high, the phyllaries imbricated in 4 or 5 series.
Phyllaries strongly graduated, appressed when
fresh (loosely spreading in fruiting heads and
when pressed and dried), those of the outer 2
series oblong-ovate, |—2(2.5) mm long, glabrous
or slightly puberulent on the back, not glandular
(rarely with a few minute glands at the tips), the
green areoles oblong to oblanceolate (rarely

178 ILLinois NATURAL History SURVEY BULLETIN

extending to the base as a broad band), the
margins usually densely lanate and with a
scarious rim, the basal (‘'/:)'/>—’/: portion of the
phyllaries scarious, indurate, and keeled or
rounded on the back; phyllaries of the inner 2
series linear-lanceolate, 4.5—6(7) mm long,
glabrous, largely scarious or only the midrib
green, the apex often reddish. Receptacle
alveolate with sharp teeth. Ray florets 6—8(12),
the corollas (8)10—12(15) mm long, white or
cream-colored, glabrous. Disk florets 15—25 (or
more), the corollas funnelform, (5)5.5—7 mm
long, glabrous, or thinly puberulent on the
slender, tubular portion of the limb, the limb
strongly flared ca 1.5 mm above the point of
insertion of the filaments, cream-colored or light
yellow aging to purple, the lobes reflexed, the
fraction of lobe/expanded limb portion 0.4—0.5,
the apparent tube (i.e., including the tubular limb
portion) distinctly longer than the expanded limb
portion. Pappus tawny, the bristles in 2 series,
those of the inner series about as long as the disk
corolla, firm, with a clavellately expanded apex,
those of the outer somewhat shorter, more
slender, and attenuate. Achenes slenderly
fusiform, 3—S mm long and 1—1.2 mm across,
dull purple or light chocolate brown, glabrous or
with a few scattered trichomes near the top, with
(7)8—10 thick (sometimes double-stranded)
straw-colored ribs. 27 = 54. Including A. chasei
G.N. Jones in Jones & Fuller (1955).

Late July-September. Mesic but usually
well-drained loamy or gravelly soil in wooded
areas: plants in Illinois found mostly on north-
facing slopes of ravines or along streams and
rivers ca 30-60 m (or more) above the water
level. Occasional in the northern one-third of the
state, south to Tazewell County and west to Rock
Island, Henry, and Knox counties. Illinois (and
Wisconsin) records represent the western outpost
of this species, disjunct by ca 450 km from the
nearest populations to the east. The species range
extends from ME west through s.e. ON to s.e. WI
and south through PA, WV, and s.e. OH to VA,
e. KY, e. TN, and n. AL.

Aster schreberi has been included in the
Illinois list of threatened species, but the plants
are of rather common occurrence in the piedmont
and mountain woods of the eastern and southeast-
er states. I have so far been unsuccessful in a
search for consistent characters whereby the
midwestern populations [sometimes treated as
Aster chasei—G.N. Jones 1963] can be distin-

Vol. 34 An. 2

guished from those to the east, at least at the
varietal level. The plants also share the same
hexaploid chromosome number of 2n (= 6x) = 54.

25. Aster sericeus Vent.
Silky Aster

Herbaceous, with a
cormoid-caudiciform
rhizome system that

turns woody with age.
New shoots initiated at or
near the base of old stems =
resulting in scattered individual lane
clumps. Stems |—several, S54
slender, erect, wiry, somewhat
fastigiately branched from near the
middle, 20—70 cm tall, aging to light brown,
glabrous in the lower half, thinly soft-pubescent
in the middle portion, and sericeous-strigose in
the branchlets. Leaves relatively uniform, sessile,
entire, appressed sericeous to densely silvery-
silky on both surfaces including the margins, the
basal and lower cauline ones often withered and
deciduous at flowering time; basal rosette leaves
oblanceolate or spatulate, with | or 2 pairs of
secondary nerves arching forward from the base
in alignment with the margins, the surfaces less
copiously pubescent than those of the cauline
leaves, the apex acute, the base sheathing:
principal cauline leaves slightly or gradually
reduced upward on the stem, oblong to linear-
lanceolate, 1.5—3(5) cm long and 4-10 mm wide,
the apex mucronulate, the base rounded but not
clasping; rameal leaves similar in contour and
vestiture, flexible, acute, spinulose-mucronulate,
those of the peduncles crowded, 4—-8(10) mm
long, intergrading with the phyllaries.
Capitulescence an open, somewhat fastigiately
branched panicle in the upper '/:—/: portion of
the stem, the branchlets often arching. Flowering
heads 2-3 cm in diameter when the rays are fully
extended, usually not crowded and not secund,
subsessile or on peduncles 0.5—3(5) cm in length.
Involucre cylindric to narrowly campanulate,
5-8(10) mm high, the phyllaries imbricated in
3—5(6) series. Phyllaries graduated or sometimes
subequal, spreading or squarrose to reflexed,
sericeous on both surfaces, including the scarious
basal portion and the margins; outer phyllaries

May 1989

(4)5—6 mm long, often largely herbaceous, ovate,
acute, mucronulate, the median ones 6—8(10) mm
long, ovate-lanceolate, acuminate or attenuate,
green in the expanded apical '/:—/: portion, the
abruptly narrowed basal portion scarious,
indurate, and rounded on the back; innermost
phyllaries very slender, attenuate, often reddish at
the base. Receptacle strongly alveolate with
sharp teeth. Ray florets (10)15—25, the corollas
12—15(18) mm long, deep purple (rarely white),
with a few trichomes near the throat. Disk florets
20-30 (or more), the corollas narrowly funnel-
form, (5)6—7 mm long, thinly puberulent on the
tube and throat, the limb bright yellow turning
reddish purple after anthesis, the lobe/limb
fraction 0.18—0).2, the tube much shorter than the
limb. Pappus simple, the bristles about as long as
the disk corolla, discolored or tawny, relatively
firm, attenuate. Achenes fusiform, plumpish,

2-3 mm long and 0.7—1 mm across, purple at
maturity or brown when weathered, glabrous,
prominently 7—10 ribbed. 2n = 10.

Late August—October. Dry sandy, loamy, or
rocky soils in unshaded situations: sand barrens,
dunes, hill prairies, and open-wooded bluffs.
Local, mostly in the northern half of the state, but
extending southward along the Mississippi River
to Randolph County. The range of the species
extends from s. ON and MI to TN, and in the
West through the eastern half of the Great Plains
from s.e. MB to TX.

Illinois plants belong in var. sericeus.

26. Aster shortii Lindley in Hooker
Short’s Aster

Herbaceous, with
branched-caudiciform or
short horizontal rhizomes
that turn woody with age.
New shoots initiated at or
near the base of old stems
resulting in scattered individual
clumps. Stems |—several, erect,
(40)80—120 cm tall, much branched
and bushy, densely and uniformly
soft-hirtellous on the upper stem and in the
branches, the indument in decurrent lines farther
down, or the stem glabrescent in the lower

ASTER AND BRACHYACTIS IN ILLINOIS 179

portion. Leaves polymorphic, the basal and
principal cauline ones petiolate; leaves of new
shoots and of the vernal rosettes with the blades
mostly ovate, 1-6 cm long and 1—3.5 cm wide,
dark green above, purplish below at least during
the cool season, with | or 2 pairs of anastomos-
ing secondary nerves curving forward from the
base and additional pinnate nerves emanating
from the midrib, the apex acute or obtuse, the
margins crenate, the base cordate or rarely
truncate; petioles twice as long as the blades,
slender, not at all or only narrowly winged, often
densely pilose or hirsute; principal cauline leaves
largely persistent throughout the flowering
period, regularly spaced, the internodes 2—3(4)
cm long, the blades ovate to lanceolate, some-
times falcate, S—10(15) cm long and 2-7 cm
wide, often conspicuously rugulose, with several
pairs of pinnate and anastomosing secondary
nerves, and a reticulum of tertiary veins with
more or less isodiametric areolae, the upper
surface glabrous, or slightly scabrous along the
midrib, the lower surface copiously hirtellous or
hirsute, the apex acute to attenuate or acuminate
and with a sharp callus point, the margins mostly
entire and scabrous, or shallowly crenate-serrate
in the lower leaves, the base cordate, truncate, or
rounded, sometimes oblique, the petioles half as
long as the blades or less, slender, not sheathing
or clasping; rameal leaves more or less abruptly
reduced in size, lanceolate to ovate, entire,
sessile, or sometimes with short petioles, densely
hirtellous, acute or obtuse with a sharp, often
purple callus point; leaves of the peduncles
minute, bracteiform, |—1.5 mm long, inter-
grading with the phyllaries. Capitulescence

an ample, diffuse, broad-topped panicle,

the head-bearing branchlets often arching.
Flowering heads |.5—2.5 cm in diameter when
the rays are fully extended, usually not crowded
and not secund, the peduncles densely bracteate,
0.2—3(5) cm long, hirtellous. Involucre
campanulate (or hemispherical when pressed
and dried), 4.5—-6 mm high, the phyllaries
imbricated in 5 or 6 series. Phyllaries appressed
when fresh, strongly graduated, the outer
triangular or lanceolate, 1—-1.5 mm long, the
inner linear-lanceolate, 4—5.5 mm long,
puberulent on both surfaces, the green areoles
oblanceolate to rhombic in the apical '/.—'/:(‘/:)
portion, usually much shorter than the scarious
basal portion (rarely the outer phyllaries with a
green band to the base), the apex acute or

180 I-tinois NATURAL History SurveEY BULLETIN

acuminate, the margins erose-hyaline and
irregularly ciliolate. Receptacle alveolate with
sharp teeth. Ray florets (15)18—25, the corollas
10-15 mm long, typically deep blue or purple,
glabrous. Disk florets 20—25 (or more), the
corollas funnelform, 5—6 mm long, glabrous or
nearly so, the limb abruptly dilated at the throat,
light yellow turning reddish purple after anthesis,
the lobe/limb fraction 0.18—0.2, the tube shorter
than the limb. Pappus simple, the bristles about
as long as the disk corolla, tawny or rose-tinged,
soft, slender, and attenuate. Achenes oblong-
obovoid, somewhat compressed, 2.5—3 mm long
and ca | mm across, dull purple at maturity or
brown when weathered, glabrous, with 4 or 5
ribs. 2n = 16; reported chromosome counts of 27
= 18 are probably in error (A.G. Jones 1977).

September—October. Well-drained soils:
edges of upland woods, thickets, wooded river

banks, and open-wooded slopes. Common nearly
throughout the state.The species range extends
from s. ON, w. PA, and n.w. MD, south to GA,
n. FL, and AL, west to e. MN and e. IA with
scattered records from AR (but apparently none
from MO).

Plants in Illinois belong in var. shortii,
including f. gronemanii Benke, described as a
plant with rose-red rays from a collection made
near Elgin, Kane County [Benke 4872 (F)]. There
is evidence of occasional hybridization and
intergradation with Aster anomalus and A.
urophyllus [e.g., Fell 53-1005 (ILL) from De
Kalb Co. and Winterringer 14794 (ISM) from
Jersey Co.], and with other related species.

27. Aster tataricus L-f.
Tartarian Aster

Herbaceous, with a branched cormoid-
caudiciform rhizome system but also producing
short, fleshy, stoloniform rhizome strands. New
shoots arising from nodes and tips of rhizomes
and also from many winter buds just below the
base of old stems, the plants aggressively
spreading by vegetative reproduction. Stems
l-several, stout, erect, 80-150 cm tall, somewhat
angled and ridged from decurrent leaf bases,
branched only in the capitulescence, uniformly
hirtellous or scabrous with antrorsely hooked
trichomes. Leaves polymorphic, the basal ones
most prominent forming convoluted clusters

Vol. 34 Art. 2

rather than rosettes, very coarse, 3—4 times as
large as the largest cauline leaves, the blades
oblanceolate, 10-30 cm long and 5—10 cm wide,
strongly rugose, with 6-12 pairs of pinnate and
anastomosing secondary nerves, the upper
surface minutely scabrous, the lower surface
densely scabrous or puberulent especially along
the nerves, the apex acute, obtuse, or rounded,
and mucronate, the margins crisp, crenate-serrate,
each tooth with a conspicuous mucro, the base of
blades gradually tapered, the petioles winged,
10-40 cm long, as long as the blades or longer,
sheathing at the base; cauline leaves gradually
reduced upward on the stem, (8)10—15(18) cm
long and 2-5 cm wide, relatively densely spaced
with internodes 24 cm long, the lower leaves
oblanceolate and subpetiolate, the upper
lanceolate and sessile, similar to the basal leaves
in surface, venation, and indument, the apex
acute or acuminate and mucronulate, the margins
serrate or entire, densely ciliolate, the base
cuneate, sheathing, and decurrent; rameal leaves
abruptly much reduced in size, lanceolate, 0.5—1
cm long, puberulent, acute or attenuate and
mucronulate, those of the peduncles few,
bracteiform but flexible, not phyllarylike.
Capitulescence an elongate or broad-topped
panicle in the upper '/,—/: of the stem, the
terminal branchlets short with few-headed
corymbiform clusters. Flowering heads 2—2.5(3)
cm in diameter when the rays are fully extended,
sessile in the axils of subtending leaves or on
puberulent and somewhat ridged peduncles
0.5—1.5(2.5) cm long. Involucre campanulate or
hemispherical, 6.5—8(10) mm high, the phyllaries
imbricated in 5 or 6 series. Phyllaries appressed
or somewhat spreading, graduated, those of the
outer series triangular or lanceolate, 34 mm
long, those of the inner series linear-lanceolate,
6-8 mm long, glabrous or minutely puberulent,
with green bands along the midrib to the base, or
the outer phyllaries sometimes largely herba-
ceous, the apex acute in those of the outer 2
series, attenuate in those farther inward, the
margins entire or erose-hyaline and with a red
rim. Receptacle alveolate with sharp teeth. Ray
florets 15—18(20), the corollas 10—15(18) mm
long, lavender, glabrous. Disk florets 20-30

(or more), the corollas funnelform, (4.5)5—6 mm
long, glabrous or with a few trichomes at the
throat, the limb abruptly dilated, light yellow
turning lavender after anthesis at least in the

May 1989

lobes, the lobe/limb fraction 0.2, the lobes notably
recurved, the slender tube slightly shorter than the
limb. Style branches somewhat atypical for the
genus in having acutish rather than attenuate sterile
appendages. Pappus simple, the bristles shorter
than the disk corolla, white or cream-colored, soft,
slender, and attenuate. Achenes obconic-obovoid,
slightly compressed or plump, 1.5—2 mm long and
ca | mm across, light brown, thinly strigillose,
with 4 or 5(6) ribs. 2n = 54.

October. Disturbed ground: roadsides,
thickets, and waste places. Occasionally escaped
from cultivation. Introduced from n.e. Asia.
Recorded by me from Champaign, Montgomery,
Sangamon, St. Clair, and Vermilion counties.

28. Aster turbinellus Lindley in Hooker

Herbaceous, with stout,
branched, caudiciform

fel

(Rat
rhizomes that turn Se iarl "v2
woody with age. New ‘the

shoots initiated at or near the
base of old stems resulting in
scattered individual clumps.
Stems |-several, 50-100 cm tall,
with many ascending or divaricate
branches from near or below the
middle, glabrous or hirtellous in thin decurrent
lines. Leaves polymorphic, notably variable in
size, the lower ones soon withered and decidu-
ous; basal rosette and lower cauline leaves
subpetiolate, the blades oblong-oblanceolate,
glabrous, the apex acute, obtuse, or rounded, the
margins shallowly crenate and ciliolate, the
subpetiolar portion coarsely ciliate, winged, and
slightly dilated toward a sheathing base;
principal leaves gradually reduced upward on the
stem, sessile, firm, linear-oblanceolate to elliptic-
lanceolate, gradually tapered at both ends, 4-12
cm long and 0.5—2 cm wide, with a prominent
midrib and weakly expressed anastomosing
secondary nerves, the upper surface glabrous, the
lower surface scabrous only along the midrib, the
apex acute with a sharp callus point, the margins
mostly entire, ciliolate, the base cuneate or
rounded and hugging the axillary branchlets;

ASTER AND BRACHYACTIS IN ILLINOIS 181

rameal leaves similar in contour, those of the
peduncles often densely spaced, bracteiform,
appressed or ascending, oblong to subulate, 1.5—4
mm long, continuous with the phyllaries but
distinct in having a pointed apex with a brown or
purple mucro. Capitulescence an open, broad,
round-topped panicle. Flowering heads 2—3 cm
in diameter when the rays are fully extended, not
crowded and not secund, usually at the ends of
wiry, ascending or divaricate, few- to many-
bracted peduncles (1)4—10(25) cm in length
(rarely sessile). Involucre slenderly campanulate
or turbinate, 7—12 mm high, the phyllaries
imbricated in 6—9 series, inserted on a prominent
obconical rachis 24 mm in height. Phyllaries
strongly graduated, appressed or somewhat
spreading, indurate, rounded on the back or
slightly keeled, the green areoles well delimited,
oblong to rhombic-oblanceolate, centered in the
apical '/s—'/; portion, usually much shorter than
the scarious basal portion, the apex obtuse,
rounded, or almost truncate, sparsely lanate,
including the top (areolar) portion of the adaxial
surface, the margins with a comparatively broad
scarious rim extending to the tip; outer (lower-
most) phyllaries ovate, 1.5 mm long, the median
and inner ones linear, to 7 mm long. Receptacle
alveolate with sharp teeth. Ray florets 15—20, the
corollas 10-15 mm long, blue or purple,
glabrous. Disk florets 15—20 (or more), the
corollas narrowly funnelform, 4.5—6(7) mm long,
glabrous, the limb yellow turning purple after
anthesis at least in the lobes, the lobe/limb
fraction 0.18, the tube sightly shorter than the
limb. Pappus simple, the bristles about as long as
the disk corolla, tawny or somewhat rose-tinged,
comparatively firm, attenuate. Achenes oblong in
contour, somewhat compressed, (1.8)2—2.8 mm
long and ca | mm across, light brown or gray,
puberulent or minutely strigillose, often puncticu-
late, with 3—5 ribs. 2n = 96, sometimes with
additional B-chromosomes.

Late August—October. Dry loamy or rocky
soils: edges of upland woods, open-wooded
slopes and bluffs, and also in somewhat disturbed
ground of pastures and roadsides. Local, mostly
in the southern half of the state, north to Macon,
Fulton, and McDonough counties. Records from
Illinois represent the northern and northeastern
limits for the species. The range extends
southward to n. LA and westward through s. MO
and AR toe. NE, s.e. KS, and e. OK.

182 ILtinots NATURAL History Survey BULLETIN

29. Aster umbellatus Miller
Flat-top Aster

Herbaceous, with a
creeping horizontal
rhizome system. New
shoots initiated at
intervals along or at the ends
of stoloniform rhizome strands
resulting in colonial stands;
autumnal rosettes lacking. Stems
mostly single at any point of emer-
gence, erect, 50-150 cm tall, usually
unbranched below the capitulescence, glabrous in
the lower portion, commonly puberulent with
antrorsely hooked trichomes above, the branches
somewhat winged or ridged from decurrent leaf
bases. Leaves (except the lowermost) more or
less uniform in contour, sessile or subpetiolate,
those of the midstem ovate to elliptic-lanceolate,
4-12 cm long and 1—3(4) cm wide, with a
prominent midrib that is abaxially keeled in the
basal or subpetiolar portion and with 6-8 pairs of
pinnate and anastomosing secondary nerves, as
well as a conspicuous reticulum of tertiary veins
with isodiametric areolae, the upper surface dark
green, glabrous or minutely scabrous, the lower
surface bluish green, scabrous-puberulent, at least
along the midrib and major nerves, the apex
acuminate, the margins entire, scabrous, the base
cuneate and decurrent; lower cauline leaves
smaller, often spatulate, those near ground level
bracteiform; rameal leaves relatively few, much
reduced in size but otherwise resembling the
larger leaves except for the few small, subulate,
puberulent bracts subtending the head. Capitu-
lescence flat- or round-topped, each individual
cluster corymbiform. Flowering heads 1—1.5(2)
cm in diameter when the rays are fully extended,
often crowded, subsessile, or on slender puber-
ulent peduncles 0.2—2.5 cm in length. Involucre
campanulate or turbinate, 3.5—4.5(5) mm high,
the phyllaries imbricated in 4 or 5 series.
Phyllaries strongly graduated, the outer ones 1-2
mm long, the inner 3—4(6) times as long, more or
less appressed at flowering time but widely
spreading in fruiting heads, puberulent or
glabrous on the back; outer (lowermost)
phyllaries subulate, the median and inner ones
oblong to linear-lanceolate, the green areoles

Vol. 34 An. 2

forming broad bands to the base as an extension
of the midrib, the apex acute to obtuse or
sometimes rounded, the margins erose-hyaline
and irregularly ciliolate. Receptacle alveolate
with long sharp teeth. Ray florets (2)6—15 (or
more), the corollas 8-10 mm long, often with a
few trichomes on the slender tube, the rays
whitish, comparatively broad, 2—3 mm wide.
Disk florets (8)12—20, the corollas funnelform,
4-7 mm long, thinly puberulent on the tube, the
limb partway slenderly tubular, abruptly flared
ca 1 mm above the point of insertion of the
filaments, cream-colored turning purple after
anthesis, the lobes relatively long and reflexed,
the fraction of lobe/expanded limb portion
0.6-0.7, the apparent tube (i.e., including the
tubular limb portion) distinctly longer than the
expanded limb portion. Pappus “double”
(according to literature references) but actually
composed of 3 series of bristles, whitish or
somewhat discolored; bristles of the inner series
about as long as the disk corolla, firm, clavel-
lately expanded toward an acute apex, those of
the middle series slightly shorter, tapered toward
an attenuate apex, and those of the outer series
minute, 1 mm long or less, slender, and attenuate.
Achenes obovate to oblanceolate in contour,
compressed, 2.5—3.5 mm long and I—1.2 mm
across, light brown, sparsely puberulent (Illinois
plants) or sometimes glabrous, with 5 or 6
prominent, glossy, golden-brown ribs. 27 = 18.
Including A. pubentior Cronquist (1947).
[Doellingeria umbellata (Miller) Nees; A.
infirmus misapplied, not of Michaux—Brendel
1887.]

(Late July) August-September (early
October). Low, damp, mostly open ground:
swamps, seep areas, thickets near streams and
rivers, and clearings in woods. Occasional in the
northern half of the state, south to Menard and
Cass counties. The range of the species extends
from NF south to GA, n. FL, LA, s. AR, and e.
TX, west to e. AB, SK, and ND, with a few
stations in IA, NE, and s. OK.

Based solely on geographic considerations,
plants of Illinois would be placed in var. pubens
A. Gray (cf. Fernald 1950) or, at the rank of
species, in Aster pubentior (cf. Gleason 1952;
Gleason and Cronquist 1963). Too much
variability is evident in the specimens examined,
however, to permit a clear separation of this
variant from var. wnbellatus. Gray (1884)
distinguished plants of var. pubens merely by

May 1989

more copious pubescence on the lower leaf
surface. Cronquist (1947), in his argument for
species rank, considered additional characteris-
tics, e.g., puberulent phyllaries and smaller heads
(12-22 florets) compared with glabrous phyllar-
ies and larger heads (23-54 florets) in A.
umbellatus var. umbellatus. In applying these
criteria, however, one may find that both taxa
occur side by side in northern Illinois. I do not,
therefore, recognize any varieties of this species
for the flora of Illinois.

30. Aster undulatus L.
Wavy-leaved Aster

Herbaceous, with
branched caudiciform
rhizomes that turn woody
with age but also some-
times with short stoloniform
rhizome strands. New shoots
arising at the base of old stems
or from the rhizomes, the plants
usually forming scattered individual
clumps. Stems |-several, erect,
40-120 cm tall, with ascending or divaricate
branches above the middle, densely and uni-
formly hirtellous or villous on the upper stem
portion and in the branches, somewhat pubescent
in lines or glabrescent in the lower stem portion.
Leaves polymorphic, the basal and lower cauline
ones petiolate, those higher up on the stem
subsessile or sessile and strongly clasping; upper
surface of the blade mostly scabrous to hirsute
(rarely glabrous), the lower surface hirtellous or
loosely villous; leaves of new shoots and of the
vernal rosettes with ovate-oblong to suborbicu-
late blades 1-6 cm long and 1-4 cm wide,
purplish below, with two or more pairs of
somewhat arching and anastomosing secondary
nerves, the apex obtuse or rounded, the margins
shallowly crenate-serrate to subentire, the base
cordate or truncate to rounded, sometimes
oblique, the petioles usually purplish, hirsute or

pilose, winged, dilated, and sheathing at the base;

principal cauline leaves variable in shape and
size, gradually reduced upward on the stem, the
lower ones petiolate, those higher up sessile and
often constricted near or below the middle, the

ASTER AND BRACHYACTIS IN ILLINOIS 183

blades ovate to lanceolate in outline, 3—12(14) cm
long and (1)2—S cm wide, the secondary venation
mostly obscured by indument, the apex acute to
attenuate or acuminate and callus-pointed, the
margins crenate-serrate or entire, and scabrous,
the petioles progressively shorter and more
broadly winged upward on the stem, conspicu-
ously dilated toward the auriculate or cordate,
clasping base; rameal leaves more or less
abruptly reduced in size, oblong or lanceolate to
linear-lanceolate, acute to attenuate, mucronulate,
subentire to entire, sessile, slightly clasping or
sheathing; leaves of the peduncles often numer-
ous, firm, bracteiform, 2—3 mm long, appressed
or ascending, intergrading with the phyllaries.
Capitulescence paniculiform, often notably
overtopping the leafy stem portion but also
sometimes with branches arising from nodes near
the middle of the stem, the ultimate head-bearing
branchlets ascending or divaricate, often
racemiform. Flowering heads 1.2—1.5(2) cm in
diameter when the rays are fully extended,
usually not crowded but sometimes secund, the
peduncles 0.3—3(5) cm long, often densely
bracteate, hirtellous. Involucre campanulate or
hemispherical, 4-6(7) mm high, the phyllaries
imbricated in 4 or 5(6) series. Phyllaries strongly
graduated, appressed, or those of the outer series
spreading, lanceolate, 1.5—2.5 mm long, the inner
ones linear-oblanceolate, 4—5(6) mm long,
puberulent on the abaxial surface and also thinly
so on the adaxial surface, the green areoles
rhombic or oblanceolate, the apex sharply acute,
attenuate, or acuminate with somewhat inrolled
margins, often mucronulate, the margins erose-
hyaline, irregularly ciliolate, and with a scarious
rim to near the tip, the basal '/>—/: portion
scarious, or sometimes the midrib green to the
base. Receptacle alveolate with mostly rounded
teeth. Ray florets (12)15—25, the corollas 8—12
mm long, purple or blue, glabrous or nearly so.
Disk florets 15—25, the corollas narrowly
funnelform, (4)5—6 mm long, glabrous or with

a few trichomes near the throat, the limb abruptly
dilated at the throat, cream-colored or light yel-
low turning purple after anthesis, the lobe/limb
fraction 0.2—0.25, the tube shorter than the limb.
Pappus simple, the bristles about as long as the
disk corolla or slightly shorter, cream-colored or
slightly rose-tinged, soft, slender, and attenuate.
Achenes oblong-obovoid, somewhat compressed,
2.5—3(3.5) mm long and 1.2—1.5 mm across, dull
purple or light brown, sparsely puberulent at least

184 ILLinois NATURAL History SurvEY BULLETIN

toward the top, with 4 or 5 straw-colored ribs.
2n = 32; reported chromosome counts of 2n = 18
and 36 are probably in error (A.G. Jones 1977,
1980b).

August—October. Dry or well-drained soils:
loamy or rocky slopes at forest edges and open-
wooded bluffs underlain by sandstone or
limestone. Uncommon, restricted to the southern
tip of the state; records from Jackson and
Alexander counties mark the western limit for
this extremely variable species. The range
extends from NS and ME toc. FL, west to s.
ON(?), OH, through s. IN, s. IL, and TN to c. MS
and s.e. LA.

Aster undulatus has been included in the
Illinois list of threatened species but is one of the
more common asters in the mountains and
woodlands to the east and southeast of our state.
Although several varieties have been described, I
am unable to assign Illinois populations to any of
them. Even though the plants seem to differ
somewhat from the type specimen of A. undula-
tus, | am treating them at this time sensu lato or
as belonging in var. undulatus. There is evidence
of occasional hybridization and intergradation
with such closely related species as A. drum-
mondii [e.g., A.G. Jones 4089 (ILL) from
Gallatin Co.] and A. oolentangiensis [e.g., K.
Wilson 3164 (ISM) from Pope Co.]. Collections
that form the basis for A. undulatus sensu Mead
(1846); Higley and Raddin (1891); Pepoon
(1927); Kibbe (1952); and Dobbs (1963) non L.
do not belong in this species.

31. Aster urophyllus Lindley in DC.

Herbaceous, with a stout
caudiciform rhizome
system lacking stoloni-
form strands. New shoots
arising at or near the base of
old stems, the plants forming
scattered individual clumps.
Stems 1|-several, erect, brittle,
40-120 cm tall, with stiffly ascend-
ing, somewhat ridged branches from
above the middle, sparsely puberulent in the
upper portion and in the branches, the indument

Bas
Sjcuaean
bd |

Vol. 34 Art. 2

usually in decurrent lines (rarely uniformly
distributed around the stem), the lower stem
portion glabrescent or glabrous. Leaves polymor-
phic, the basal and larger cauline ones petiolate,
those of the upper stem subsessile to sessile:
leaves of new shoots and of the vernal rosettes
with ovate or lanceolate blades 4-12 cm long and
2-5 cm wide, nearly glabrous, or often minutely
scabrous above and thinly pubescent below, the
secondary venation weakly expressed, the apex
acute to acuminate, the margins shallowly
crenate-serrate, the base cordate or truncate to
rounded, the petioles S—15 cm long, slender or
narrowly winged, dilated, and sheathing at the
base; principal cauline leaves gradually reduced
in size upward on the stem, the blades ovate to
lanceolate, 5—12 cm long and 2-5 cm wide,
similar to the basal leaves in venation and
vestiture, the apex acuminate to attenuate with a
sharp, often purple callus point, the margins
crenate-serrate to subentire, the base truncate or
rounded, the petioles progressively shorter and
more broadly winged upward on the stem,
slightly sheathing; rameal leaves abruptly
reduced in size, lanceolate to linear-lanceolate,
glabrous or nearly so, attenuate and callus-
pointed, entire, scabrous-margined, cuneate and
slightly decurrent; leaves of the peduncles
bracteiform, linear or subulate, 2—4+ mm long,
mostly ascending (rarely spreading to recurved),
intergrading with the phyllaries. Capitulescence
typically a dense, narrow, ovoid or pyramidal
panicle in the upper '/: —/:(‘/:) portion of the
stem, commonly overtopping the leafy portion,
the head-bearing branchlets stiffly ascending,
often racemiform. Flowering heads |—1.2(1.5)
cm in diameter when the rays are fully extended,
usually crowded and sometimes secund, sub-
sessile or on often densely bracteate, puberulent
peduncles 0.3—2 cm long. Involucre cylindrical
or turbinate, 4.5—6(7) mm high, the phyllaries
imbricated in 4 or 5 series. Phyllaries appressed
or somewhat recurved-spreading, graduated, the
outer subulate, 2—3 mm long, the inner linear-
lanceolate, 5—6(7) mm long, glabrous, scarious
over most of the surface area, the light green
areoles very slender, linear to linear-oblanceolate,
the apex long-attenuate, terminating in an often
recurved, spinulose mucro, the margins erose-
hyaline and irregularly ciliolate. Receptacle
alveolate with sharp teeth. Ray florets
(8)10—12(14), the corollas 6—-8(10) mm long.
typically white (rarely lavender), sparsely

May 1989

puberulent on the tube and throat. Disk florets
10-15, the corollas funnelform, abruptly dilated,
(3.5)4-5 mm long, glabrous or with a few
trichomes near the throat, the limb cream-colored
turning pinkish after anthesis, the lobe/limb
fraction 0.2—0.25, the tube much shorter than the
limb. Pappus simple, the bristles shorter than the
disk corolla, whitish or slightly discolored to
rose-tinged, soft, slender, and attenuate. Achenes
oblong-obovoid, somewhat compressed, 1.8—2.5
mm long and ca | mm across, dull purple or
brown, glabrous, with 4 or 5 ribs. 2” = 16. [A.
Sagittifolius sensu auct. (pro parte) non Wedem.
ex Willd. (see A.G. Jones 1980b; Jones and
Hiepko 1981). A. hirtellus Lindley in DC.
A.sagittifolius var. hirtellus (Lindley in DC.)

ASTER AND BRACHYACTIS IN ILLINOIS 185

Burgess in Britton & Brown f. hirtellus (Lindley
in DC.) Shinners (1941 )—Steyermark 1963.

A. sagittifolius var. urophyllus (Lindley in DC.)
Burgess in Britton & Brown—Deam 1940.

A. cordifolius var. moratus sensu Fern. (1950)
non (Shinners) Shinners. ]

August—October. Loamy or rocky soils in
dry or mesic situations: woodland edges, open
upland woods, and thickets, also in disturbed
ground of pastures, roadsides, and railroad rights-
of-way. Common throughout much of the state.
The range extends in the East from s.e. Canada
and ME south to GA, AL, and n. FL, and in the
West from MN through IA, MO, and s.e. NE to
s.e. KS, with scattered stations recorded from AR
and MS.

186 ILLinois NATURAL History SuRVEY BULLETIN

Vol. 34 Art. 2

Description of Brachyactis Species

Brachyactis ciliata (Ledeb.) Ledeb.
Rayless Aster

Taprooted and somewhat
succulent annual. Stem
erect, 10-70 cm tall,
bluish or yellowish
green, often red-tinged,
glabrous or with a few bristly
trichomes in the leaf axils and
along the decurrent leaf bases;
branches mostly ascending, often
from near the base. Leaves bluish
green, with a conspicuous midrib, the
secondary venation faintly expressed; basal
leaves spatulate, soon withered; principal cauline
leaves linear, 3-10 cm long and 0.1—-0.9 cm wide,
gradually reduced in size upward on the stem,
sessile, with clusters of smaller leaves produced
in the axils, glabrous, the apex acute to short-
attenuate, the margins entire and appressed
ciliate, the base slightly dilated and sheathing;
rameal leaves similar in contour, those of the
peduncles few. Capitulescence a contracted
panicle, or racemiform in poorly developed
plants. Heads several to many, short-peduncled
or subsessile. Involucre 6—10 mm high, the
phyllaries imbricated in 3 or 4 series. Phyllaries
subequal or somewhat graduated, occasionally
those of the outer series longer than those of the
middle and inner series, glabrous, linear to
oblanceolate, (4)5—8 mm long, the outer and
median ones herbaceous except for a narrow
scarious margin, the apex acute or obtuse and
mucronulate. Receptacle flat, shallowly alveolate
with rounded teeth, or merely pitted in fruiting
heads. Florets 40-80 (or more). Pistillate florets
fertile, in several series, usually more numerous
than the disk florets, the corollas slender, tubular,
with a long-exserted pink style and no stamens;
rays (ligules) absent or rudimentary. Disk florets
perfect and fertile, the corollas tubular or
narrowly funnelform, 34.5 mm long, the limb
whitish aging to pink, with 5 very short lobes, the
tube slender, yellowish green, longer than the
limb. Style branches of the disk florets with
attenuate, papillate, sterile appendages that are

longer than the stigmatic lines. Pappus very
prominent, considerably overtopping the corolla,
the capillary bristles numerous, in 2 or 3 series,
pure white or faintly pink, subequal in length,
4-6 mm long, soft and silky, attenuate, and
minutely barbellate. Achenes oblong-obovoid,
slender, slightly compressed, 1.5—2.5 mm long
and 0.40.5 mm across, gray or whitish, often
with purple streaks, pilose or strigillose,
obscurely 24 ribbed. 2n = 14. Including B.
angusta (Lindley in Hooker) Britton in Britton &
Brown (see A.G. Jones 1984). [Aster brachyactis
S.F. Blake—G.N. Jones 1945, 1950, 1963; Jones
and Fuller 1955; Swink 1974; Mohlenbrock
1975, 1986; Swink and Wilhelm 1979; and
others. ]

August—October. Disturbed ground: waste
places, roadsides and railroad tracks. Occasional,
recorded from only five counties of the Chicago
region; adventive from the North and the West.
The species range extends from NT and James
Bay, ON, east to PQ, PE, and NB, and west
through the northern half of the Great Plains to
n. WA; the plants were also recorded from
scattered stations in s.e. WI, n. [A, w. MO,

w. NE, w. KS, w. OK, and s. and e. CO.

The species was originally described (under
the name Erigeron ciliatus Ledeb.) from
collections made in Siberia. North American
plants may be recognized as Brachyactis ciliata
subsp. angusta (Lindley in Hooker) A.G. Jones
[based on Tripolium angustum Lindley in
Hooker. Aster angustus (Lindley in Hooker)
Torrey & Gray non Nees—Pepoon 1927. B.
angusta—Hill 1902]. This taxon, however, can
probably not be distinguished from typical subsp.
ciliata on any basis other than geography.

Brachyactis is a small genus of two or three
species with the center of distribution probably
in North America. The genus can be distin-
guished from Aster and other members of the
Astereae by a combination of the following:
pistillate florets that lack or have inconspicuous
ray corollas, a prominent pappus that consider-
ably overtops the disk corollas and is composed
of numerous subequal bristles in several series,
and the uncommon basic chromosome number of
x = 7. Most authors retain the taxon as a section
of Aster (cf. Houle and Brouillet 1985; Semple
and Brouillet 1980a, b).

May 1989

Glossary of Descriptive Terms

abaxial side of an organ away from the axis

achene dry, indehiscent, one-seeded fruit

aciculiform needle-shaped

acuminate tapering more or less abruptly to a
slender sharp point, the margins somewhat
curved inward

acute terminating in a sharp point at an angle of

between 45° and 90°, the margins straight (see
also attenuate)

adaxial side of an organ facing the axis

adventive introduced from an adjacent or
nearby region and spreading in the new
region

alveolate honeycombed

anastomosing veins (or nerves) connected by
cross veins (or nerves)

anthesis flowering time, i.e., period during
which pollination takes place

apex; apical tip; pertaining to the tip

areole small space clearly marked out on a
surface

ascending directed upward after arising at an
oblique angle or on a curve

attenuate gradually tapering to a narrow and
slender tip at an angle of less than 45°, the
margins more or less straight (see also acute)

auriculate having auricles, i.e., with ear-shaped
appendages

barbellate with minute barbs

bracteate having bracts

bracteiform bractlike

bulliform appearing blistered, bubblelike

callus a hard protuberance or thickening

campanulate bell-shaped

capitulescence cluster (often a large assem-
blage) of flower heads (commonly syn-
onymized with inflorescence)

caudex the usually condensed, thick, tough,
persistent, largely underground base of an
otherwise herbaceous stem

caudiciform stem base shaped like a caudex,
i.e., condensed, thick, and tough

cauline pertaining to or arising from the stem

cespitose growing in tufts, e.g., several stems
from a common base

ciliate fringed with slender trichomes (see also
fimbriate)

ciliolate minutely ciliate, i.e., the trichomes
short

cinereous with ash-colored (light gray)
indument

ASTER AND BRACHYACTIS IN ILLINOIS 187

clasping the base of a leaf or petiole nearly or
completely surrounding the stem

clayellate minutely club-shaped, i.e., expanded
toward the apex

cordate heart-shaped, the base with rounded
lobes and a sinus

corolla perianth whorl composed of petals
(united in the Asteraceae)

corymbiform shaped like a corymb, i.e., a flat-
topped capitulescence with the lower (outer)
branchlets and peduncles longer than the
upper (inner) ones

crenate toothed with rounded teeth

cuneate wedge-shaped with the narrow end at
the point of attachment

cymiform shaped like a cyme, i.e., resembling
an inflorescence in which the central or
terminal flower opens first

cymule a small few-flowered cyme:; refers in
the Asteraceae to an ultimate cluster of the
cymiform capitulescence

deciduous falling off at maturity or after
withering

decumbent reclining on the ground but with
the tips ascending

decurrent pertaining to leaf bases or veins that
continue downward from the nodes in the
form of wings, ribs, or lines of indument

dichotomous forked with two nearly equal
branches

dilated expanded in width or widened in
diameter

disk floret flower of the central portion of a
head, the corolla regular, 5-lobed

divaricate spreading at a wide angle

erose with the margin appearing eroded or
gnawed

falcate sickle-shaped

fastigiate with the branches partway parallel
and close together; broomlike

filament the stalk portion of a stamen that
supports the anther

fimbriate fringed with coarse trichomes or
processes (see also ciliate)

foliaceous leaflike or leafy

funnelform resembling or shaped somewhat
like a funnel

fusiform spindle-shaped, swollen near the
middle and tapering at both ends

glabrescent nearly glabrous or becoming
glabrous

188 ILtinois NATURAL History SurvEY BULLETIN

glabrous smooth, i.e., without vestiture

glandular invested with glands, i.e., with
structures that produce a viscid sap

glaucous covered with a waxy, bluish bloom
that often rubs off easily

habit general appearance

head capitulum = unit of capitulescence (the
individual florets sessile)

hirsute having coarse or stiff, spreading or
ascending trichomes

hirtellous minutely hirsute, i.e., with short,
spreading trichomes

hispid having rigid spreading bristles or
spinules

hispidulous minutely hispid

hyaline translucent or colorless

imbricated overlapping, i.e., shinglelike in a
spiral arrangement

indument hairy covering

indurate hardened

inrolled rolled inward or downward at the edges

involucre one or more series of bracts (phyllar-
ies) subtending the florets of a head (in the
Asteraceae)

isodiametric with all the diameters of an areole
about equal in length

keel; keeled a central abaxial ridge formed by
the midrib of a compressed phyllary, bract, or
leaf; forming a keel

lanate woolly, with long, tangled, curly
trichomes

lanceolate lance-shaped; much longer than
broad, widest near the base and tapering to the
apex

ligule the strap-shaped limb of a ray floret in the
Asteraceae

limb upper, expanded portion of a corolla (in
disk florets, the portion above the zone of
attachment of the filaments) as distinct from
the tube portion

mucro_a sharp, short, and abruptly narrowed
point or protuberance

mucronate tipped with a mucro

mucronulate diminutive of mucronate

node _ the place on the stem where leaves are
attached and branches arise

ob- Latin prefix meaning inverted, e.g., obovate
= inverse of ovate

oblique slanting, not straight up, or at an angle
that is not 90°

oblong at least two times longer than broad with
nearly parallel sides

Vol. 34 Art. 2

obtuse blunt; if sharp-pointed, with an angle of
more than 90°

ovate; ovoid egg-shaped, with the broader part
near the base

panicle a variously branched compound
inflorescence (in the Asteraceae a compound
capitulescence with pedunculate flower heads)

paniculiform resembling or shaped like a
panicle

papillate nipplelike

Pappus the specialized outer perianth whorl in
the Asteraceae composed of bristles (in
Aster), awns, or scales

peduncle the stalk of a head

perfect having both functional stamens and
pistils

petiole; petiolate the stalk of a leaf; having a
stalk

phyllary bract of the involucre

pilose invested with long, soft, slender
trichomes

pinnate arising from both sides of the axis or
midrib

pistillate having pistils but no functional
stamens

polymorphic exhibiting several forms (morphs)
of the same organ

pro parte in part

pro sp. described as a species

puberulent minutely or finely hairy

pubescence; pubescent hairiness: hairy

pulvinate swollen or cushion-shaped

puncticulate having minute colored or
translucent dots

pustulate with pimplelike or blisterlike raised
areas

racemiform resembling or shaped like a
raceme, i.e., an elongate capitulescence or
branch with pedunculate heads

rameal of or pertaining to the branches

ray floret of the outer series of florets in a head,
the corolla strap-shaped (= ligulate)

receptacle the expanded end of the peduncle,
i.e., the end bearing the aggregate of florets

reflexed abruptly bent or tumed downward

remote distantly spaced

reticulate; reticulum interconnected like a
network

rhizome an elongate, often branched under-
ground stem, usually with minute scale leaves
and rooting at or producing new shoots from
the nodes

May 1989

rhombic shaped like a rhombus, 1.e., like an
equilateral, oblique-angled parallelogram;
more or less diamond-shaped

rosette a cluster of leaves with very short
internodes, arranged in a compact, spiral
(near-circular) series

rugose; rugulose wrinkled

scabrellous minutely scabrous

scabrous rough to the touch, the stiff trichomes
pointing in one direction

searious thin, dry, membranous, not green

secund directed to one side of the stem or
branchlet

sensu auct. according to author(s)

sensu lato ina broad sense

sensu stricto in a narrow sense

sericeous silky, with appressed, soft, glossy
trichomes

serrate with sharp teeth that point forward

serrulate finely serrate

sessile lacking a stalk

sheathing closely enveloping

spatulate spatula-shaped or spoon-shaped

spiciform shaped like a spike, i.e., a simple,
elongate capitulescence with sessile heads

spinule; spinulose short spine; minutely spiny

squarrose having the parts recurved at the tip
(mostly applied to phyllaries)

stamen pollen-bearing organ

stigmatic pertaining to the portion of the style
branches receptive to pollen

stoloniform shaped like stolons (applied to
long-creeping horizontal rhizomes)

ASTER AND BRACHYACTIS IN ILLINOIS 189

striate marked with fine longitudinal lines or
ridges

strigillose minutely strigose, i.e., the trichomes
very short

strigose with the trichomes appressed and lined
up in one direction

style the elongated part of the pistil above the
ovary

subulate awl-shaped; slender, tapering to a
sharp point

suffruticose plants woody only at the base and
herbaceous over most of the above-ground
portion

terete circular in cross section

throat expanding zone in a corolla at the
junction of tube and limb

thyrsiform shaped like a thyrse, i-e., like a
compact or contracted panicle

tomentose invested with short-haired, matted,
woolly pubescence

trichome an epidermal structure (hair, bristle,
or prickle)

truncate ending abruptly as if cut off

turbinate top-shaped, i.e., inversely conical

umbelliform resembling or shaped like an
umbel, i.e., a flat-topped capitulescence in
which several peduncles more or less arise
from a common point

urceolate urn-shaped, e.g., a corolla that is
abruptly contracted just below the mouth

vestiture any covering of the surface, i.e.,
indument and/or glands

villous woolly, the trichomes long, soft, and
curly but not matted or tangled

190 ILtinois NATURAL History SURVEY BULLETIN

Literature Cited

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Botany 72:268—277.

BENKE, H.C. 1928. Some Illinois asters and a
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BENKE, H.C. 1930. Aster amethystinus an
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Benke, H.C. 1932a. Some field notes: a new
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BENKE, H.C. 1932b. New asters from Illinois.
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FELL, E.W. 1955. Flora of Winnebago County,
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Vol. 34 An. 2

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HicLey, W.K., AND C.S. RappIN. 1891. The flora
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HILL, E.J. 1902. Notes on migratory plants.
Bulletin of the Torrey Botanical Club
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HoLMGREN, P. K., W. KEUKEN, AND E.K.
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Hou_e, F., AND L. BROUILLET. 1985. Chromo-
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37:369-372.

Jones, A.G. 1974. Aster ericoides L. east of the
Mississippi River. The American Midland
Naturalist 92:466—475.

Jones, A.G. 1977. New data on chromosome
numbers in Aster section Heterophylli
(Asteraceae) and their phylogenetic implica-
tions. Systematic Botany 2:334—-347.

Jones, A.G. 1978a. Observations on reproduction
and phenology in some perennial asters. The
American Midland Naturalist 99:184—197.

Jones, A.G. 1978b. The taxonomy of Aster
section Multiflori (Asteraceae).—I. Nomen-
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taxa. Rhodora 80:319-357.

Jones, A.G. 1978c. The taxonomy of Aster
section Multiflori (Asteraceae).—II. Biosys-
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Jones, A.G. 1980a. A classification of the New
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Jones, A.G. 1980b. Data on chromosome
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certain North American species. Brittonia
32:240-261.

Jones, A.G. 1982. Virgulus Raf. vis-a-vis Aster
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Jones, A.G. 1983. Nomenclatural changes in
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Jones, A.G. 1984. Nomenclatural notes on Aster
(Asteraceae)—II. New combinations and
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May 1989

Jones, A.G. 1985. Chromosomal features as
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Jones, A.G. 1987. New combinations and status
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Jones, A.G., AND P. HiepKo. 1981. The genus
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Jones, A.G., AND P.P. Lowry. 1986 [1987].
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Jones, A.G., AND D.A. YOUNG. 1983. Generic
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JONES, R.L. 1983. A systematic study of Aster
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ASTER AND BRACHYACTIS IN ILLINOIS 19]

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cytogeography of Aster pilosus (Composi-
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(Compositae: Astereae): a multivariate
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Wisconsin. The American Midland Naturalist
26:398-420.

SHINNERS, L.H. 1949. Aster ontarionis the same
as A. pantotrichus (A. missouriensis).
Rhodora 51:89-91.

STEYERMARK, J.A. 1963. Flora of Missouri. lowa
State University Press, Ames.

192 ILtinois NATURAL History SURVEY BULLETIN

Swink, F. 1974. Plants of the Chicago region.
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SwINK, F., AND G. WILHELM. 1979. Plants of the
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Torrey, J., AND A. Gray. 1841. Flora of North
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New York.

Vol. 34 Art. 2

WETMORE, R.H., AND A.L. DELISLE. 1939.
Studies in the genetics and cytology of two
species in the genus Aster and their polymor-
phy in nature. American Journal of Botany
26:1-12.

WIEGAND, K.M. 1928. Aster lateriflorus and
some of its relatives. Rhodora 30:161-179.

WIEGAND, K.M. 1933. Aster paniculatus and
some of its relatives. Rhodora 35:16-38.

May 1989 ASTER AND BRACHYACTIS IN ILLINOIS 193

Index to Scientific Species Names, Including Synonyms

The names of accepted Illinois species are given
in roman type. Bold-faced page numbers indicate
species descriptions.

Aster X amethystinus 141, 146, 156, 166, 166

angustus 186

anomalus 144, 145, 149, 180

azureus 139, 169

borealis 139, 141, 144, 148, 150-151, 161

brachyactis 139, 143, 186

carneus 174

chasei 139, 141, 178

ciliolatus 141, 144, 146, 152, 152

commutatus 156

cordifolius 141, 144, 146, 151-152, 152, 185

depauperatus 170

diffusus 162

divaricatus 159

drummondii 144, 145, 152, 153-154, 169,
184

dumosus 141, 144, 148, 151, 154-155, 158,
162

eatonii 151

ericoides 139, 141, 144, 147, 155-156, 156,
166, 170, 172

exiguus 156

falcatus 141, 144, 147, 156

finkii 154

firmus 176

fragilis 140, 144, 148, 155, 157-158, 161,
162, 163, 168

furcatus 141, 144, 145, 158-159

hirsuticaulis 162

hirtellus 185

horizontalis 162

infirmus 182

interior 161

junceus 150

junciformis 139, 150

Kumleini 167

laevis 144, 145, 147, 152, 159-160, 161,
169, 177

lanceolatus 139, 144, 148, 151, 155, 158,
160, 160-161, 163, 168, 174, 177

lateriflorus 144, 148, 158, 160, 161,
162-163, 168, 177

linariifolius 144, 146, 163

lindleyanus 152

longifolius 151, 176, 177

lucidulus 176

lutescens 140

macrophyllus 141, 144, 145, 164-165

miser 162

missouriensis 168

multiflorus 156

novae-angliae 141, 144, 146, 156, 165-166,
166

novi-belgii 176

occidentalis 151

oblongifolius 144, 146, 166-167

ontarionis 139, 144, 148, 158, 163, 167-168

oolentangiensis 139, 144, 145, 148, 160,
168-169, 184

paniculatus 161

pantotrichus 139, 168

parviceps 139, 141, 144, 147, 169-170

patens 144, 146, 170-171, 176, 177

patentissimus 171

pendulus 162

pilosus 139, 144, 147, 156, 170, 171-173

polyphyllus 172

praealtus 144, 147, 173-174, 177

prenanthoides 141, 144, 147, 174-175

pringlei 172

ptarmicoides 139, 140, 143

pubentior 182

puniceus 144, 147, 160, 161, 174, 175-177

sagittifolius 152, 153, 185

salicifolius 174

schreberi 141, 144, 145, 177-178

sericeus 144, 146, 178-179

shortii 144, 145, 149, 154, 179-180

simplex 139, 161

subasper 174

tataricus 139, 141, 144, 147, 180-181

tenuifolius 170

tradescantii 161

turbinellus 144, 147, 181

umbellatus 144, 146, 182-183

undulatus 141, 144, 145, 153, 154, 183-184

urophyllus 144, 146, 152, 154, 180,
184-185

villosus 172

vimineus 157, 161, 162

Brachyactis angusta 186

ciliata 140, 143, 145, 186

Diplopappus linariifolius 163

lutescens 140

Doellingeria umbellata 182
Erigeron ciliatus 186
Solidago x lutescens 140

ptarmicoides 143
riddellii 140

Tripolium angustum 186

194

I-cinois NATURAL History SurveY BULLETIN

Index to Vernacular (Common) Names

Aromatic Aster 166
Azure Aster 168
Big-leaved Aster 164
Blue Aster 149

Blue Wood Aster 151
Brittle Aster 140, 157
Bushy Aster 154
Calico Aster 162
Crooked Aster 174
Crooked-stemmed Aster 174
Drummond's Aster 153
Flat-top Aster 182
Flax-leaved Aster 163
Forked Aster 158
Frost-flower 143
Frost-weed Aster 171
Hairy Aster 171
Heart-leaved Aster 151
Heath Aster 155
Large-leaved Aster 164
Late Purple Aster 170
Lindley’s Aster 152
Many-rayed Aster 149
Michaelmas Daisy 143
New England Aster 165
Ontario Aster 167
Panicled Aster 160
Purple-stemmed Aster 175

Rayless Aster 186
Red-stemmed Aster 175
Rice-button Aster 154
Rush Aster 150
Savory-leaved Aster 163
Schreber’s Aster 177
Short’s Aster 179
Side-flowered Aster 162
Silky Aster 178
Sky-blue Aster 168
Small-headed Aster 169
Small White Aster 157
Smooth Aster 159
Smooth Blue Aster 159
Spreading Aster 170
Starved Aster 162
Starwort 143

Stuff Aster 139

Swamp Aster 175
Tartarian Aster 180
Wavy-leaved Aster 183
Western Heath Aster 156
White Prairie Aster 155
White Woodland Aster 162
Wild Aster 143

Willow Aster 173
Willow-leaved Aster 173
Wreath Aster 155

Vol. 34 An. 2

Illinois Natural History Survey
607 East Peabody Drive
Champaign, Illinois 61820

A Division of the Illinois Department of Energy and Natural Resources

NATL §

R ILLINOIS

NOV <0 199 NATURAL

“HISTORY

LIBRARY of eee
wee

ILLINOIS
NATURAL
HISTORY
SURVEY

A Nomenclator of Leptosphaeria
V. Cesati & G. de Notaris

(Mycota-Ascomycotina-Loculoascomycetes)

J.L. Crane

Center for Biodiversity
Illinois Natural History Survey

C.A. Shearer

Department of Plant Biology
University of Illinois at Urbana-Champaign

Illinois Natural History Survey Bulletin
Volume 34, Article 3
March 199]

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

A catalog of the publications of the Illinois Natural History Survey is
available without charge from the address below. A price list and an
order blank are included with the catalog.

Illinois Natural History Survey
Distribution Center

Natural Resources Building
607 East Peabody Drive
Champaign, Illinois 61820

Citation:

Crane, J.L., and Shearer, C.A. 1991. A Nomenclator of Leptosphaeria
V. Cesati & G. de Notaris (Mycota-Ascomycotina-Loculoascomycetes).
Illinois Natural History Survey Bulletin 34(3):195—355.

Editor: John P. Ballenot
Designer: Gail Glende Rost

US ISSN 0073-4918

Printed by Authority of the State of Illinois
(X11281-MD-3-91)

Contents

Acknowledgments _ iv

Introduction and Historical Background 195

List of Leptosphaeria Species, Varieties, and Forms 198
Host Index 291

Host Family Index 308

Substrate Index 318

Geographic Index 328

Appendix |. Taxonomic Division of Leptosphaeria 339
Appendix 2. Genera Historically Allied to Leptosphaeria 341
Appendix 3. Synonyms of Leptosphaeria 347

Appendix 4. Anamorphs of Leptosphaeria 348

Literature Cited 351

Acknowledgments

We express sincere appreciation to our col-
leagues at the Illinois Natural History Survey
and the University of Illinois at Urbana-
Champaign who assisted us in the completion
of this nomenclator. Carla G. Heister and
Monica A. Lusk located many references in the
University of Illinois library system from
cryptic citations. Martha Landis is noted for
her ability to locate references that cannot be
found by others. Helen F. Sullivan and Susan
Burke assisted us with many of the slavic
references. Dr. Ken Robertson assisted with the
verification of host nomenclature.

Mycological literature is so diverse that
no one library can encompass it all. Therefore,
we are especially indebted to Dr. S. Udagawa
at the National Institute of Hygienic Sciences in
Tokyo, Japan, who so kindly supplied copies of
the Japanese papers. Dr. Robbin C. Moran at
the Missouri Botanical Garden in St. Louis
provided articles from journals not available in
the University of Illinois library system. Drs.
E. Miiller of Switzerland, F.A. Stafleu of the
Netherlands, J. Rammeloo of Belgium, P.
Fanton of the University of Padua, Italy, Clark
T. Rogerson and G.J. Samuels of the New York
Botanical Garden, Bronx, and Richard P. Korf
of Cornell University, Ithaca, New York,
supplied copies of species descriptions from

rare mycological books, journals, and exsiccati
labels. We are most grateful to Drs. Lekh Batra
and David Farr at the National Fungus Collec-
tions in Beltsville, Maryland, and to Drs.
Donald Pfister and Jean Boise at the Farlow
Herbarium and Library of Harvard University
in Cambridge, Massachusetts, for their kind
hospitality during our visits, and for making
available to us their rich mycological libraries
and collections.

Several individuals helped us in a variety
of ways, and without their willingness to con-
tribute time and expertise this project would not
have been completed. Betty A. Nelson typed
the original manuscript and its several revisions
and checked for inconsistencies in author and
journal citations. She and Patty L. Duzan spent
many hours on computer-related problems in
sorting this nomenclator. We also thank Sabine
M. Huhndorf, who supplied several names of
Leptosphaeria species.

We gratefully acknowledge Dr. Richard
T. Hanlin of the University of Georgia, Athens,
for critically reviewing the manuscript and
John P. Ballenot for editing the manuscript.
Support for this study was provided by the
National Science Foundation, Systematic
Biology Program, grant NSF-BSR-87-00065.

Introduction and Historical Background

This nomenclator of Leptosphaeria is an index
to names published in the genus through 1989
and includes the source of publication, nomen-
clatural history, substrate and/or host, and
geographical location for each species name.
Orthographic errors of specific epithets were
corrected, with the original spelling given in
brackets. Full names and initials of authorities
are given. Standardized, complete names of
periodicals are cited following the system of
Brown and Stratton (1963). Porter and Koster
(1970), Koster and Gascoigne (1971), and
Smits (1968). The titles of books and pam-
phlets are cited according to Stafleu and Cowan
(1976-1988). Obligate synonomy is indicated
by the symbol =. Articles of the International
Code of Botanical Nomenclature cited in this
work follow Greuter et al. (1988). Several
authors have described new species in Lepto-
sphaeria as trinomials in which the subgenus or
group to which the new species belongs is
indicated as a middle name in parentheses; an
example is Leptosphaeria (Clypeosphaeria)
hendersoniae. Such trinomials are alphabet-
ized by specific epithet; however, the group
name precedes the epithet.

The scientific names of host plants are
given as reported in the protologue of each
Leptosphaeria species, with the currently
accepted name in parentheses. Several host
epithets reported in the original descriptions
were never published or could not be verified.
These are changed to “sp.” Plant host families
and genera were verified in Willis (1973), Farr
et al. (1979), and Cronquist (1981). Specific
epithets of vascular plants were confirmed in
Halliday and Beadle (1983), Kartesz and
Kartesz (1980), the Gray Herbarium Card
Index (1894—present), and Index Kewensis
(1895-present). Names of pteridophytes were
substantiated in Christensen (1905-1906), and
names of mosses, in Wijk et al. (1959-1960).
Lichenized fungi names follow Zahlbruckner
(1921-1940) and Lamb (1963), and fungal host
names were verified in Saccardo (1882-1931)

and the Index of Fungi (1920-1987). For
convenience, Leptosphaeria species reported
from Algae, Fungi, Lichenes, and Musci are
listed under these headings and their respective
hosts in the host index. When available,
specific substrate information, such as leaf,
stem, water, etc., is given.

The geographical location of species is
usually described by country and is based on
information in the protologue and new combi-
nations. Geographical names were verified in
Seltzer (1952).

Cesati and de Notaris (1863) established
the genus Leptosphaeria and included 26
species; among these was Leptosphaeria hirta
(G.L. Rabenhorst) V. Cesati & G. de Notaris,
the type species of an earlier genus, Nodu-
losphaeria G.L. Rabenhorst, 1858. Lepto-
sphaeria was conserved against Nodu-
losphaeria with Leptosphaeria doliolum (C.H.
Persoon:E.M. Fries) V. Cesati & G. de Notaris
as the type species (Greuter et al. 1988). The
original description of Leptosphaeria was
superficial by modern taxonomic standards,
and the genus was delimited largely by asco-
spore characteristics. The ascospore character-
istics selected (oblong or fusoid, two- to many-
celled, hyaline, becoming yellowish or dark
brown) and the poor characterization of other
structural features resulted in the inclusion of a
wide range of Ascomycetes in this genus.
Leptosphaeria now comprises approximately
1,689 taxa. These taxa represent, according to
present-day concepts of ascomycete classifica-
tion, a melange of Euascomycetes and Locu-
loascomycetes.

Few mycologists have attempted to deal
with the systematics of Leptosphaeria. Sac-
cardo (1878, 1883, 1891, 1895, 1899, 1913,
1928) recognized 800 species, which he
grouped primarily according to host (parasites
of dicotyledons, monocotyledons, and crypto-
gams) and habit (species on stems and/or
branches and leaves). Species were further
subdivided, based on the external features of

196 Illinois Natural History Survey Bulletin

the pseudothecium (glabrous, hairy, setose) and
ascospore septation. Saccardo’s concept of
Leptosphaeria was broad, and his emended
description specified the presence of pseu-
doparaphyses (as paraphyses).

H6hnel (1907a) was the first to use
information on centrum structure for the
classification of Leptosphaeria-like fungi. He
established the family Pseudosphaeriaceae for
species in which the asci grow up into a cellular
tissue that occupies the inner space of the
fruiting body. As the asci develop, the cellular
tissue becomes compressed and at fruiting-
body maturity simulates paraphyses similar to
those found in the Sordariales (Pyrenomy-
cetes). In the Sordariales, however, the
paraphyses originate prior to the asci, and the
asci grow up among them. Hohnel included
four genera in the Pseudosphaeriaceae [Pseu-
dosphaeria F. v. Hohnel, Pyrenophora F. v.
Hohnel, Scleroplea (P.A. Saccardo) C.A.
Oudemans, and Wettsteinina F. v. Hohnel].
Later, Hohnel (1918a, 1918b) divided Lepto-
sphaeria into three genera based on centrum
structure (Leptosphaeria, Scleropleella F. v.
Hohnel, and Nodulosphaeria G.L. Rabenhorst)
(Appendix 1). Petrak (1923) subsequently
provided evidence that Leptosphaeria is related
to Wettsteinina and belongs in the Pseudo-
sphaeriaceae.

Wehmeyer (1942) reported the occur-
rence of 13 species of Leptosphaeria from
North America and described two new species.
He noted, “Many of the species show minor
differences of spore structure, often correlated
with the host. In the descriptive literature,
these details are not always given. Large
numbers of species have been described,
largely on host distinction, on the one hand,
whereas many host varieties have been ob-
scured by inclusion in one of the ubiquitous
species on the other hand. As a result it is
difficult to be sure of species determinations
without a comparative study of the entire
genus.” This statement remains valid today.
Wehmeyer (1946) also redescribed seven
species of Leptosphaeria and discussed
evolutionary trends within the genus. He noted
that 382 species have the 3-septate type of
ascospore and believed that the 3-septate, dark
brown spore type is the primitive state. Ac-
cording to Wehmeyer, the spores of more
recently evolved species are lighter in color,

Vol. 34 Art. 3

longer, narrower, and more septate. These
trends lead directly, without a break, to
Ophiobolus H. Riess.

Miiller (1950), circumscribing Lepto-
sphaeria broadly, considered 114 species oc-
curring in Switzerland. Within his broad
generic concept, Miiller divided Leptosphaeria
into four sections (Appendix 1). These sections
were circumscribed based on pseudothecial and
centrum structure and ascospore morphology.
Three of his sections correspond, in part, to the
three groups of Hohnel (Appendix 1).

Munk (1957) accepted Miiller’s sections
with certain reservations and divided Lepto-
sphaeria into four somewhat different sections
(Appendix 1). Sections I (Eu-Leptosphaeria),
Ill (Scleropleella), and IV (Nodulosphaeria)—
which correspond to Miiller’s Sections II, I, and
IV, respectively—were more restricted than
those of Miiller. The remaining species of Lep-
tosphaeria were treated in Section II (Para-
Leptosphaeria).

Holm (1957), in a treatment of 62
Swedish species of Leptosphaeria, rejected
Miiller’s broad concept of Leptosphaeria and
limited the genus to those species most closely
related to Leptosphaeria doliolum, the type of
the genus (Appendix 1). Excluded species
were distributed primarily in Nodulosphaeria
H. Riess, Phaeosphaeria 1. Miyake, and
Entodesmium H. Riess. Holm placed much
emphasis on substrata and relation of the
pseudothecium to the substratum. Holm’s
disbursement of Leptosphaeria species was
rejected by both Dennis (1978) and Sivanesan
(1984) but was accepted by Hedjaroude (1969),
v. Arx and Miiller (1975), Eriksson (1967), and
Shoemaker (1984).

In the past 50 years, there have been a
large number of intergeneric transfers of
Leptosphaeria species. Many of these transfers
have come about by the partitioning of species
groups into new (Paraphaeosphaeria O.
Eriksson) or existing (Entodesmium H. Riess
and Phaeosphaeria I. Miyake) genera. Other
species have been transferred to genera very
similar to and integrating with Leptosphaeria
(Lidophia J.C. Walker & B.C. Sutton, Massaria
G. de Notaris, Massarina P.A. Saccardo,
Melanomma T.R.J. Nitschke ex L. Fuckel,
Ophiobolus H. Riess, and Wettsteinina F. v.
HGhnel). Descriptions of related genera and a
synopsis of their relationships to Leptosphaeria

March 199]

are summarized in Appendix 2. Genera
synonymous with Leptosphaeria are listed in
Appendix 3.

In recent years, Leptosphaeria has been
included in two orders in the Bitunicatae or
Loculoascomycetes. Luttrell (1973) placed
Leptosphaeria in the Pleosporales under the
Pleosporaceae. Von Arx and Miiller (1975)
retained Leptosphaeria in the Pleosporaceae
under the Dothideales. Hawksworth et al.
(1983) placed Leptosphaeria in the Dothideales
in either the Phaeosphaeriaceae or the Pleospo-
raceae. Eriksson and Hawksworth (1986)
classified the genus in Dothideales under
Phaeosphaeriaceae. Most recently, Barr
(1987a) has placed Leptosphaeria in the

Nomenclator of Leptosphaeria

Leptosphaeriaceae of the Pleosporales in the
Loculoascomycetes.

One of the most intriguing aspects of
Leptosphaeria is the diversity of anamorphic
States attributed to this genus. Anamorphs of
Leptosphaeria have been demonstrated for 70
species, and most of these associated ana-
morphs are Coelomycetes (Appendix 4). The
largest numbers of Leptosphaeria associations
are with Phoma P.A. Saccardo and Stagono-
spora (P.A. Saccardo) P.A. Saccardo.

It is intended that this nomenclator will
be the base for a reappraisal of Leptosphaeria
using modern concepts and techniques appli-
cable to Loculoascomycetes.

197

List of Leptosphaeria Species, Varieties, and Forms

abbreviata (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:26. 1883.
=Sphaeria abbreviata M.C. Cooke, Handbook
of British Fungi, p. 893. 1871. On dead stems
of Rubus sp., Rosaceae. Great Britain.

abuensis K.S. Panwar & S.J. Kaur, Kavaka, Madras
3:67-68. 1975. On dead wood of unknown
host, unknown family. India.

abutilonis M. Chochrjakov in V. Tranzschel, L.
Gutner, and M. Chochrjakov, Trudy Instituta
Novogo Lubyanogo Syr’ya, Moskva 4:133.
1933. On leaves of Abutilon avicennae
Gaertner, Malvaceae. U.S.S.R.

abutilonis L.E. Wehmeyer & S. Ahmad, Biologia.
Biological Society of Pakistan, Lahore 10:12.
1964. Nom. illegit. Art. 64.1. On unknown
substrate of Abutilon indicum (L.) Sweet,
Malvaceae. Pakistan.

acanthi N.T. Patouillard, Revue Mycologique,
Toulouse 8:181. 1886. On dead stems of
Adhatoda sp., Acanthaceae. China.

aceris N.N. Woronichin, Vestnik Tiflisskogo
Botanicheskogo Sada, Tiflis (Moniteur du
Jardin Botanique de Tiflis) 35:5. 1914. On
living leaves of Acer laetum C.A. Mey.,
Aceraceae. U.S.S.R. (Caucasus).

acheniarum K. Starback [see Leptosphaeria agnita
var. acheniarum K. Starback].

achilleae (B. Auerswald) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:236. 1863. =Sphaeria
achilleae B. Auerswald in G.L. Rabenhorst,
Klotzschii Herbarium Vivum Mycologicum
Sistens Fungorum Per Totam Germaniam

Cresentium Collectionem Perfectam, Dresden,

Edition 1, Century 15, No. 1448. Anno 1850.
On stems of Achillea millefolium auct.,
Compositae. Germany.

acicola (L. Fuckel) P.A. Saccardo, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:55. 1883. =Ple-
ospora? acicola L. Fuckel, Symbolae
Mycologicae, Erster Nachtrag, p. 301. 1871.
On fallen needles of Pinus sylvestris L.,
Pinaceae. Germany.

aconiti P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica

Italiana, Firenze 7:315. 1875. On dry stems
of Aconitum napellus L., Ranunculaceae.
Italy.

(Metasphaeria) acorella M.C. Cooke, Grevillea,
London 13:99. 1885. =Metasphaeria acorella
(M.C. Cooke) A.N. Berlese & P. Voglino,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo Addi-
tamenta Ad Volumina I-IV, p. 158. 1886. On
leaves of Acorus calamus L., Araceae. Great
Britain.

acori P.A. Karsten, Hedwigia, Dresden 22:179.
1883. [Ad interim.] Nom. inval. Art. 34.1.
On decaying leaves of Acorus calamus L.,
Araceae. Finland.

aculeorum G. Passerini, Atti della R. Accademia dei
Lincei Memoriae, Rome, Series 4, 6:458-459.
(1889) 1890. On stems of Rosa canina L.,
Rosaceae. Italy.

acuta H. Rehm, Ascomyceten, Fascicle 16, No. 783.
Anno 1884. Nom. illegit. Art. 64.1. =Lepro-
sphaeria acutiuscula A.N. Berlese.

acuta (G.F. Hoffmann:E.M. Fries) P.A. Karsten,
Mycologia Fennica Pars 2, Pyrenomycetes,

p. 98. 1873. =Sphaeria acuta GF.
Hoffmann:E.M. Fries, Vegetabilia Crypto-
gama, Fascicle 1, p. 22. 1787; E.M. Fries,
Systema Mycologicum Sistens Fungorum
2:507. 1823. =Pleospora acuta (G.F.
Hoffmann:E.M. Fries) L. Fuckel, Symbolae
Mycologicae, p. 135. 1870. =Ampullina
acuta (G.F. Hoffmann:E.M. Fries) L. Quélet.
Mémoires de la Société d’Emulation de
Montbéliard, Series 2, No. 5, pp. 523-524.
1875; Les Champignons du Jura et des
Vosges 3:95. 1875. On fallen stems of Urtica
dioica L., Urticaceae. Austria, Belgium,
Finland, France, Germany, Great Britain,
Italy, U.S.A.

acuta (G.F. Hoffmann:E.M. Fries) P.A. Karsten
forma insignis F. Fautrey in C. Roumeguére,
Fungi Selecti Gallici Exsiccati, Century 72,
No. 7137. Anno 1897; Revue Mycologique,
Toulouse 19:149. 1897. On dry, decorticated
stems of Urtica dioica L., Urticaceae. France.

acuta (G.F. Hoffmann:E.M. Fries) P.A. Karsten
forma urticae F. Fautrey in C. Roumeguére,
Revue Mycologique, Toulouse 16:7. 1894;

March 199]

Fungi Selecti Gallici Exsiccati, Century 65,
No. 6434. Anno 1894. On diseased stalks of
Urtica dioica L., Urticaceae. France.

acutispora §. Toth, Omagiu Lui Traian Savulescu
cu Prilejul Implinirii A 70 De Ani, Academia
Republicii Populare Romane, Bucharest, p.
778. 1959. On dry stems of Erysimum
diffusum Ehth., Erysimum erysimoides (L.)
Fritsch ex Janchen, Sisymbrium strictissimum
L., Cruciferae. Romania.

acutiuscula A.N. Berlese, cones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:86.
1894. =Leptosphaeria acuta H. Rehm. On
stems of Urtica sp., Urticaceae. Germany.

adesmicola (C.L. Spegazzini) L. Holm, Svensk
Botanisk Tidskrift, Stockholm 62:224—225.
1968. =Gibberidea adesmicola C.L.
Spegazzini, Anales del Museo Nacional de
Historia Natural de Buenos Aires 19(Series 3,
12):385. 1909. On dead branches of Adesmia
sp., Leguminosae. Argentina.

advenula (W. Nylander) P.A. Saccardo & D.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 17:731. 1905. On thallus of Lecidea
excentrica Roehling, Lichenes (Lecideaceae).
Ireland, Italy.

aegira P.A. Saccardo & C.L. Spegazzini in P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
1:37. 1877. On wilting leaves of Populus
alba L., Salicaceae. Italy.

aeluropodis A.1. Lobik, Materialy po Floristicheskim
i Faunisticheskim Obsledovaniyam Terskogo
Okruga, pp. 22-23. 1928 [as aeluropi]. On
leaves of Aeluropus littoralis (Gouan.) Parl.,
Gramineae. U.S.S.R.

aerea C.L. Spegazzini, Anales de la Sociedad
Cientifica Argentina, Buenos Aires
12:178-179. 1881. Fungi Argentini Pugillus
4, No. 166. 1881. =Sphaerulina (Leptosphae-
rella) aerea (C.L. Spegazzini) M.C. Cooke,
Grevillea, London 18:80. 1890. =Pleospora
pellita (E.M. Fries) G.L. Rabenhorst var.
pellita, fide L.E. Wehmeyer, A World
Monograph of Pleospora, p. 47. 1961. On
branches and dead leaves of Eryrhrina crista-
galli L., Tillandsia bicolor Brongn., Legumi-
nosae, Bromeliaceae. Argentina.

aetnensis G. Scalia, Prima Contribuzione alla
Conoscenza delle Flora Micologica della
Provincia di Catania, p. 16. 1899. On dry
branches of Smilax aspera L., Smilacaceae.
Italy.

affinis P.A. Karsten, Fungi Fenniae Exsiccati,
Century 9, No. 896. Anno 1869. =Metas-
phaeria affinis (P.A. Karsten) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Diggesit P.A. Saccardo 2:159.

Nomenclator of Leptosphaeria 199

1883. On dead stems of Rhinanthus sp.,
Scrophulariaceae. Finland.

africana A.M. Saccas, Etude de la Flore Crytoga-
mique des Caféiers en Afrique Centrale.
Bulletin de Institut Frangais du Café du Cacao
et d’Autres Plantes Stimulantes (Bulletin
IFCC No. 16), pp. 219-221. 1981. Nom.
inval. Art. 37.1. On dead branches of Coffea
robusta L. Linden (=Coffea canephora Pierre
ex Froehn.), Rubiaceae. South Africa.

agaves H. Sydow, P. Sydow, & E.J. Butler, Annales
Mycologici, Berlin 9:409. 1911. On wilted
leaves of Agave rigida Mill. var. sisalana
(Perrine) Engelm., Agavaceae. India.

aglaja P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:310. 1875. On leaves of
Oxalis stricta L., Oxalidaceae. Italy.

agminalis P.A. Saccardo & P. Morthier in P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Ilustrans
1:498-499. 1879. =Dothideopsella agminalis
(P.A. Saccardo & P. Morthier) F. v. Héhnel,
Sitzungsberichte der Akademie der Wissen-
schaften in Wien, Mathematisch-
naturwissenschaftliche Klasse, Abt. I, 124(1
& 2):70. 1915. On stems of Clematis vitalba
L., Ranunculaceae. France, Italy.

agminalis P.A. Saccardo & P. Morthier forma minor
F. Fautrey in C. Roumeguére, Revue
Mycologique, Toulouse 13:168. 1891; Fungi
Selecti Gallici Exsiccati, Century 59, No.
5849. Anno 1891. On unknown substrate of
Clematis vitalba L., Ranunculaceae. France.

agnita (J. Desmaziéres) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:236. 1863. =Sphaeria
(Caulicola) agnita J. Desmaziéres, Annales
des Sciences Naturelles, Paris, Botanique,
Series 3, 16:313. 1851. On stems of Valeri-
ana officinalis L., Valerianaceae. Italy.

agnita (J. Desmaziéres) V. Cesati & G. de Notaris
subsp. /abens P.A. Saccardo & G. Scalia in
P.A. Saccardo, C.H. Peck, and W. Trelease,
Harriman Alaska Expedition 5:330. 1904. On
dead herbaceous stems of unknown host.
U.S.A.

agnita (J. Desmaziéres) V. Cesati & G. de Notaris
var. acheniarum K. Starbick, Arkiv for
Botanik, Uppsala, Stockholm 5(7):23. 1905.
[Raised to species rank; Leptosphaeria
acheniarum K, Starbick in reprint on same
page.] On dry achenes of Mikania sp.,
Compositae. Bolivia.

agnita P.A. Saccardo var. ambigua A.N. Berlese,
Atti dell’ Accademia Scientifica Veneto-
Trentino-Istriana, Padova, Series 1,
9:247-248. 1886; Ricerche intorno alla
Leptosphaeria agnita (Desm.) Ces. et de Not.,
ed alla L. ogilviensis (B. et Br.) Ces. et de

200

Not., pp. 4-5. 1886. On dry stems of
Eupatorium cannabinum L., Compositae.
Germany.

agnita (J. Desmazieres) V. Cesati & G. de Notaris
var. bupleuri P.A. Saccardo, Bulletin. Société
R. de Botanique de Belgique, Bruxelles 28:91.
1889. On dead stems of Bupleurum falcatum
L., Umbelliferae. U.S.S.R.

agnita P.A. Saccardo var. chrysanthemi A.N.
Berlese, Atti dell’ Accademia Scientifica
Veneto-Trentino-Istriana, Padova, Series 1,
9:246-247. 1886; Ricerche intorno alla
Leptosphaeria agnita (Desm.) Ces. et de Not.,
ed alla Leptosphaeria ogilviensis (B. et Br.)
Ces. et de Not., pp. 3-4. 1886. On dry stems
of Chrysanthemum corymbosum L., Composi-
tae. Germany.

agnita (J. Desmazieres) V. Cesati & G. de Notaris
var. erigerontis A.N. Berlese, Atti dell’
Accademia Scientifica Veneto-Trentino-
Istriana, Padova, Series 1, 9:247. 1886;
Ricerche intorno alla Leptosphaeria agnita
(Desm.) Ces. et de Not., ed alla Leptosphaeria
ogilviensis (B. et Br.) Ces. et de Not., p. 4.
1886. On dead stems of Erigeron canadensis
L., Compositae. U.S.A.

agnita (J. Desmazieres) V. Cesati & G. de Notaris
var. major P.A. Saccardo & A.N. Berlese, Atti
del Istituto Veneto di Scienze, Lettere ed Arti,
Venezia, Series 6, 3:730. 1885. On stems of
Chaerophyllum aureum L., Umbelliferae.
Switzerland.

agnita P.A. Saccardo var. major A.N. Berlese, Atti
dell’ Accademia Scientifica Veneto-Trentino-
Istriana, Padova, Series 1, 9:247. 1886;
Ricerche intorno alla Leptosphaeria agnita
(Desm.) Ces. et de Not. ed alla Leptosphaeria
ogilviensis (B. et Br.) Ces. et de Not., p. 4.
1886. On dry stems of Chaerophyllum
aureum L., Umbelliferae. Germany.

agnita (J. Desmaziéres) V. Cesati & G. de Notaris
var. trifolii R. Gonzalez Fragoso, Memorias
de la R. Sociedad Espanola de Historia
Natural, Madrid 11:90. 1919. On dry stems of
Trifolium angustifolium L., Leguminosae.
Spain.

ahmadii F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 8:167. 1954. On dry branches of
Capparis aphylla Roth, Capparaceae.
Pakistan.

ailanthi P.A. Karsten & P.A. Hariot, Journal de
Botanique, Paris 3:206. 1889. On branches of
Ailanthus glandulosa Dest., Simaroubaceae.
France.

akagiensis E.A. Vainio, Botanical Magazine, Tokyo
35:79. 1921. On tree bark of unknown host,
unknown family. Japan.

albopunctata (G.D. Westendorp) J.B. Ellis & B.M.
Everhart, The North American Pyrenomy-
cetes. A Contribution to Mycologic Botany,

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

p- 375. 1892. =Sphaeria albopunctata G.D.
Westendorp.

albopunctata (G.D. Westendorp) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:72.
1883. =Sphaeria albopunctata G.D.
Westendorp, Bulletins de L’ Académie Royale
des Sciences, des Lettres et des Beaux-Arts de
Belgique, Brussels, Series 2, 7:87—88. 1859.
=Heptameria (Leptosphaeria) albopunctata
(G.D. Westendorp) M.C. Cooke, Grevillea,
London 17:32. 1889. =Phaeosphaeria
albopunctata (G.D. Westendorp) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1566. 1989. On seeds of Phrag-
mites communis Trin., Gramineae. Belgium.

albulae E. Miller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):281. 1950. On dead stems
of Lonicera alpigena L., Caprifoliaceae.
Switzerland.

(Leptosphaerella) alcides P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 8:176.
1876. On underside of leaves of Populus alba
L., Salicaceae. Italy.

alcides P.A. Saccardo forma quercina R. Ciferni,
Annales Mycologici, Berlin 20:51. 1922. On
dry leaves of Quercus robur L., Fagaceae.
Italy.

alexandrinis A. Negru, Mycopathologia et Mycolo-
gia Applicata, Den Haag 33:365—366. 1967.
On leaves of Cucumis sativus L., Cucurbita-
ceae. Romania.

algarbiensis M.T. Lucas, Memorias da Sociedade
Broteriana, Coimbra 21:34. 1970. On dead
leaves of Chamaerops humilus L., Palmae.
Portugal.

algida E. Rostrup, Meddelelser om Gronland,
Kjobenhavn 3:558. 1888. On leaves of
Catabrosa algida Fr., Gramineae. Greenland.

alhagii 1.P. Frolov, Novosti Sistematiki Nizshikh
Rastenij, Novitates Systematicae Plantarum
Non Vascularium 7:185—186. 1970. On dry
branches of Al/hagi sp., Leguminosae.
U.S.S.R.

alhaginis 1.P. Frolov in E.N. Koschkelova, LP.
Frolov, and Z. Dzhuraeva, Mikoflora
Badkhyza, Karabilya i Yuzhnoi Chasti Mur-
gabskogo Oazisa (Mikromitsety) [The
Mycoflora of Badkhyz, Karabil and the
Southern Part of the Murgab Oasis (Micromy-
cetes)], p. 86. 1970. Nom. nud. Art. 32.1.
=Leptosphaeria alhagii \.P. Frolov.

aliena H. Sydow, Annales Mycologici, Berlin
35:28-30. 1937. On leaves and culms of
Cladium articulatum R. Br., Cyperaceae.
Australia (New South Wales).

alliariae (L. Fuckel) J. Schroter in F.J. Cohn,
Kryptogamen-Flora Von Schlesien. Im
Namen Der Schlesischen Gesellschaft fiir

March 1991

vaterlandische Cultur herausgegeben von
Prof. Dr. Ferdinand Cohn, Secretair der
Botanischen Section, Breslau 3(2):364. 1894.
=Sphaeria aliriae L. Fuckel, Symbolae
Mycologicae, p. 135. 1870, sub. Pleospora
maculans. On stems of Alliaria officinalis
Andrz ex Bieb., Cruciferae. Poland.
alliariae (B. Auerswald) H. Rehm, Ascomyceten,
Fascicle 14, No. 686. Anno 1882; Hedwigia,
Dresden 22:56. 1883. =Sphaeria alliariae B.
Auerswald in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi

Mycologici Continuatio, Edition 3, Century 3,

No. 261. Anno 1860. On dry stems of
Sisymbrium alliaria (L.) Scop., Cruciferae.
Germany.

alliariae (B. Auerswald) G. Linhart, Fungi Hungar-
ici Exsiccati [Magyarorszag Gombai],

Century 2, No. 164. Anno 1883. Nom. illegit.

Art. 64.1. =Sphaeria alliariae B. Auerswald

in G.L. Rabenhorst, Fungi Europaei Exsiccati,

Klotzschii Herbarii vivi Mycologici Continu-
atio, Edition 3, Century 3, No. 261. Anno
1860. On dry stems of Brassica campestris
L., Cruciferae. Germany, Hungary.

allorgei A.L. Guyot, Revue de Mycologie, Paris
14:69. 1949. On dry stems of /satis tinctoria
L., Cruciferae. France.

almeidae M. de Sousa da Camara, Revista
Agronomica, Lisbon, No. 1:11—12. 1928.
{From reprint.] On leaves of Cinnamomum
zeylanicum Blume, Lauraceae. Portugal.

almeidana M. de Sousa da Camara, Boletim de
Agricultura, Lisboa 2(1):14. 1936. [Pagina-
tion on reprint, p. 21.) On branchlets of
Pinecenectria sp., Agavaceae. Portugal.

aloes M.R. Sousa de Dias & M. de Sousa da
Camara, Agronomia Lusitana, Sacavém 16:8.
1954. On leaves of Aloe arborescens Miller,
Liliaceae. Portugal.

alopecuri N.N. Lavrov, Trudy Tomskogo Gosu-
darstvennogo Universiteta. Tomsk. Series
Biologicheskii 110:76. 1951. On dead leaves
of Alopecurus borealis Trin., Gramineae.
U.S.S.R.

alpiniae A. Maublanc, Bulletin de la Société
Mycologique de France, Paris 21:89. 1905S.
On leaves of Alpinia speciosa K. Sch.,
Zingiberaceae. Brazil.

altaica A. Nannizzi, Atti dell’ Accademia dei
Fisiocritici di Siena, Series 10, 3:961. 1928.
On dried sheaths of leaves on Polygonum
bistorta L., Polygonaceae. Yugoslavia.

alvariensis K. Starbick, Bikang till K. Svenska
Vetenskapsakademiens Handlingar, Stock-
holm, Series 3, 15(2):28. 1889. [Raised to
specific rank in legend of Figure 4.] =Lepro-
sphaeria vagabunda P.A. Saccardo subsp.
alvariensis K. Starbiick.

Nomenclator of Leptosphaeria 201

ambiens H. Rehm, Philippine Journal of Science,
Manila, Section C, 8:257. 1913. On dead
branches of unknown host, unknown family.
Philippines.

ammophilae (W.G. Lasch) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:236. 1863.
=Sphaeria ammophilae W.G. Lasch in G.L.
Rabenhorst, Klotzschii Herbarium Vivum
Mycologicum Sistens Fungorum Per Totam
Germaniam Cresentium Collectionem
Perfectam, Dresden, Edition I, Century 14,
No. 1340. Anno 1850; Flora, Jena und
Regensburg 8:282. 1850; Botanische Zeitung,
Berlin & Leipzig 8:439. 1850.
=Phaeosphaeria ammophilae (W.G. Lasch) J.
Kohlmeyer & E. Kohlmeyer, Icones Fun-
gorum Maris Lief. 3, Table 55. 1965.
=Amarenomyces ammophilae (W.G. Lasch)
O. Eriksson, Nordic Journal of Botany,
Copenhagen [Opera Botanica] 60:124. 1981.
On leaves of Ammophila arenaria (L.) Link,
Elytrigia juncea (L.) Nevski, Gramineae.
Denmark, Finland, Germany, Netherlands,
U:S'S:R:

ammophilae H. Rehm ex H. Rehm, Ascomyceten,
Fascicle 14, No. 691. Anno 1882. Nom. nud.
32.1 [as (Lasch) Cesati & de Notaris];
Hedwigia, Dresden 22:57. 1883. Non
Leptosphaeria ammophilae (W.G. Lasch) V.
Cesati & G. de Notaris. On dry leaves of
Ammophila arenaria (L.) Link, Gramineae.
Germany.

ammophilae H. Rehm, falsely so cited in A.N.
Berlese and P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
139. 1886; P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:790. 1891. =Leptosphaeria
ammophilae (W.G. Lasch) V. Cesati & G. de
Notaris.

ammothamni E.N. Koschkelova in E.N. Kosch-
kelova, I.P. Frolov, and Z. Dzhuraeva,
Mikoflora Badkhyza, Karabilya i Yuzhnoi
Chasti Murgabskogo Oazisa (Mikromitsety)
[The Mycoflora of Badkhyz, Karabil and the
Southern Part of the Murgab Oasis (Micromy-
cetes)], p. 87. 1970. Nom. nud. Art. 32.1. On
dry branches of Ammothamnus lehmannii
Bunge, Leguminosae. U.S.S.R. (Badkhyza,
Karabilia).

ammothamni E.N. Koschkelova, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 7:176—177.
1970. On dry branches of Ammothamnus
lehmannii Bunge, Leguminosae. U.S.S.R.

amorphae E. Mitroshina, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-

202

rum U.R.S.S., Petropolis 6:82. 1949. On dead
branches of Amorpha fruticosa L., Legumino-
sae. U.S.S.R.

ampelina M. Curzi & M. Barbaini, Atti dell’ Istituto
Botanico della Universita e Laboratorio
Crittogamica di Pavia, Milano, Series 3,
3:160. 1927. On dry vines of Vitis vinifera L.,
Vitaceae. Italy.

amphibola P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:322. 1875. =Leptosphaeria
culmifraga P.A. Saccardo non (E.M. Fries) V.
Cesati & G. de Notaris. On stems of Sorghum
vulgare Pers., Gramineae. Italy.

amphiloga F. Petrak in H. Sydow and F. Petrak,
Annales Mycologici, Berlin 29:202. 1931. On
dead culms of Bambusa sp., Gramineae.
Philippines.

anacycli R. Gonzalez Fragoso, Asociacion Espanola
Para el Progresso de las Ciencias, Congreso
de Coimbra 1925:11—12. 1925. Nom. inval.
Art. 34.1. On dry stems of Anacylus radiatus
Loisel., Compositae. Portugal.

anarithma (M.J. Berkeley & C.E. Broome?) P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Ilustrans
1:395. 1878; Fungi Italici autographice
delineati (additis nonnullis extra-italicis
asterisco notatis), Patavii, Table 400. 1878.
=Sphaeria (Foliicolae) anarithma M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
3, 3:374. 1859. =Sphaerella anarithma (M.J.
Berkeley & C.E. Broome) M.C. Cooke,
Handbook of British Fungi, p. 920. 1871.
=Sphaerella anarithma P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 8:175. 1876
{as (B. & Br.) Cooke]. =Metasphaeria
anarithma (M.J. Berkeley & C.E. Broome)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:175. 1883. This new combination or new
species is based on material collected on dried
leaves of Aira cespitosa L. from Montello .
(Treviso), September 1873, and may be
identical to the species described by M.J.
Berkeley and C.E. Broome, also on Aira
cespitosa from Batheaston, March 1853, and
Bowood, October 1853. On dried leaves of
Aira cespitosa L., Gramineae. Great Britain,
Italy.

anarithmoides P.A. Saccardo & C.L. Spegazzini in
P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:395. 1878. =Metasphaeria
anarithmoides (P.A. Saccardo & C.L.
Spegazzini) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:175. 1883. On decaying leaves of
Poa sp.?, Gramineae. Italy.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

anarrhini R. Gonzalez Fragoso, Boletim da
Sociedade Broteriana, Coimbra, Series 2.
2:22—23. 1923. On dry stems of Anarrhinum
bellidifolium (L.) Willd., Scrophulariaceae.
Portugal.

anceps P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:120. 1878. On not yet dead
branches of Ribes nigrum L., Saxifragaceae.
Germany.

andrijevicensis F. Bubak, Bulletin de |’ Herbier
Boissier, Genéve and Bale, Series 2,
6:400-401. 1906. On old stems of Ranuncu-
lus villarsii DC., Ranunculaceae. Yugoslavia.

andromedae (B. Auerswald) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:49-SO. 1883.
=Sphaerella andromedae B. Auerswald in W.
Gonnermann and G.L. Rabenhorst, Mycologia
Europaea, Abbildungen Sammtlicher Pilze
Europa’s, Hefte 5 und 6, Synopsis Pyrenomy-
cetum Europaeorum, p. 12. 1869 [as Spae-
rella}. On dry leaves of Andromeda tetragona
L., Ericaceae. Canada, Norway.

anemones L. Hollés, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 4:334.
1906. [Anamorph: Rhabdospora anemones L.
Hollos.] On dry petioles of Anemone
sylvestris L., Ranunculaceae. Hungary.

anisomeres L.E. Wehmeyer, Canadian Journal of
Research, Ottawa 20(Section C):582. 1942.
On unknown substrate of Agropyron repens
(L.) Beauv., Gramineae. Canada (Nova
Scotia).

anomala J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 3:117. 1887.
=Didymosphaeria anomala (J.B. Ellis & B.M.
Everhart) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:730. 1891. =Microthelia anomala
(J.B. Ellis & B.M. Everhart) O. Kuntze,
Revisio Generum Plantarum 3(3):498. 1898.
=Wertsteinina anomala (J.B. Ellis & B.M.
Everhart) M.E. Barr, Contributions from the
University of Michigan Herbarium, Ann
Arbor 9:548. 1972. On dead herbaceous
stems of unknown host, unknown family.
U.S.A. (Utah).

antarctica C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cordoba 11:218.
1888; Fungi Fuegani, No. 228. Anno 1888.
On leaves of unknown host, unknown family.
Argentina (Tierra del Fuego).

anthelmintica (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:21. 1883.
=Sphaeria (Caulicolae) anthelmintica M.C.
Cooke, Grevillea, London 7:54. 1878. On
stems of Chenopodium anthelminticum Bert.
ex Steud., Chenopodiaceae. U.S.A.

March 1991

antherici L. Holl6s, Matematikai és Természettu-
domanyi Kézlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:31. 1926. On dry
stems of Anthericum ramosum L., Liliaceae.
Hungary.

anthophila P.A. Saccardo & C.L. Spegazzini,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 1:396.
1878. On lower portion of fallen, rotting
capitulum of Carlina acaulis L., Compositae.
Italy.

anthostomella C.L. Spegazzini, Anales del Museo
Nacional de Historia Natural de Buenos Aires
6:281. 1898; Fungi Argentini Novi vel Critici,
No. 624. Anno 1898. On fallen decayed
culms of Panicum grumosum Nees, Gra—
mineae. Argentina.

anthostomoides (H. Rehm) H. Rehm, Bericht des
Naturwissenschaftlichen Vereins fiir Sch-
waben und Neuberg, Augsburg 26:84. 1881.
=Massaria anthostomoides H. Rehm,
Ascomyceten, Fascicle 7, No. 339. Anno
1876. =Montagnula anthostomoides (H.
Rehm) A. Leuchtmann, Sydowia. Annales
Mycologici, Horn, N.O. 37:175. 1984. On
decayed stems of Aconitum napellus L.,
Ranunculaceae. Italy.

aparines (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:21. 1883. =Ple-
ospora aparines L. Fuckel, Symbolae
Mycologicae, p. 136. 1870 [as aparinae]. On
dry stems of Galium aparine L., Rubiaceae.
Austria.

apios M. Miura ex I. Hino & K. Katumoto, Transac-
tions of the Mycological Society of Japan
{Nihon Kin Gakkai Kaiho] 7:47. 1966 [as
apiosis|. On living leaves of Apios fortunei
Maxim., Leguminosae. Japan.

apios-fortunei (M. Miura] ex I. Hino & K. Katu-
moto, Transactions of the Mycological
Society of Japan [Nihon Kin Gakkai Kaiho]
7:46-47. 1966 [as apiosis-fortunei (M. Miura)
I. Hino & K. Katumoto comb. nov.]. =Guig-
nardia apios-fortunii M. Miura, Report of the
Akita Prefectural Agricultural Experiment
Station 8:9. 1957. On living leaves of Apios
fortunei Maxim., Leguminosae. Japan.

apocalypta (H. Rehm) ex H.G. Winter, Die Pilze
Deutschlands, Oesterreichs und der Schweiz,
Abt. IL. (Ascomyceten) in G.L. Rabenhorst,
Kryptogamen-Flora von Deutschland,
Oesterreich und der Schweiz 1(2):442. 1885
[as Leptosphaeria? apocalypta|. =Xeno-
sphaeria apocalypta H. Rehm, Ascomyceten,
Fascicle 7, No. 346. Anno 1876. Nom. nud.
Art. 32.1. On thallus of Stereocaulon alpinum
Laur., Lichenes (Stereocaulaceae). Italy.

apogon P.A. Saccardo & C.L. Spegazzini in P.A.
Saccardo, Michelia Commentarium Mycol-

Nomenclator of Leptosphaeria

203

ogicum Fungos in Primis Italicos Illustrans
1:398. 1878. =Leptosphaeria culmorum P.A.
Saccardo & C.L. Spegazzini forma apogon
(P.A. Saccardo & C.L. Spegazzini) H. Rehm.
=Heptameria apogon (P.A. Saccardo & C.L.
Spegazzini) M.C. Cooke, Grevillea, London
18:32. 1889 [as apogon Sacc.]. =Lepto-
sphaeria eustoma (E.M. Fries:E.M. Fries)
P.A. Saccardo forma apogon (P.A. Saccardo
& C.L. Spegazzini) A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:56. 1892. On culms of Scirpus holo-
schoenus L., Cyperaceae. Italy.

appendiculata R. Pirotta, Atti dell’ Istituto Botanico
della Universita e Laboratorio Crittogamica di
Pavia, Milano 2 & 3:161—162. 1879.
=Leptosphaeria vitis §. Schulzer v.
Miiggenburg. Nom. illegit. Art. 63.1. [A
superfluous name change. ]

aquatica §.T. Tilak & R.L. Kulkarni, Beihefte zur
Nova Hedwigia Weinheim 47:456—-457. 1974.
On living leaves of Typha angustata Bory &
Chaub., Typhaceae. India.

aquilana D., Saccardo ex D. Saccardo in P.A.
Saccardo, Mycotheca Italica, Sistens Fungos
Venetos Exsiccatos, Century 15, No. 1485.
Anno 1881. Nom. nud. Art. 32.1; Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 17:724-725. 1905.
=Nodulosphaeria aquilana (D. Saccardo) L.
Holm, Symbolae Botanicae Upsalienses
14(3):83. 1957. On dry stems of unknown
host, Scrophulariaceae. Italy.

aquilegiae (A.N. Berlese & G. Bresadola) F.
Hazslinszky, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 25(2):145. 1892.
=Metasphaeria sepincola (E.M. Fries:E.M.
Fries) P.A. Saccardo var. aquilegiae A.N.
Berlese & G. Bresadola, Micromycetes
Tridentini Contribuzione Allo Studio dei
Funghi Microscopici del Trentino Per Cura
Del Dr. A.N. Berlese e dell’ Ab. G. Bresadola,
p. 39. 1889. =Metasphaeria aquilegiae A.N.
Berlese & G. Bresadola, Revue Mycologique,
Toulouse 12:185. 1890. On stems of
Aquilegia vulgaris L., Ranunculaceae.
Germany, Hungary.

aquilina (B. Auerswald) G. Passerini, Atti della
Societa Crittogamologica Italiana, Milano,
Series 2, 2:47. 1879. =Sphaerella aquilina B.
Auerswald in W. Gonnermann and G.L.
Rabenhorst, Mycologia Europaea, Ab-
bildungen Sammtlicher Pilze Europa’s, Hefte
5 und 6, Synopsis Pyrenomycetum Eu-
ropaeorum, p. 20. 1869. On dry frond of
Preris aquilina L., Pteridaceae. Italy.

arabidis A. Allescher, Bericht der Bayerischen
Botanischen Gesellschaft zur Erforschung der

204 Illinois Natural History Survey Bulletin

Heimischen Flora, Miinchen 5:14. 1897. On
dead stems of Arabis alpina L., Cruciferae.
Germany.

arbuti A. Clotilde dos Santos & M. de Sousa da
Camara, Agronomia Lusitana, Sacavém
17:139-140. 1955. On branches of Arbutus
unedo L., Ericaceae. Portugal.

arctalaskana Y. Kobayasi, Annual Report of the
Institute for Fermentation Osaka 3:32. 1967.
On bract of inflorescence on Lycopodium
selago L. var. appressum Desv., Lycopodi-
aceae. U.S.A.

arecae G. Mariani, Atti della Societa Italiana di
Scienze Naturali e del Museo Civile di Storia
Naturale, Milano 50:165-166. 1911. On
wilted and dead leaves of Areca sapida
Soland. ex Hook. f., Palmae. Portugal.

arenaria A.L. Guyot, Revue de Mycologie, Paris
14:69, 71. 1949. Nom. illegit. Art. 64.1.
=Phaeosphaeria arenaria (A.L. Guyot) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1538. 1989. On dry leaves; on
sheaths and culms of Festuca arenaria
Osbeck, Phleum arenarium L., Gramineae.
France.

arenaria (E.C. Bommer, M.H. Rousseau & P.A.
Saccardo) J. Lind, Danish Fungi as Repre-
sented in the Herbarium of E. Rostrup,
Nordisk Forlag, Copenhagen, p. 220. 1913 [as
arenariae]. =Metasphaeria arenaria E.C.
Bommer, M.H. Rousseau & P.A. Saccardo,
Annales Mycologici, Berlin 3:509. 1906. On
dead leaves of Elymus arenarius L., Hordeum
arenarium Ascherson, Gramineae. Belgium,
Denmark.

argentina C.L. Spegazzini, Anales de la Sociedad
Cientifica Argentina, Buenos Aires 9:183.
1880. =Heptameria argentina (C.L. Spegazz-
ini) M.C. Cooke, Grevillea, London 18:29.
1889. On fallen stems of Oenothera
longiflora L., Onagraceae. Argentina.

argentinensis (C.L. Spegazzini) F. Petrak in F.
Petrak and H. Sydow, Annales Mycologici,
Berlin 33:171. 1935. =Venturiella argenti-
nensis C.L. Spegazzini, Anales del Museo
Nacional de Historia Natural de Buenos Aires
19(Series 3, 12), p. 379. 1909; Mycetes
Argentinenses, Series IV, No. 619. 1909.
=Neoventuria argentinensis (C.L. Spegazzini)
H. Sydow & P. Sydow, Annales Mycologici,
Berlin 17:44. 1910. On wood of Pircunia
dioica Mogq. (=Phytolacca), Phytolaccaceae.
Argentina.

arnoldii H. Rehm, Osterreichische Botanische
Zeitschrift, Wien 54:84. 1904 [as arnoldi].
On thallus of Peltigera malacea (Ach.)
Funck., Lichenes (Peltigeraceae). Italy.

arrhenatheri F. Hazslinszky, Matematikai és
Természettudomanyi Kézlemenyek Von-
atkozolag a Hazai Viszonyokra, Budapest

Vol. 34 Art. 3

25(2):132. 1892. On culms of Arrhenatherum
avenaceum Beauv., Gramineae. Hungary.

arrhenatheri F. Hazslinszky var. italica C. Massa in
T. Ferraris and C. Massa, Annales Mycol-
ogici, Berlin 10:287. 1912. On leaves of
Arrhenatherum elatius (L.) Beauv. ex J. Presl.
& C. Presl., Gramineae. Italy.

artemisiae (L. Fuckel) B. Auerswald in G.L.
Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio Edita Nova, Series 2, Century 18, No.
1725. Anno 1874; Hedwigia, Dresden
17:46-47. 1878 [as in G. Niessl v.
Mayendorf]. =Sphaeria artemisiae L. Fuckel,
Fungi Rhenani Exsiccati A Leopoldo Fuckel
Collecti, Fascicle 9, No. 896. Anno 1864.
=Pleospora helminthospora L. Fuckel,
Symbolae Mycologica, p. 138. 1870;
Symbolae Mycologicae Zweiter Nachtrag, p.
24. 1873. [Erroneously cited in both these
works as (Ces.) Fuckel. Not based on
Sphaeria helminthospora V. Cesati.] On dry
stems of Artemisia campestris L., Composi-
tae. Austria, Germany.

arthrophyma (C.E. Fairman) L. Holm, Svensk
Botanisk Tidskrift, Stockholm 62:228—229.
1962. =Gibberidia arthrophyma C.E.
Fairman, Mycologia, Lancaster, Pennsylvania
10:246—247. 1938. On old stems of Chryso-
thamnus graveolens (Nutt.) Greene, Composi-
tae. U.S.A.

arunci §.M. Zeller, Mycologia, Lancaster, Pennsyl-
vania 19:134—-135. 1927. On dead stems of
Aruncus silvestris Kostel. [=Aruncus dioicus
(Walter) Fernald], Rosaceae. U.S.A.
(Oregon).

arundinacea (M.J. Sowerby:E.M. Fries) P.A.
Saccardo, Atti dell’ Accademia Scientifica
Veneto-Trentino-Istriana, Padova 2(2):155.
1873. =Sphaeria arundinacea M.J.
Sowerby:E.M. Fries, Coloured Figures of
English Fungi or Mushrooms, Table 336.
1803; E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:429. 1823. =Pleospora
arundinacea (M.J. Sowerby:E.M. Fries) L.
Fuckel, Symbolae Mycologicae, p. 137. 1870.
=Melogramma (?) arundinacea (M.J.
Sowerby:E.M. Fries) G. Niessl v. Mayendorf
in G.L. Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio, Edition 3, Century 19, No. 1840. Anno
1874; Hedwigia, Dresden 13:185. 1874.
=Sphaeria striaeformis Auctores var.
arundinis J.B. Albertini & L.D. v. Schweinitz,
Conspectus Fungorum in Lusatiae Superioris
Agro Niskiensi Crescentium e Methodo
Persoonia, p. 15. 1805. [This variety ts
considered a synonym by E.M. Fries.]
=Phaeosphaeria arundinacea (M.J. Sowerby)
A. Hedjaroude, Sydowia. Annales Mycol-

March 199]

ogici, Horn, N.O. 22:78. (1968) 1969.
=Massarina arundinaceae (M.J.
Sowerby:E.M. Fries) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:179. 1984. On fallen culms of Phragmites
communis Trin., Gramineae. France, Great
Britain, Italy.

arundinacea (M.J. Sowerby:E.M. Fries) H. Rehm,
Bericht des Naturwissenschaftlichen Vereins
fiir Schwaben und Neuberg, Augsburg 26:50.
1881. Nom. illegit. Art. 64.1. [Combination
previously made as Leptosphaeria
arundinacea (M.J. Sowerby:E.M. Fries) P.A.
Saccardo.] On fallen culms of Phragmites
communis Trin., Gramineae. Great Britain,
Italy.

arundinacea (M.J. Sowerby:E.M. Fries) P.A.
Saccardo var. godini (J. Desmaziéres) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:63. 1883. =Sphaeria godini J. Desmaziéres,
Plantes Cryptogames Du Nord De La France,
Edition 1, Fascicle 9, No. 439. Anno 1829;
Annales des Sciences Naturelles, Paris,
Botanique, Series 3, 5:49. 1846. On culms;
on stubble of Phragmites communis Trin.,
Phragmites sp., Gramineae. France.

arundinacea (M.J. Sowerby:E.M. Fries) P.A.
Saccardo var. godini (J. Desmaziéres) P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
2:65. 1880. =Sphaeria godini J. Desmaziéres,
Plantes Cryptogames Du Nord De La France,
Edition 1, Fascicle 9, No. 439. Anno 1829;
Annales des Sciences Naturelles, Paris,
Botanique, Series 3, 5:49. 1846. =Lepto-
sphaeria godini (J. Desmaziéres) B. Auers-
wald. On stubble of Phragmites communis
Trin., Gramineae. France.

arvensis C.L. Spegazzini in P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 1:459. 1879. On
dead, rotting stems of Equisetum arvense L.,
Equisetaceae. Italy.

asclepiadis R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2693-2694. 1984. On
stems of Asclepias sp., Asclepiadaceae.
Canada, U.S.A.

asparagi C.H. Peck, Bulletin of the New York State
Museum, Albany 219-220:68. (1919) 1920.
On dead stems of Asparagus officinalis L.,
Liliaceae. U.S.A. (New York).

asparagi G. Passerini, Rendiconti della Sedute della
R. Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:90. 1887. =Leptosphaeria
passeriniana P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 9:785. 1891. On dry
stems of Asparagus officinalis L., Liliaceae.
Italy.

Nomenclator of Leptosphaeria 205

asparagi C.H. Peck, Report. New York State
Museum of Natural History, Albany 40:70.
(1886) 1887. On dead stems of Asparagus
officinalis L., Liliaceae. U.S.A.

asparagina P.A. Karsten, Hedwigia, Dresden 23:1.
1884; Acta Societatis pro Fauna et Flora
Fennica, Helsingforsiae 2(6):55. 1885. On
stems of Asparagus officinalis L., Liliaceae.
Finland.

asperellae K. Hara, By6chu-gai Zasschi (Journal of
Plant Protection), Tokyo 5:883-884. 1918.
On leaves of Asperella japonica Hack.,
Gramineae. Japan.

aspidistrae F. Tassi, Bolletino del R. Orto Botanico,
Siena 3:14—-15. 1900. On leaves of Aspidistra
elatior Blume, Liliaceae. China.

asplenti (G.L. Rabenhorst) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:82. 1883. =Sphae-
rella asplenii G.L. Rabenhorst in G. Niessl v.
Mayendorf, Verhandlungen des Naturforsch-
enden Vereins in Briinn 3:178. 1865;
Vorarbeiten zu einer Kryptogamenflora von
Mahren und Oesterr II. Pilze und Myxomy-
ceten, p. 121. 1865. On dry fronds of
Asplenium septentrionale (L.) Hoffm.,
Aspleniaceae. Germany.

associata H. Rehm, Annales Mycologici, Berlin
10:356-357. 1912; Ascomyceten, Fascicle 50,
No. 1994. Anno 1912. =Phaeosphaeria
associata (H. Rehm) O. Eriksson, Arkiv for
Botanik, Uppsala, Stockholm, Series 2, 6:412.
1967. Parasitic on Epichloe typhina
(Pers.:Fr.) Tul., Fungi (Clavicipitaceae).
Canada, Sweden, U.S.A.

astericola J.B. Ellis & B.M. Everhart, Fungi
Columbiani, Century 16, No. 1537. Anno
1901. Nom. nud. Art. 32.1. On dead stems of
Aster multiflorus Ait. (=Aster ericoides L.),
Compositae. U.S.A.

astericola J.B. Ellis & B.M. Everhart ex J.B. Ellis &
B.M. Everhart, Journal of Mycology,
Columbus, Ohio 8:17. 1902. On dead stems
of Aster multiflorus Ait. (=Aster ericoides L.),
Compositae. U.S.A.

asteris J.H. Miller & G. Burton, Mycologia,
Lancaster, Pennsylvania 34:34. 1942. On
dead stems of Aster sagittifolius Wedem. ex
Willd., Compositae. U.S.A.

atraphaxidis M.P. Vasyagina in M.P. Vasyagina,
Z.M. Byzova, and M.A. Tartenova, Flora
Sporovykh Rastenii Kazakhstana 12
Sumchatye Griby 2. Lokuloaskomitsely
(Loculoascomycetes) Alma Alta: “Nauka”
AH Kazakhskoi SSR, p. 125. 1987. On
unknown substrate of Atraphaxis musch-
ketovii Krasnov, Polygonaceae. U.S.S.R.

atriplicis A.L. Lobik, Material¥ po Floristicheskim i
Faunisticheskim Obsledovaniyam Terskogo
Okruga [Data from Investigations on the Flora
and Fauna of the Ter Region], p. 23. 1928.

206

On unknown substrate of Atriplex verrucifera
Bieb., Chenopodiaceae. U.S.S.R.

atropurpurea F., Petrak, Annales Mycologici, Berlin
25:286—287. 1927. On stems of Urtica dioica
L., Urticaceae. Germany.

aucta G. Niessl v. Mayendorf in G.L. Rabenhorst,
Fungi Europaei Exsiccati, Klotzschii Herbarii
vivi Mycologici Continuatio, Series 2,
Century 23, No. 2240. Anno 1876; Hedwigia,
Dresden 16:118. 1877. =Nodulosphaeria
aucta (G. Niessl v. Mayendorf ) L. Holm,
Svenska Botanisk Tidskrift, Stockholm 55:73.
1961. On stems of Clematis recta L.,
Ranunculaceae. Czechoslovakia.

auerswaldii P.A. Saccardo & J.B. Traverso in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
19:1096. 1910. Nom. illegit. Art. 63.1. Name
erroneously changed to avoid a tautonym.
=Sphaerella leptosphaerioides B. Auerswald
in W. Gonnermann and G.L. Rabenhorst,
Mycologia Europaea, Abbildungen Sammtli-
cher Pilze Europa’s, Hefte 5 und 6, Synopsis
Pyrenomycetum Europaeorum, p. 13. 1869.
On either side of dry leaves and sepals of
Arenaria ciliata L., Caryophyllaceae. Europe.

aulica (M.C. Cooke & J.B. Ellis) J.B. Ellis in N.L.
Britton, Catalogue of Plants Found in New
Jersey. Geological Survey of New Jersey,
Final Report of the State Geologist, Trenton
2(1):525. 1889. =Sphaeria (Obtecta) aulica
M.C. Cooke & J.B. Ellis, Grevillea, London
6:95. 1878. On dead stems of Lonicera sp.,
Solidago sp., Caprifoliaceae, Compositae.
U.S.A. (New Jersey).

australiensis (A.B. Cribb & J.W. Cribb) G. Hughes,
Syesis, British Columbia Provincial Museum,
Victoria, British Columbia 2:132. 1969.
=Metasphaeria australiensis A.B. Cribb &
J.W. Cribb, Papers from the Department of
Botany, University of Queensland, Brisbane
3:79. 1955. On roots of Avicennia marina
(Forsk.) Vierh. var. resiniferae (Forst.) Bakh.,
Avicenniaceae. Australia, Canada.

australis (L. Crié) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:54. 1883. =Depazea australis L.
Crié, Annales des Sciences Naturelles, Paris,
Botanique, Series 6, 7:46. 1878. On living
leaves of Eustephia sp., Amaryllidaceae.
Australia, New Caledonia.

austro-americana C.L. Spegazzini, Annales de la
Sociedad Cientifica Argentina, Buenos Aires
10:20. 1880; Fungi Argentini Pugillus 2, No.
82. Anno 1880. On living or wilting leaves of
Plantago macrostachys Decne., Plantag-
inaceae. Argentina.

avellanae J.H. Fabre, Annales des Sciences
Naturelles, Paris, Botanique, Series 6, 9:89.
1878. On dead branches of Corylus avellana
L., Betulaceae. France.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

avenae B. Auerswald in W. Gonnermann and G.L.
Rabenhorst, Mycologia Europaea, Abbildugen
Sammtlicher Pilze Europa’s, Hefte 5 und 6,
Synopsis Pyrenomycetum Europaeorum,
Table 12, Figure 165. 1869. Nom. nud. Art.
32.1. =Metasphaeria avenae (B. Auerswald)
ex P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:176. 1883 [as (Auersw.) Sacc.]. On leaves
of Avena sp., Gramineae. Germany.

avenaria G.F. Weber, Phytopathology, Lancaster,
Pennsylvania 12:454-455. 1922.
=Phaeosphaeria avenaria (G.F. Weber) O.
Eriksson, Arkiv for Botanik, Uppsala,
Stockholm, Series 2, 6:408. 1967. On leaves
of Agropyron repens (L.) Beauy., Bromus
inermis Leysser, Poa pratensis L., Gramineae.
U.S.A.

avenaria G.F. Weber form sp. triticea T. Johnson,
Canadian Journal of Botany, Ottawa 25:
262-263. 1947. =Phaeosphaeria avenaria
(G.F. Weber) O. Eriksson form sp. triticea (T.
Johnson) R.A. Shoemaker, Canadian Journal
of Botany, Ottawa 67:1522. 1989. On leaves
and sheaths of Triticum durum Desf., Triticum
vulgare Vill., Gramineae. Canada.

avicenniae J. Kohlmeyer & E. Kohlmeyer, Nova
Hedwigia. Zeitschrift fiir Kryptogra-
menkunde, Weinheim 9:98—99. 1965. On
bark of living pneumatophores of Avicennia
nitida Jacq., Avicenniaceae. U.S.A.

azaleae L. Hollés, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 6:529.
1908. On dead branches of Azalea sp.
(=Rhododendron sp.), Ericaceae. Hungary.

bacillifera (P.A. Karsten) A. Munk, Dansk Botanisk
Arkiv, Kjobenhavn 17(1):378. 1957. Nom.
inval. Art. 33.2. =Sphaerella bacillifera P.A.
Karsten, Hedwigia, Dresden 22:179. 1883.
On dead leaves of Scheuchzeria palustris L.,
Scheuchzeriaceae. Denmark.

baeomycearia (W. Lindsay) P.A. Saccardo & A.
Trotter in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 22:235. 1913. =Microthelia
baeomycearia W.L. Lindsay, Transactions of
the Royal Society of Edinburgh 25:541—542.
1869; Proceedings of the Royal Society of
Edinburgh 6:535. 1869. Nomen. On thallus
of Baeomyces rufus (Huds.) Rebent., Lichenes
(Baeomycetaceae). Great Britain.

baggei (B. Auerswald ex G. Niessl v. Mayendorf)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:35. 1883. =Cryptospora baggei (B.
Auerswald) ex G. Niessl v. Mayendorf. Ver-
handlungen des Naturforschenden Vereins in
Briinn 10:200. 1872. =Sphaeria baggei B.
Auerswald, Tauschverein (unpublished
manuscript?). Nom. nud. Art. 32.1c? On
branches of Salix sp., Salicaceae. Austria,
Czechoslovakia, Germany.

March 1991

balcarica Z.D. Savintzeva, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 8:109—110.
1971. On dead stems of Trifolium
trichocephalum Bieb., Leguminosae.
U.S.S.R.

baldingerae F. Fautrey & J.B. Lambotte in F.
Fautrey, Revue Mycologique, Toulouse
19:53. 1897. =Phaeosphaeria baldingerae (F.
Fautrey & J.B. Lambotte) A. Hedjaroude,
Sydowia. Annales Mycologici, Horn, N.O.
22:87. (1968) 1969. On dry culms of
Baldingera arundinacea (L.) Dumort.,
Verbenaceae. France.

baldratiana P. Baccarini, Annali di Botanica, Roma
4:274. 1906. On leaves of Sansevieria
ehrenbergii Schweinf. ex Baker, Agavaceae.
Ethiopia.

ballotae J. Politis, Akademia Athénon, Pragmateiai,
Athens 1935:16. 1935. On dried stems of
Ballota acetabulosa (L.) Bentham, Labiatae.
Greece.

bambusae (1. Miyake & K. Hara) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo
24:995-996. 1928. =Phaeosphaeria
bambusae K. Miyake & K. Hara, Botanical
Magazine, Tokyo 24:340. 1910. =Lepto-
sphaerella bambusae (1. Miyake & K. Hara) I.
Miyake & K. Hara, Botanical Magazine,
Tokyo 24:(249). 1913. =Trematosphaerella
bambusae (K. Miyake & K. Hara) I. Hino &
K. Katumoto, Icones Fungorum Bambusicolo-
rum Japonicorum, p. 160. 1961. Non
Trematosphaerella bambusae (K. Miyake &
K. Hara) F.L. Tai, Sylloge Fungorum
Sinicorum, Science Press, Academia Sinica,
Peking, p. 330. 1979. Nom. illegit. Art. 64.1.
On leaves of Phyllostachys bambuoides
Siebold & Zucc., Pleioblastus hindsii (Munro)
Nakai, Pleioblastus simoni (Carriere) Nakai,
Pseudosasa japonica (Siebold & Zucc. ex
Steudel) Makino, Sasa paniculata (F.
Schmidt) Makino [=S. senanensis (Fr. & Sav.)
Rehd.], Sasaella ramosa (Makino) Makino,
Semiarundinaria tatebeana Muroi, Grami-
neae. Japan.

bambusae L. Rolland, Bulletin de la Société
Mycologique de France, Paris 12:4. 1896 [as
banbusae} . On cortex (culm) of Bambusa sp.,
Gramineae. France.

bambusicola S.C. Teng & S. Ou, Sinensia. Special
Bulletin of the Metropolitan Museum of
Natural History, Nanking 7:500. 1936. On
decaying culms of Bambusa sp., Gramineae.
China (Chekiang).

bardanae (K.F.W. Wallroth) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:34. 1883.
=Sphaeria bardanae K.F.W. Wallroth, Flora
Cryptogamica Germaniae 2:805. 1833. On

Nomenclator of Leptosphaeria 207

dried stems of Bardana sp., Compositae.
Germany.

barriae R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2694. 1984. On stems of
Lupinus sp., Leguminosae. Canada.

basalduai C.L. Spegazzini, Anales del Museo
Nacional de Historia Natural de Buenos Aires
8(Series 3, 1):71. 1902; Mycetes Argenti-
nenses, Series II, No. 133. Anno 1902. On
decaying, fallen branches of Lippia seriphioi-
des A. Gray, Verbenaceae. Argentina.

bataticola M. Chochrjakov & M.S. Dunin in MLS.
Dunin and E.D. Yakimovick, Bolezni Batata i
Mery Bor’by s Nimi, pp. 89-90. 1934. On
leaves of Ipomoea batatas (L.) Lam.,
Convolvulaceae. U.S.S.R.?

batumensis W. Siemaszko, Acta Societatis Botani-
corum Poloniae, Warszawa 1:21. 1923. On
living leaves of Phoenix canariensis Hort. ex
Chabaud, Palmae. U.S.S.R.

baumii J.A. v. Arx & E. Miiller, Studies in Mycol-
ogy, Baarn, Netherlands 9:76. 1975.
=Baumiella caespitosa P.C. Hennings in H.
Baum, Kunene-Sambesi Expedition, Berlin, p.
165. 1903. Non Leptosphaeria caespitosa G.
Niessl v. Mayendorf. On leaves of Monotes
dasyanthus Gilg., Dipterocarpaceae. Angola.

beaumontii (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:87. 1883. =Sphaeria beaumontii M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:145. 1876. On grass
culms of unknown host of Gramineae,
Gramineae. U.S.A.

bella G. Passerini, Rendiconti della Sedute della R.
Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:90. 1887. On branchlets of
Chondrilla juncea L., Compositae. Italy.

bellynckii (G.D. Westendorp) B. Auerswald,
General-Doubl. Verzeichn-tauschverein, p. 4.
1870. Non vidi. =Sphaeria bellynckii G.D.
Westendorp, Bulletins de L’ Académie Royale
des Sciences, des Lettres et des Beaux-Arts de
Belgique, Brussels, Series 2, 7:86. 1859.
=Metasphaeria bellynckii (G.D. Westendorp)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:178. 1883. =Scleropleella bellynckii (G.D.
Westendorp) A. Munk, Dansk Botanisk
Arkiv, Kjgbenhavn 15(2):107. 1953. On
dead stems of Convallaria polygonatum L.,
Liliaceae. Belgium, Germany, Italy, Sweden.

berberidicola C.L. Spegazzini, Boletin de la
Academia Nacional de Ciencias en Cordoba
27:369-370. 1923. On leaves of Berberis
ilicifolia Forst., Berberidaceae. Argentina.

berberidis C. Richon, Catalogue Raisonné des
Champignons Qui Croissent Dans le Départe-

ment de la Marne, p. 300. 1889. On young,
living branches of Berberis vulgaris L.,
Berberidaceae. France.

berlesei P. Larsen & A. Munk, Dansk Botanisk
Arkiv, Kj@benhavn 14(7):23. 1952.
=Phaeosphaeria berlesei (P. Larsen & A.
Munk) A. Hedjaroude, Sydowia. Annales
Mycologici, Horn, N.O. 22(1-4):87. (1968)
1969. On dead stems of Equisetum hyemale
L., Equisetaceae. Canada, Denmark,
Germany, Hungary, U.S.A.

betulina F. Hazslinszky, Matematikai és Természet-
tudomanyi Ko6zlemenyek Vonatkozolag a
Hazai Viszonyokra, Budapest 25(2):152.
1892. =Metasphaeria betulina (F.A.
Hazslinszky) P.A. Saccardo & P. Sydow in
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Diggesit P.A. Saccardo
14:583-584. 1899. On branches of Betula sp.,
Betulaceae. Hungary.

betulina 1.E. Brezhnev, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 7:184. 1951. Nom.
illegit. Art. 64.1. On leaf spots of Betula
verrucosa Ehrh., caused by Phyllosticta
betulina Sacc., Betulaceae. U.S.S.R. (Kursk).

bicolor D. Hawksworth, W. Kaiser & B.N.
Ndimande, Mycologia, Lancaster, Pennsylva-
nia 71:483. 1979. On leaves of Saccharum
officinarum L., Gramineae. Kenya.

bicuspidata (M.C. Cooke & W.H. Harkness) A.N.
Berlese & P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
133. 1886. =Sphaeria (Leptosphaeria)
bicuspidata M.C. Cooke & W.H. Harkness,
Grevillea, London 13:19-20. 1884. On twigs
of Baccharis sp., Compositae. U.S.A.

biebersteinii S.A. Gucevicz, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 6:128—129.
1969. On dry branches of Cerastium
biebersteiniti DC., Caryophyllaceae. U.S.S.R.

biseptata B. Auerswald [cited without place of
publication in P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:58. 1883, sub.
Leptosphaeria michotii (G.D. Westendorp)
P.A. Saccardo]. On unknown substrate of
unknown host, unknown family. Unknown
country.

bispora (P. Larsen) A. Munk, Dansk Botanisk Arkiv,
Kjgbenhavn 17(1):376. 1957. =Metasphaeria
bispora P. Larsen, Dansk Botanisk Arkiv,
Kj@benhavn 14(7—8):33—34. 1952. On dead
leaves of Typha latifolia L., Typhaceae.
Denmark.

bitorulosa [erroneously so cited by P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 15:190.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

1901; and as Metasphaeria bitorulosa for
Leptosphaeria tritorulosa (M.J. Berkeley &
C.E. Broome) V. Cesati & G. de Notaris].

blumeri E. Miller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):275. 1950. On dead stems
of Medicago falcata L., Leguminosae.
Switzerland.

bocconiae (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:16.
1883. =Sphaeria (Caulicolae) bocconiae
M.C. Cooke & J.B. Ellis, Grevillea, London
7:10. 1876 [as bokoniae). =Heptameria
bocconiae (M.C. Cooke & J.B. Ellis) M.C.
Cooke, Grevillea, London 18:29. 1889. On
stems of Bocconia sp., Papaveraceae. U.S.A.
(New Jersey).

bomareae F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):494-496. 1950. On leaves
of Bomarea caldasii (HBK) Asch. & Graebn.,
Alstroemeriaceae. Ecuador.

bondari A.A. Bitancourt & A.E. Jenkins, Phytopa-
thology, Lancaster, Pennsylvania 25:886.
1935. On leaves and twigs of Citrus auran-
tium L., Citrus grandis Osb., Citrus limonia
Osb., Citrus medica L., Citrus sinensis Osb.,
Citrus sp., Rutaceae. Brazil.

borealis J.B. Ellis & B.M. Everhart, The North
American Pyrenomycetes. A Contribution to
Mycologic Botany, p. 353. 1892. On
decorticated wood of Salix sp., Salicaceae.
U.S.A.

borealis J.B. Ellis & B.M. Everhart var. populi J.
Dearness & H.D. House, Bulletin of the New
York State Museum, Albany 266:74. 1925.
On decorticated branch of Populus balsamif-
era L., Salicaceae. U.S.A. (New York).

bornmuelleri O. Jaap, Annales Mycologici, Berlin
14:15. 1916. On dry leaves of Rubia
peregrina L., Rubiaceae. Italy.

borziana P.A. Saccardo & F. Cavara, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze, New Series
7:283. 1900. On dead branches of Spartina
juncea auct., Gramineae. Italy.

boucera (M.C. Cooke & J.B. Ellis) J.B. Ellis in N.L.
Britton, Catalogue of Plants Found in New
Jersey. Geological Survey of New Jersey,
Final Report of the State Geologist, Trenton
2(1):525. 1889. =Sphaeria (Caulicolae)
boucera M.C. Cooke & J.B. Ellis, Grevillea,
London 8:15. 1879 [as bucera]. =Meta-
sphaeria boucera (M.C. Cooke & J.B. Ellis)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:161. 1883 [as bucera]. On dead herbaceous
stems of unknown host, unknown family.
U.S.A. (New Jersey).

brachyasca E. Rostrup, Meddelelser om Gronland,
Kjobenhavn 3(3):618. 1893 [published
1892?]. On stems and leaves of Saxifraga
oppositifolia L., Saxifragaceae. Greenland.

March 1991

brachypodii G. Passerini in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi
Mycologici Continuatio Edita Nova, Series 2,
Century 24, No. 2334. Anno 1876; Hedwigia,
Dresden 17:45. 1879. =Metasphaeria
brachypodii (G. Passerini) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:176.
1883. On dry leaves of Brachypodium
sylvaticum (Hudson) Beauv., Gramineae.
Italy.

brachysperma A.N. Berlese, Bulletin Trimestriel de
la Société Mycologique de France, Paris
5:41-42. 1889. On dead herbaceous stems of
unknown host, unknown family. Italy.

bractearum (P.A. Saccardo) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:58. 1883. =Lepto-
sphaeria carpophila P.A. Saccardo var.
bractearum P.A. Saccardo, Michelia Com-
mentarium Mycologicum Fungos in Primis
Italicos Illustrans 1:397. 1878. On rotting
bracts of heads on Dipsacus sylvestris
Hudson, Dipsacaceae. Italy.

brasiliensis C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cordoba 11:521.
1889; Fungi Puiggariani 1:143, No. 280.
1889. On living leaves of Xanthoxylum sp.
(=Zanthoxylum sp.), Anacardiaceae. Brazil.

brauni E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):246. 1950. On stems of
Hieracium murorum L., Compositae.
Switzerland.

bresadolaeana O. Jaap, Annales Mycologici, Berlin
14:14-15. 1916. On dry stems of preceding
year on Euphorbia wulfenii Hoppe ex Koch,
Euphorbiaceae. Yugoslavia.

brightonensis R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2695. 1984. On stems of
Agastache urticifolia (Benth.) Ktze., Labiatae.
U.S.A.

briosiana G. Pollacci, Atti dell’ Istituto Botanico
della Universita e Laboratorio Crittogamica di
Pavia, Milano, Series 2, 5:36. 1896. On living
leaves of Jubaea spectabilis Humb., Bonpl.,
& Kunth, Palmae. Italy.

brizae G. Passerini, Rendiconti della Sedute della R.
Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:91. 1887. =Phaeosphaeria
brizae (G. Passerini) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1523.
1989. On dry culms and peduncles of Briza
media L., Gramineae. Italy.

brunellae J.B. Ellis & B.M. Everhart, Proceedings of
the Academy of Natural Sciences of Philadel-
phia 1890:237. 1890. On dead stems of
Prunella vulgaris L., Labiatae. Canada.

bryophila P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:319. 1875. On stalks of

Nomenclator of Leptosphaeria

209

Tortula tortuosa Ehrh. ex Hedw. [=Tortella
tortuosa (Hedw.) Limpr.], Musci (Pottiaceae).
Italy.

bryzae G. Passerini, vide Leptosphaeria brizae G.
Passerini.

bubakii J. Klika, Annales Mycologici, Berlin 24:134.
1926. On dead stems of Zea mays L.,
Gramineae. Bulgaria.

buddlejae |. Hino & K. Katumoto, Bulletin of the
Faculty of Agriculture, Yamaguchi Univer-
sity, Shimonoseki 8:644-645. 1957. On
leaves of Buddleja davidii Franch., Logani-
aceae. Japan.

buddlejae S.A. Gucevicz, Trudy Gosudarstvennogo
Nikitskogo Botanicheskogo Sada, Yalta
29:190. 1959. Nom. illegit. Art. 64.1. On
dried stems of Buddleja davidii Franch.,
Loganiaceae. U.S.S.R. (Yalta).

bulgarica F. Petrak, Annales Mycologici, Berlin
29:368. 1931. On dry stems of Ruta grave-
olens L., Rutaceae. Bulgaria.

bupleuri P. Sydow, Hedwigia, Dresden 38:(142).
1899. On dry stems of Bupleurum petraeum
L., Umbelliferae. France.

buxina M. de Sousa da Camara, Revista
Agronomica, Lisbon 20(1):21—22. 1932. On
bark of trunks of Buxus sempervirens L.,
Buxaceae. Portugal.

byssincola R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2695. (1984) 1985. On
unknown substrate of Lupinus sp., Legumino-
sae. U.S.A. (Colorado).

caballeroi R. Gonzalez Fragoso, Memorias de la R.
Sociedad Espanola de Historia Natural,
Madrid 11:90. 1919. On dry stems of Datura
stramonium L., Solanaceae. Spain.

cacuminispora C.E. Fairman, Proceedings of the
Rochester Academy of Science, Rochester,
New York 6:98-99. 1921. On nuts of Carya
sp., Juglandaceae. U.S.A.

cadubriae C.L. Spegazzini, Atti della Societa
Crittogamologica Italiana, Milano, Series 2,
3(1):55. 1881. =Heptameria cadubriae (C.L.
Spegazzini) M.C. Cooke, Grevillea, London
18:31. 1889. =Nodulosphaeria cadubriae
(C.L. Spegazzini) L. Holm., Svensk Botanisk
Tidskrift, Stockholm 55:73-74. 1961. On
rotten, fallen stems of Epilobium dodonaei
Vill., Onagraceae. Italy.

caespitosa G, Niessl v. Mayendorf in J. Kunze,
Fungi Selecti Exsiccati, Century 1, No. 77.
Anno 1876. Nom. nud. Art. 32.1. On dry
stems and humid tissues of Artemisia
campestris L., Compositae. Germany.

caespitosa G. Niessl vy. Mayendorf ex P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:35.
1883. =Phaeoderris caespitosa (G. Niessl v.
Mayendorf ex P.A. Saccardo) F. v. Héhnel,
Sitzungsberichte der Akademie der Wissen-
schaften in Wien, Mathematisch-naturwissen-
schaftliche Klasse, Abt. 1, 120:462. 1911.

210 Illinois Natural History Survey Bulletin

On dry stems and humid tissues of Artemisia
campestris L., Compositae. Germany.

caespitosa G. Niessl y. Mayendorf forma salviae J.
Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(2):161.
1901. [Description effectively published:
Vorstudien zu einer Pilz-Florades Grossher-
zogthums, Luxemburg 1:262. 1899.] On dry
stems of Salvia pratensis L., Labiatae.
Luxembourg.

caffra F. v. Thiimen, Flora, Jena und Regensburg
59:363-364. 1876; Mycotheca Universalis,
Century 4, No. 351. Anno 1876. On living
fronds of Marattia sp., Marattiaceae (Fern).
South Africa.

californica (M.C. Cooke & W.H. Harkness) A.N.
Berlese & P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
137. 1886. =Sphaeria (Leptosphaeria)
californica M.C. Cooke & W.H. Harkness,
Grevillea, London 13:20. 1886. On twigs and
leaves of Araucaria imbricata Pav.,
Euonymus sp., Rhododendron sp., Sarotham-
nus sp., Araucariaceae, Celastraceae,
Ericaceae, Leguminosae. U.S.A.

calligoni I.P. Frolov in E.N. Koschkelova, I.P.
Frolov, and Z. Dzhuraeva, Mikoflora
Badkhyza, Karabilya i Yuzhnoi Chasti
Murgabskogo Oazisa (Mikromitsety) [The
Mycoflora of Badkhyz, Karabil and the
Southern Part of the Murgab Oasis (Micromy-
cetes)], p. 87. 1970. Nom. nud. Art. 32.1. On
bare, unsheathed (uncorticated?) wood of
Calligonum sp., Polygonaceae. U.S.S.R.
(Karabilia).

calligoni I.P. Frolov, Novosti Sistematiki Nizshikh
Rastenij, Novitates Systematicae Plantarum
Non Vascularium 7:192—193. 1970. On
decorticated twig of Calligonum sp., Poly-
gonaceae. U.S.S.R.

calopogonii R. Gonzalez Fragoso & R. Ciferri,
Boletin de la R. Sociedad Espanola de
Historia Natural, Madrid 27:272. 1927;
Publicaciones Estacion Agronomica de Moca,
Santa Domingo. Series B, Botanica, No.
11:21-22. 1928. Nom. inval. Art. 34.1. On
living or wilted leaves of Calopogonium
mucunoides Desv., Leguminosae. Dominican
Republic.

calvescens (E.M. Fries) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:315.
1875. =Sphaeria calvescens E.M. Fries,
Scleromyceti Sueciae. Collegit, Digessit et
Evulgaviti No. 401. [Apparently not
published; specimens of number 401 may be
scattered in herbaria.] =Pleospora calvescens
(E.M. Fries) L.R. Tulasne & C. Tulasne,
Selecta Fungorum Carpologica 2:266. 1863.
=Pyrenophora calvescens (E.M. Fries) P.A.
Saccardo, Sylloge Fungorum Omnium

Vol. 34 Art. 3

Hucusque Cognitorum Diggesit P.A. Saccardo
2:279. 1883. =Cucurbitaria calvescens (E.M.
Fries) V. Cesati & G. de Notaris, Commen-
tario della Societa Crittogamologica Italiana,
Milan 1:215. 1863. =Chaetoplea calvescens
(E.M. Fries) F.E. Clements in F.E. Clements
and C.L. Shear, The Genera of Fungi, p. 275.
1931 [as (Fr.) Sacc.]. On dry stems of
Atriplex hortensis L., Chenopodium album L.,
Chenopodiaceae. France.

calvescens (E.M. Fries) P.G. Crivelli, Ueber die
Heterogene Ascomycetengattung Pleospora
Rabh.; Vorschlag fiir eine Aufteilung.
Abhandlung zur Erlangung des Titels eines
Doktors der Naturwissenschaften der
Eidgenodssischen Technischen Hochschule,
Ziirich No. 7318:177. 1983. Nom. illegit. Art.
64.1.

camelliae M.C. Cooke & G.E. Massee, Grevillea,
London 16:5. 1877. On living leaves of
Camellia sp., Theaceae. Australia.

camelliae-japonicae W. Siemaszko, Acta Societatis
Botanicorum Poloniae, Warszawa 1:21. 1923.
On leaves of Camellia japonica L., Theaceae.
ULS.S.R.

camilla P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:312. 1875. On wilted
leaves of Scrophularia aquatica L., Scrophu-
lariaceae. Italy.

camphorata G. Passerini, Atti dell’ Reale Accademia
Nazionale dei Lincei. Rendiconti, Rome,
Series 4, 7(2):44. 1891. On dry stems of
Artemisia camphorata Vill., Compositae.
Italy.

camphorosmae A.\. Lobik, Materialy po Floristich-
eskim i Faunisticheskim Obsledovaniyam
Terskogo Okruga [Data from Investigations
on the Flora and Fauna of the Ter Region],
p- 23. 1928. On leaves of Camphorosma
perennis Pall., Chenopodiaceae. U.S.S.R.

campisilii C.L. Spegazzini, Revue Mycologique,
Toulouse 2:32. 1880; Michelia Commentar-
ium Mycologicum Fungos in Primis Italicos
Illustrans 2:162. 1880. On dead leaves of
Lycopodium annotinum L., Lycopodiaceae.
Italy.

canadensis G. de Notaris [as Leptosphaeria
canadensis G. de Notaris in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:163.
1883]. Nom. nud. Art. 32.1. =Metasphaeria
canadensis P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:163. 1883 [as (De Not.) Sacc.].
On dry stems of Erigeron canadensis L.,
Compositae. Canada, Italy.

canephorae R.L. Steyaert, Bulletin. Société R. de
Botanique de Belgique, Bruxelles 80(Series 2.
30):22. 1948. On leaves of Coffea canephora
Pierre ex Froehn., Rubiaceae. Zaire.

March 1991

caninae (W. Phillips & C.B. Plowright) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:81. 1883. =Sphaeria caninae W. Phillips &
C.B. Plowright, Grevillea, London 6:27. 1877.
On thallus of Pe/tigera canina (L.) Willd.,
Lichenes (Peltigeraceae). Great Britain.

cannabina T. Ferraris & C. Massa, Annales
Mycologici, Berlin 10:286. 1912. On wilted
leaves of Cannabis sativa L., Cannabaceae.
Italy.

cannae D. McAlpine, Proceedings of the Linnean
Society of New South Wales, Sydney 27:375.
1902. On upper surface of wilted leaves of
Canna indica L., Cannaceae. Australia.

capparidicola B.B. Mundkur & S. Ahmad, Myco-
logical Papers. Commonwealth Mycological
Institute, Kew 18:4. 1946. On dead branches
of Capparis aphylla Roth, Capparaceae.
India.

capparidis G. Passerini, Revue Mycologique,
Toulouse 2:34—35. 1880. On rotten branches
of Capparis spinosa L., Capparaceae. Italy.

caprifolii P. Brunaud, Bulletin de la Société
Botanique de France, Paris 34:429. 1887. On
dead vines of Lonicera caprifolium L.,
Caprifoliaceae. France.

capsularum F, Cavara, Fungi Longobardiae
Exsiccati Sive Mycetum Specimina in
Longobardia Collecta, exsiccata et Speciebus
Novis Vel Criticis, Iconibus Illustrata,
Pugillus 4, No. 174. Anno 1894. On dry
capsules of Oenothera biennis L., Ona-
graceae. Italy.

capsularum F. Cavara, Hedwigia, Dresden 34:(37).
1895; Botanisches Zentralblatt, Jena &
Dresden 61:316. 1895. On dried capsules of
Oenothera biennis L., Onagraceae. Italy.

carduina G. Passerini, Atti del Reale Istituto Veneto
di Lincei Rendiconti, Rome, Series 4,
4(2):58-59. 1888. On bracts of Carduus
nutans L., Compositae. Italy.

carduorum (K.F.W. Wallroth) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:235. 1863.
=Sphaeria carduorum K.F.W. Wallroth, Flora
Crypogamica Germaniae 2:805. 1833. On dry
stems of Carduus sp., Compositae. Belgium,
France, Germany, Great Britain.

caricicola F, Fautrey in C. Roumeguere, Revue
Mycologique, Toulouse 15:20. 1893; Fungi
Selecti Gallici Exsiccati, Century 63, No.
6243. Anno 1893. =Phaeosphaeria cari-
cicola (F. Fautreyy A. Leuchtmann, Sydowia.
Annales Mycologici, Horn, N.O. 37:109.
1984. On leaves of Carex pendula Hudson,
Carex riparia Curtis, Cyperaceae. France,
Switzerland.

caricina J, Schroter in F.J. Cohn, Kryptogamen-
Flora Von Schlesien. Im Namen Der Schlesis-
chen Gesellschaft fiir vaterlandische Cultur

Nomenclator of Leptosphaeria 211

herausgegeben von Prof. Dr. Ferdinand Cohn,
Secretair Der Botanischen Section, Breslau
3(2):361. 1894. On leaves of Carex panicea
L., Carex silvatica auct., Cyperaceae. Poland.

caricinella P.A. Karsten, Ofversigt af K. Vetenskap-
sakademiens Forhandlingar, Stockholm
29(2): 100-101. 1872. =Phaeosphaeria
caricinella (P.A. Karsten) O. Eriksson, Arkiv
for Botanik, Uppsala, Stockholm, Series 2,
6:414. 1967. On dried leaves of Carex pulla
Good., Cyperaceae. Canada, Norway.

cariciphila C.A. Oudemans, Nederlandsch kruidkun-
dig Archief. Verslagen en Mededeelingen der
Nederlandsche Botanische Vereeniging,
Leiden, Series 3, 2:870. 1903. On sheaths of
Carex arenaria L., Cyperaceae. Netherlands.

caricis J. Schroter, Jahresbericht der Schlesischen
Gesellschaft fiir vaterlandische Kultur,
Breslau 58:175. (1880) 1881.
=Phaeosphaeria caricis (J. Schréter) A.
Leuchtmann, Sydowia. Annales Mycologici,
Horn, N.O. 37:147. 1984. On leaves of Carex
hirta L., Carex pendula Hudson, Carex
vaginata Tausch, Dactylis glomerata L.,
Luzula sylvatica (Hudson) Gaudin, Typha
latifolia L., Cyperaceae, Gramineae, Jun-
caceae, Typhaceae. Canada, Czechoslovakia,
Sweden, Switzerland, U.S.A.

caricis-firmae F. Petrak, Sydowia. Annales Mycol-
ogici, Horn, N. O. 1(4-6):315. 1947. On
living or dead leaves of Carex firma Host,
Cyperaceae. Austria.

caricis-vulpinae A.I. Lobik, Bolezni Rastenii S.
Peterburg (Morbi Plantarum Scripta Sectionis
Phytopathologiae Horti Botanici Principalis)
17:161. 1928. On leaves of Carex vulpina L.,
Cyperaceae. U.S.S.R.

carlinoides M. Losa, Anales del Jardin Botanico de
Madrid 8:305. (1947) 1948. On dead stems of
Carduus carlinoides Gouan, Compositae.
Spain.

carneomaculans F. Petrak, Annales Mycologici,
Berlin 25:287—288. 1927. On dry stems of
Valeriana officinalis L., Valerianaceae.
Czechoslovakia.

carpogena P.A, Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:313. 1875. On dried
capsules of Veronica urticifolia Jacq.,
Scrophulariaceae. Italy.

carpophila P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:313—314. 1875. On rotting
capsules of Tecoma radicans Juss., Bignoni-
aceae. Italy.

carpophila P.A. Saccardo var. bractearum P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
1:397. 1878. =Leptosphaeria bractearum
(P.A. Saccardo) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:58. 1883. On rotting

bracts of heads of Dipsacus sylvestris Hudson,
Dipsacaceae. Italy.

cassiaecola J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 2:41. 1886. On
dead stems of Cassia sp., Leguminosae.
U.S.A.

casta Iu. Voronov, Leningrad. Vsesoiuzny! Institut
Zashchity Rastenii. Otdel. Fitopatologii
Materialy po Mikologii i Fitopatologii
4(1):60. 1922. On twigs of Vitex agnus-
castus L., Verbenaceae. U.S.S.R.

castagnei (M.C. Durieu de Maisonneuve & J.P.
Montagne) P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:317. 1875.
=Sphaeria castagnei M.C. Durieu de
Maisonneuve & J.P. Montagne in M.C.
Durieu de Maisonneuve, Exploration
Scientifique de L’ Algérie Publi¢ée Par Ordre
du Gouvernement. Series Naturelles.
Botanique I, Premiere Partie, p. 528. 1869.
=Sphaeria jasmini J.L. Castagne, Catalogue
Des Plantes qui Croissent Naturellement aux
Environs de Marseille, p. 167. 1845. Non
Sphaeria jasmini L.D. v. Schweinitz.
=Pleospora jasmini (J.L. Castagne) L. Fuckel,
Symbolae Mycologicae, p. 138. 1870.
=Paraphaeosphaeria castagnei (M.C. Durieu
de Maisonneuve & J.P. Montagne) O.
Eriksson, Arkiv for Botanik, Uppsala,
Stockholm, Series 2, 6:406. 1967. On
branches of Euonymus europaeus L.,
Celastraceae. France.

castilleiae F.E. Clements, Bulletin of the Torrey
Botanical Club (and Torreya), New York
30:85. 1903. On dead stems of Castilleja
pallida (L.) Sprengel, Scrophulariaceae.
U.S.A.

castillejae R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2695-2696. 1984. On
stems of Castilleja miniata Dougl. ex Hook.,
Scrophulariaceae. U.S.A.

castrensis R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2696. 1984. On stems of
Lupinus obtusilobus Heller, Leguminosae.
U.S.A.

catalaunica R. Gonzalez Fragoso, Introduccion al
Estudio de la Fl6rula de Micromicetos de
Catalufa, Musei Barcinonensis Scientiarum
Naturalium Opera. Series Botanica 2, pp.
89-90. 1917. Nom. inval. Art. 34.1; Florula
de Micromicetos de Cataluna (Publicacions de
la Junta de Ciéncies Naturals de Barcelona),
pp. 89-90. 1917. Nom. inval. Art. 34.1. On
thin branches of Smilax aspera L., Smila-
caceae. Spain.

cattanei F. vy. Thiimen, Die Pilze der Reispflanze
(Oryza sativa Lin.). Eine Monographie, p. 5.
1889 [as catanei]; Verlag der K.K. Versuchs-
Station fiir Wein-und Obstbau zu Klos-

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

terneuburg bei Wein, No. 12, p. 5. 1889.
=Phaeosphaeria cattanei (F. v. Thiimen) I.
Miyake, Journal of the College of Agriculture,
Imperial University of Tokyo 2(4):247. 1910.
=Trematosphaerella cattanei (F. v. Thimen)
G. Padwick, Manual of Rice Diseases,
Commonwealth Mycological Institute. Kew,
Surrey, p. 153. 1950. On wilted culms,
sheaths, and leaves of Oryza sativa L.,
Gramineae. Italy.

cattanei (P.A. Saccardo) A. Jaczewski, Opredielitel”
Gribov’ 2, Fungi Imperfecti (Petrograd), p.
608. 1917. =Metasphaeria cattanei P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:176. 1883. =Pleospora endiusae (L.
Fuckel) A. Cattaneo var. major A. Cattaneo,
Atti dell’ Istituto Botanico della Universita e
Laboratorio Crittogamica di Pavia, Milano.
Series 1, 2:125. 1879. On wilted leaves of
Oryza sativa L., Gramineae. Italy.

caucana F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 5(3—-6):244. 1951. In the stromata
of Phyllachora sp., Fungi (Phyllachoraceae).
Colombia.

caulincola (L.D. v. Schweinitz) M.A. Sherwood,
Mycotaxon. An International Journal
Designed to Expedite Publication of Research
on Taxonomy & Nomenclature of Fungi &
Lichens, Ithaca, New York 5:165. 1977.
=Stictis caulincola L.D. v. Schweinitz,
Synopsis Fungorum in America Boreali
Media Digentium. Secundum Observationes
Ludovici Davidis de Schweinitz, p. 179. 1832.
=Phragmonae viacaulicola (L.D. v. Schwein-
itz) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 8:676. 1889. On stems of unknown
host, unknown family. U.S.A. (Pennsyl-
vania).

caulium P.A. Saccardo. Nom. nud. [Falsely cited in
literature as published in Reliquiae Liber-
tianae II, No. 119. Anno 1881.] Vide
Leptosphaeria vagabunda P.A. Saccardo var.
caulium C. Roumeguére & P.A. Saccardo.

cavanillesii M.J. de Urries y Azara, Anales del
Jardin Botanico de Madrid 6:337-397. 1946.
Nom. inval. Art. 36.1. On branch of
Lavandula sp., Labiatae. Spain.

cavarae M. Curzi, Atti dell’ Istituto Botanico della
Universita e Laboratorio Crittogamica di
Pavia, Milano, Series 3, 3:62. 1927. In living
leaves of Thea sinensis L., Theaceae. Italy.

ceanothi (M.C. Cooke & H.W. Harkness) A.N.
Berlese & P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
136. 1886. =Sphaeria (Didymosphaeria)
ceanothi M.C. Cooke & H.W. Harkness,
Grevillea, London 13:19. 1895. On twigs of
Ceanothus sp., Rhamnaceae. U.S.A.

March 199]

ceballosi L.M. Unamuno Yrigoyen, Boletin de la R.
Sociedad Espanola de Historia Natural,
Madrid 33:230—231. 1933. On dried leaves of
Cyperus schoenoides Griseb., Cyperaceae.
Spain.

cecropiae C.E. Chardon, Farlowia. A Journal of
Cryptogamic Botany, Cambridge, Massachu-
setts 2:466. 1946. On leaves of Cecropia
peltata L., Urticaceae. Dominican Republic.

centaureae E. Miiller, Sydowia. Annales Mycol-
ogici, Horn, N.O. 4(1-6):299. 1950. =Nodu-
losphaeria centaureae (E. Miller) L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):85. 1957. On dead stems of Carduus
defloratus L., Centaurea scabiosa L.,
Compositae. Switzerland.

centrafricana A.M. Saccas, Etude de la Flore
Cryptogamique des Caféiers en Afrique
Centrale. Bulletin de Institut Frangais du Café
du Cacao et d’Autres Plantes Stimulantes
(Bulletin IFCC No. 16), pp. 223-225. 1981.
Nom. inval. Art. 37.1. On dead stems of
Coffea excelsa Cheval., Rubiaceae. Central
African Republic.

cephalariai-uralensis [as cephalariae-uralensis|
N.A. Naumov & T.L. Dobrozrakova,
Materialy po Mikologii i Fitopatologii Rossi,
Petrograd 8(2):134. 1929. On dry stems of
Cephalaria uralensis (Murray) Roemer &
Schultes, Dipsacaceae. U.S.S.R.

cerastii J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg 1(3):221.
1903. On dry stems of Eupatorium can-
nabinum L., Compositae. Luxembourg.

ceratispora (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:87. 1883. =Sphaeria ceratispora M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:150-151. 1876. On
herbaceous stem of Zea sp., Gramineae.
U.S.A.

cercocarpi H. Sydow & P. Sydow, Annales
Mycologici, Berlin 5:339. 1907. On dead
leaves of Cercocarpus ledifolius Nutt. ex
Torr. & Gray, Rosaceae. U.S.A.

cerei-peruvianit W. Siemaszko, Acta Societatis
Botanicorum Poloniae, Warszawa 1:22. 1923.
On cladodes of Cereus peruvianus (L.) Miller,
Cactaceae. Poland, U.S.S.R.

cerlettii C.L. Spegazzini, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:398. 1878; Rivista di Viticoltura,
Enologia ed Agraria, Conegliano 3:54—56.
1879. =Metasphaeria cerlettii (C.L. Spegazz-
ini) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:167. 1883. =Chaetopyrena
cerletti (C.L. Spegazzini) A. Trotter in P.A.
Saccardo, Sylloge Fungorum Omnium

Nomenclator of Leptosphaeria 213

Hucusque Cognitorum Digessit P.A. Saccardo
24:961. 1928. On dead vines of Vitis vinifera
L., Vitaceae. Italy.

cervispora (K. Kalchbrenner & M.C. Cooke) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:44. 1883. =Sphaeria cervispora K.
Kalchbrenner & M.C. Cooke, Grevillea,
London 9:29. 1880. On dead stems of
Artemisia sp., Compositae. South Africa.

cesatiana (J.P. Montagne ex V. Cesati & G. de
Notaris) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):39. 1957.
=Rhaphidospora cesatiana J.P. Montagne ex
V. Cesati & G. de Notaris, Commentario della
Societa Crittogamologica Italiana, Milan
1:234. 1863. =Ophiobolus cesatianus (J.P.
Montagne ex V. Cesati & G. de Notaris) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:44. 1883. On stems of Echium vulgare L.,
occasionally on Brassica oleracea L.,
Hypericum perforatum L., Lycopus europaeus
L., Boraginaceae, Cruciferae, Guttiferae,
Labiatae. Austria, Great Britain, Italy.

chaetostoma P.A. Saccardo, Michelia Commentar-
ium Mycologicum Fungos in Primis Italicos
Illustrans 2:533-534. 1882. =Metasphaeria
chaetostoma (P.A. Saccardo) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:165.
1883. On vines (stems) of Vitis vinifera L.,
Vitaceae. Italy.

chamaeropis M. Chochrjakov, Notulae Systematicae
e Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komaroy Academiae Scientia-
rum U.R.S.S., Petropolis 7:143—144. 1951.
On living leaves of Chamaerops humilis L.,
Palmae. U.S.S.R.

chelidonti F. Fautrey in F. Fautrey and J.B. Lam-
botte, Revue Mycologique, Toulouse 17:168.
1895. On dried stems of Chelidonium majus
L., Papaveraceae. France.

chenopodii-albi W. Siemaszko, Acta Societatis
Botanicorum Poloniae, Warszawa 1:21. 1923.
On leaves of Chenopodium album L.,
Chenopodiaceae. U.S.S.R.

chilensis C.L. Spegazzini, Fungi Chilenses, p. 78.
1910; Revista de La Facultad de Agronomia
Y Veterinaria, Universidad Nacional de La
Plata, Series 2, 6:78. 1910. On dead branches
of Lomatia obliqua R. Br., Proteaceae. Chile.

chochrjakovii (L.N. Vasil’eva) L.N. Vasil’eva,
Pirenomitsety 1 Lokuloaskomisety Severa
Dal’nego Vostoka (Leningrad:*“Nauka’):125.
1987. =Nodulosphaeria chochrjakovii L.N.
Vasil’eva, Mikologiia i Fitopatologia
Akademiia Nauk SSSR, Leningrad 13:278.
1979. On dead leaves of Hedysarum
obscurum L., Leguminosae. U.S.S.R.

214 Illinois Natural History Survey Bulletin

chondri L.K. Rosenvinge, Botanisk Tidsskrift,
Kj@benhavn 27:XXXIII-XXXV. 1906.
=Leptosphaeria marina E. Rostrup. Non
Leptosphaeria marina J.B. Ellis & B.M.
Everhart.

chrysanthemi F.E. Clements & E.S. Clements,
Cryptogamae Formationum Coloradensium,
Century 1, No. 24. Anno 1906. Nom. inval.
Art. 36.1. On aged stems of Carduus
scopulorum Greene, Compositae. U.S.A.

chusqueae H. Sydow, Annales Mycologici, Berlin
37:376-377. 1939. On dead, deciduous
leaves of Chusquea serrulata Pilger, Grami-
neae. Philippines.

cibostii (G. de Notaris) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:235. 1863. =Sphaeria
cibostii G. de Notaris, Memorie della
Accademia delle Scienze di Torino, Series 2,
13:103. 1853. On dead stems of unknown
host, Umbelliferae. Italy.

cinclidoti A. Racovitza, Mémoires du Muséum
Nationale d’Histoire Naturelle, Paris, Series
B, Botanique 10:150. 1959. On dead
phylloids of Cicclidotus fontinaloides (Hedw.)
P. Beauv. [Cinclidotus is an orthographic
variant.], Musci (Pottiaceae). France.

(Metasphaeria) cinerea (L. Fuckel) H.G. Winter,
Hedwigia, Dresden 26:58. 1887. =Cucurbi-
taria cinerea L. Fuckel, Fungi Rhenani
Exsiccati A Leopoldo Fuckel Collecti
Supplement, Fascicle 6, No. 2044. Anno
1867. =Sphaeria cinerea (L. Fuckel) L.
Fuckel, Symbolae Mycologicae, p. 114. 1870.
=Metasphaeria cinerea (L. Fuckel) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:166. 1883. On branches of Salix aurita L.,
Salicaceae. Austria.

cinnamomi M. Shirai & K. Hara, Botanical Maga-
zine, Tokyo 25:72. 1911. On diseased
branches of Cinnamomum camphora T. Nees
ex Eberm., Lauraceae. Japan.

circinans (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:88. 1883. =Bysso-
thecium circinans L. Fuckel, Botanische
Zeitung, Berlin & Leipzig 19:251. 1861.
=Trematosphaeria circinans (L. Fuckel) H.G.
Winter, Dr. L. Rabenhorst’s Kryptogaman-
Flora von Deutschland, Oesterreich und der
Schweiz, Second edition, 1(2) Ascomyceten:
Gymnoasceen und Pyrenomyceten, p. 277.
1885. =Passeriniella circinans (L. Fuckel)
A.N. Berlese in P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 11:326. 1895. On
roots and stems of Medicago sativa L.,
Leguminosae. Germany.

cirricola G, Passerini, Rendiconti della Sedute della
R. Accademia dei Lincei, Classe di Scienze

Vol. 34 Art. 3

Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:90. 1887. On dry denuded
stems of Vitis vinifera L., Vitaceae. Italy.

cirstt-arvensis M. Losa, Anales del Jardin Botanico
de Madrid 8:306. (1947) 1948. On dry culms
of Cirsium arvense (L.) Scop., Compositae.
Spain.

cisti L. Celotti, Miceti del Parco e Dintorni della
Scuola Nazionale di Agricoltura di Montpel-
lier Raccolti e Studiati, Congliano, p. 32.
1887. On branches of Cistus albidus L.,
Cistaceae. France.

cisticola M. de Sousa da Camara, Agronomia
Lusitana, Sacavém 9:93—94. 1947. On
branches of Cistus monspeliensis L., Cista-
ceae. Portugal.

cistina M.J. de Urries y Azara, Boletin de la R.
Sociedad Espanola de Historia Natural,
Madrid 33:95. 1933. On dead branchlets of
Cistus ladanifer L., Cistaceae. Spain.

citricola P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:416. 1882. On living leaves of
Citrus sp., Rutaceae. Italy.

cladii D. Cruchet, Bulletin de la Société Vaudoise
des Sciences Naturelles, Lausanne 55:161.
1923. On roots of Cladium mariscus (L.)
Pohl, Cyperaceae. Switzerland.

cladophila J. Schroter, Jahresbericht der Schlesi-
schen Gesellschaft fiir vaterlandische Kultur,
Breslau 58:174. (1880) 1881. On living stems
of Ribes alpinum L., Saxifragaceae. Sweden.

clara (B. Auerswald) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:73. 1883.
=Sphaeria clara B. Auerswald in M.C.
Cooke, Grevillea, London 5:121. 1877. On
leaves of Carex sp., Sparganium sp..
Cyperaceae, Sparganiaceae. Germany, Great
Britain.

clarkii D. Hawksworth, Transactions of the British
Mycological Society, London 74:376. 1980.
On thallus of Peltigera cf. rufescens (Weis.)
Humb., Lichenes (Peltigeraceae). Great
Britain.

clavata A.L. Guyot, Revue de Mycologie, Paris
11:62. 1946. =Massariosphaeria clavata
(A.L. Guyot) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1582. 1989. On
dry stems and leaves of Koeleria albescens
DC., Koeleria gracilis Pers., Gramineae.
France.

clavicarpa J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 1:43—44. 1885;
Hedwigia, Dresden 25:109. 1886.
=Heptameria clavicarpa (J.B. Ellis & B.M.
Everhart) M.C. Cooke, Grevillea, London
18:32. 1889. On dead culms of Phragmites
communis Trin., Gramineae. U.S.A.

March 1991

clavigera (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:44.
1883. =Sphaeria (Caulicolae) clavigera M.C.
Cooke & J.B. Ellis, Grevillea, London 6:16.
1877. On stems of Phytolacca americana L.,
Phytolaccaceae. U.S.A.

clavispora J.H. Miller & M.G. Burton, Mycologia,
Lancaster, Pennsylvania 34:1—2. 1942. On
dead stems of Eupatorium capillifolium
(Lam.) Small, Compositae. U.S.A.

clelandii C.G. Hansford, Proceedings of the Linnean
Society of New South Wales, Sydney 82:216.
1957. On dead branches of Acacia kempeana
F. Muell., Leguminosae. Australia.

clerodendri S.A. Gucevicz, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 7:159. 1970. On
bark and denuded wood of Clerodendrum
foetidi Bunge, Verbenaceae. U.S.S.R.

clivensis (M.J. Berkeley & C.E. Broome) G.L.

’ Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio. II] Ausgabe (Editio nova, Series
Secunda), Century 10, No. 947. Anno 1866.
=Sphaeria clivensis M.J. Berkeley & C.E.
Broome, Annals and Magazine of Natural
History, London, Series 2, 9:379. 1852.
=Pleospora bardanae G. Niessl v. Mayendorf,
Verhandlungen des Naturforschenden Vereins
in Briinn 14:178. 1876. On stems of Senecio
atratus Greene, Compositae. U.S.A.

clivensis (M.J. Berkeley & C.E. Broome) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:16. 1883. Nom. illegit. Art. 64.1.
=Sphaeria (Caulicolae) clivensis M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 9:379. 1852. =Diapleella clivensis (M.J.
Berkeley & C.E. Broome) A. Munk, Dansk
Botanisk Arkiv, Kj@benhavn 15(2):75. 1953.
=Kalmusia clivensis (M.J. Berkeley & C.E.
Broome) M.E. Barr, Mycotaxon. An Interna-
tional Journal Designed to Expedite Publica-
tion of Research on Taxonomy & Nomencla-
ture of Fungi & Lichens, Ithaca, New York
29:504. 1987. On dead stems of Pastinaca
sativa L., Senecio atratus Greene, Umbellif-
erae, Compositae. Germany, Great Britain,
U.S.A.

clivensis (M.J. Berkeley & C.E. Broome) P.A.
Saccardo var. constricta F.E. Clements & E.S.
Clements, Cryptogamae Formationum
Coloradensium, Century 5, No. 433. Anno
1908. Nom. nud. Art. 32.1. On aged stems of
Senecio atratus Greene, Compositae. U.S.A.

coccodes P.A. Karsten, Fungi Fenniae Exsiccati,
Century 10, No. 963. Anno 1870. =Mera-
sphaeria coccodes (P.A. Karsten) P.A.

Nomenclator of Leptosphaeria 215

Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:174. 1883. On culms of Calamagrostis sp.,
Gramineae. Finland.

coccothrinacis R. Ciferri & R. Gonzalez Fragoso,
Boletin de la R. Sociedad Espanola de
Historia Natural, Madrid 26:251—252. 1926.
On living leaves of Coccothrinax argentata
(Jacq.) Bailey, Palmae. Dominican Republic.

cocoes J. Verissimo d’Almeida & M. de Sousa da
Camara, Revista Agronémica, Lisbon
2(12):384. 1904; Boletim da Sociedade
Broteriana, Coimbra 24:163—164. 1909. On
leaves of Cocos romanzoffiana Cham.
{=Arecastrum romanzoffianum (Cham.)
Beccari], Palmae. Portugal.

coffaeicida C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cérdoba 23:560-S61.
1918. On living leaves of Coffea sp.,
Rubiaceae. Costa Rica.

coffeigena (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:51. 1883. =Sphaeria (Foliicolae) cof-
feigena M.J. Berkeley & M.A. Curtis, Journal
of the Linnean Society (Botany) London
10:389. 1868; Fungi Cubenses No. 862. Anno
1868. On leaves of Coffea arabica L.,
Rubiaceae. Cuba.

coffeigena (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo var. longirostrata C. Moreau & M.
Moreau, Revue de Mycologie, Paris (Supple-
ment Colonial No. 1) 16:21—25. 1951. On
trunk and branches of Coffea canephora
Pierre ex Froehn., Rubiaceae. Ivory Coast.

coicis K. Sawada, Report. Government Research
Institute, Department of Agriculture, Formosa
87:30. 1944. [Descriptive catalogue of the
Formosan Fungi...Taihoku, Formosa 10:30.
1944.] Nom. inval. Art. 36.1. On leaves of
Coix lacryma-jobi L. var. susutama Honda,
Gramineae. Taiwan.

coleosanthi C.E. Fairman, Mycologia, Lancaster,
Pennsylvania 10:246. 1918. On old stems of
Coleosanthus reniformis (Gray) Rydb.
(=Brickellia), Compositae. U.S.A. (New
Mexico).

collinsoniae J. Dearness & H.D. House, Bulletin of
the New York State Museum, Albany
233-234:36-37. (1920) 1921. On dead,
usually blackened stems of Collinsonia
canadensis L., Labiatae. U.S.A.

collumiae (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:50. 1883. =Sphaeria collumiae M.J.
Berkeley & M.A. Curtis, Proceedings of the
American Academy of Arts and Sciences,
Boston 4:130. 1860 [as cullumiae}]. On leaves
of Collomia squarrosa Nutt., Polemoniaceae.
South Africa.

216 Illinois Natural History Survey Bulletin

colocasiae L.M. Unamuno Yrigoyen, Memorias de
la R. Sociedad Espanola de Historia Natural,
Madrid 15:346. 1929. On living leaves of
Colocasia antiquorum Schott, Araceae.
Spain.

comatella (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:32.
1883. =Sphaeria (Caulicolae) comatella
M.C. Cooke & J.B. Ellis, Grevillea, London
5:52. 1876. On stems of Asparagus sp.,
Baptisia sp., Daucus sp., Solanum sp.,
Liliaceae, Leguminosae, Umbelliferae,
Solanaceae. U.S.A.

complanata (H.J. Tode:E.M. Fries) V. Cesati & G.
de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:236. 1863.
=Sphaeria complanata H.J. Tode:E.M. Fries,
Fungi Mecklenburgenses Selecti 2:21. 1791;
E.M. Fries, Systema Mycologicum Sistens
Fungorum 2:508. 1823. =Metasphaeria
complanata (H.J. Tode:E.M. Fries) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:161. 1883. On herbaceous stems of
unknown host, unknown family. Great
Britain, U.S.A.

compositarum E. Miiller, Sydowia. Annales
Mycologici, Horn, N.O. 4(1-6):213. 1950.
=Massariosphaeria compositarum (E. Miiller)
A. Leuchtmann, Sydowia. Annales Mycol-
ogici, Horn, N.O. 37:173. 1984. On dead
stems of unknown host, Compositae.
Switzerland.

compressa (H. Rehm) L. Holm, Symbolae Botanicae
Upsalienses 14(3):29. 1957. =Ophiobolus
compressus H. Rehm, Bericht des Naturwis-
senschaftlichen Vereins fiir Schwaben und
Neuberg, Augsburg 26:49. 1881.
=Rhaphidospora compressa H. Rehm,
Ascomyceten, Fascicle 4, No. 189. Anno
1873. Nom. nud. Art. 32.1. On dry stems of
Achillea millefolium auct., Artemisia
austriaca Jacq., Artemisia campestris L.,
Cirsium altissimum (L.) Spreng., Compositae.
Hungary, Sweden, U.S.A.

concentrica J.B. Ellis & B.M. Everhart, The North
American Pyrenomycetes. A Contribution to
Mycologic Botany, p. 354. 1892. On leaves
of Malus sp., Rosaceae. U.S.A.

concinna J.B. Ellis & B.M. Everhart, Proceedings of
the Academy of Natural Sciences of Philadel-
phia 1895:423. 1895. On dead stems of
Castilleja miniata Doug. ex Hook., Scrophu-
lariaceae. U.S.A.

conferta G. Niessl v. Mayendorf in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:20.
1883. On stems of Farsetia incana (L.) R.
Br., Cruciferae. Czechoslovakia.

Vol. 34 An. 3

congesta M.T. Lucas, Transactions of the British
Mycological Society, London 46:362. 1963.
On stems of Erigeron canadensis L.,
Compositae. Portugal.

coniformis (E.M. Fries:E.M. Fries) J. Schréter in F.J.
Cohn, Kryptogamen-Flora Von Schlesien. Im
Namen Der Schlesischen Gesellschaft fiir
vaterlandische Cultur herausgegeben von
Prof. Dr. Ferdinand Cohn, Secretair der
Botanischen Section, Breslau 3(2):369. 1894
{as conformis}]. On dead stems of Urtica
dioica L., Urticaceae. Poland.

coniformis (E.M. Fries:E.M. Fries) H.G. Winter, Dr.
L. Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):469. 1885 [as (L. Fuckel)
Winter]. =Sphaeria coniformis E.M.
Fries:E.M. Fries, Systema Mycologicum
Fungorum 2:508. 1823. =Pleospora coni-
formis (E.M. Fries:E.M. Fries) L. Fuckel,
Symbolae Mycologicae, p. 136. 1870.
=Metasphaeria coniformis (E.M. Fries:E.M.
Fries) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:159. 1883. On dry stems of
Euphrasia lutea L., Scrophulariaceae.
Austria.

(Metasphaeria) coniformis (E.M. Fries:E.M. Fries)
H.G. Winter, Hedwigia, Dresden 26:58. 1887
{as (L. Fuckel) Winter]. Nom. illegit. Art.
64.1. On stems of Euphrasia lutea L.,
Scrophulariaceae. Austria.

conti C.L. Spegazzini, Anales del Museo Nacional
de Historia Natural de Buenos Aires 19(Series
3, 12):382—383. 1909; Mycetes Argenti-
nenses, Series 4, p. 382, No. 628. Anno 1909.
Nom. illegit. Art. 64.1. Non Leptosphaeria
conii E. Rostrup. =Leptosphaeria coniigena
P.A. Saccardo & A. Trotter. On decaying
stems of Conium maculatum L., Umbelliferae.
Argentina.

conti E. Rostrup, Botanisk Tidsskrift, Kjobenhavn
26:311. 1905. On stems of Conium macula-
tum L., Umbelliferae. Denmark.

coniigena P.A. Saccardo & A. Trotter, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 22:220. 1913.
=Leptosphaeria conti C.L. Spegazzini, nec
Leptosphaeria conii E. Rostrup. On stems of
Conium maculatum L., Umbelliferae.
Argentina.

conimbricensis A.N. Berlese & F. Saccardo, Revue
Mycologique, Toulouse 11:121. 1889. On
dead culms of unknown host of ?grass,
?Gramineae. Portugal.

coniothyrium (L. Fuckel) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:317.
1875. =Sphaeria coniothyrium L. Fuckel,
Symbolae Mycologicae, p. 115. 1870.

March 1991

=Melanomma coniothyrium (L. Fuckel) L.
Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):56. 1957. On stems of Alnus
glutinosa (L.) Gaertner, Broussonetia sp.,
Chimonanthus sp., Kerria sp., Menispermum
canadense L., Rosa sp., Rubus fruticosus L.,
Salix vitellina L., Sambucus nigra L.,
Betulaceae, Moraceae, Calycanthaceae,
Rosaceae, Menispermaceae, Salicaceae,
Caprifoliaceae. Austria, France, Germany,
Italy, Portugal.

coniothyrium (L. Fuckel) P.A. Saccardo forma
berberidis (C.E. Richon) R. Ciferri, Annales
Mycologici, Berlin 20:51. 1922. =Lepto-
sphaeria berberidis C.E. Richon, Catalogue
Raisonné des Champignons Qui Croissent
Dans le Département de la Marne, p. 300.
1889. On leaves and branchlets of Berberis
vulgaris L., Berberidaceae. France, Italy.

coniothyrium P.A. Saccardo var. foliicola N.N.
Woronichin, Vestnik Tiflisskogo Botanich-
eskogo Sada, Tiflis (Moniteur du Jardin
Botanique de Tiflis) 28:17. 1913. On living
leaves of Laurus sp., Lauraceae. U.S.S.R.
(Caucasus).

conoidea (V. Cesati & G. de Notaris) P.A. Saccardo,
Nuovo Giornale Botanico Italiano e Bolletino
della Societa Botanica Italiana, Firenze 7:314.
1875. =Sphaeria doliolum (C.H. Persoon:
E.M. Fries) V. Cesati & G. de Notaris var.
conoidea G, de Notaris, Memorie della
Accademia delle Scienze di Torino, Series 2,
16:466. 1856. On stems of Angelica sylvestris
L., Umbelliferae. Italy.

conoidea (V. Cesati & G. de Notaris) P.A. Saccardo
forma angelicae F. Fautrey in C.
Roumeguére, Fungi Selecti Gallici Exsiccati,
Century 56, No. 5536. Anno 1891; Revue
Mycologique, Toulouse 13:8. 1891. On dry
stems of Angelica sylvestris L., Umbelliferae.
France.

conoidea (V. Cesati & G. de Notaris) P.A. Saccardo
forma astferis F. Fautrey, Revue Mycologique,
Toulouse 16:112. 1894; Fungi Selecti Gallici
Exsiccati, Century 66, No. 6552. Anno 1894.
On dry stems of Aster salignus Willd.,
Compositae. France.

conoidea (V. Cesati & G. de Notaris) P.A. Saccardo
forma macrospora F. Fautrey, Revue
Mycologique, Toulouse 11:152. 1889. On
rotten stems of Urtica dioica L., Urticaceae.
France.

consessa (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:17.
1883. =Sphaeria (Caulicolae) consessa M.C.
Cooke & J.B. Ellis, Grevillea, London 6:95.
1878. On stems of Helianthus sp., Composi-
tae. U.S.A.

Nomenclator of Leptosphaeria

217

consimilis J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 2:41. 1886. On
dead limbs of Salix sp., Salicaceae. U.S.A.

consobrina P.A. Karsten, Ofversigt af K. Vetenskap-
sakademiens Forhandlingar, Stockholm
29(2):102—103. 1872. =Phaeosphaeria
consobrina (P.A. Karsten) O. Eriksson, Arkiv
for Botanik, Uppsala, Stockholm, Series 2,
6:415. 1967. On dried leaves of Carex pulla
Good., Cyperaceae. Norway.

consocians (W. Nylander) P.A. Saccardo & D.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 17:730. 1905. =Mycoporum
consocians W. Nylander, Flora, Jena und
Regensburg 55:364. 1872. On thalli and
apothecia of Lecidea vernalis (L.) Ach.,
Lichenes (Lecideaceae). U.S.S.R.

consociata H. Rehm, Hedwigia, Dresden 35:149.
1896; Ascomyceten, Fascicle 24, No. 40.
Anno 1896. =Monographella consociata (H.
Rehm) O. Eriksson & J. Yue, Mycotaxon. An
International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens, Ithaca,
New York 38:205. 1990. On leaves of
Chusquea sp., Gramineae. Ecuador.

constricta (G. Bresadola) F. Hazslinszky, Matemati-
kai és Természettudomanyi K6zlemenyek
Vonatkozolag a Hazai Viszonyokra, Budapest
25(2):148. 1892. =Metasphaeria constricta
G. Bresadola, Revue Mycologique, Toulouse
12:185. 1890. On branches of Euonymus
europaeus L., Celastraceae. Hungary.

contecta J. Kohlmeyer, Nova Hedwigia. Zeitschrift
fiir Kryptogramenkunde, Weinheim 6:314.
1963. On rotten wood in marine waters on
unknown host, unknown family. Germany.

controversa (J. Desmaziéres) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:235. 1863.
=Sphaeria controversa J. Desmazieres,
Annales des Sciences Naturelles, Paris,
Botanique, Series 2, 17:102. 1842. =Di-
aporthe controversa (J. Desmazieéres) T.R.J.
Nitschke in L. Fuckel, Symbolae Mycologi-
cae, Erster Nachtrag, p. 319. 1871. =Di-
aporthe eres T.R.J. Nitschke fide L.E.
Wehmeyer, The Genus Diaporthe Nitschke
and Its Segregates, p. 63. 1933. On plant
stems, on dry branches of Acer sp., Fraxinus
sp., Rubus sp., Aceraceae, Oleaceae, Ro-
saceae. France.

convallariae J.V. d’ Almeida & M. de Sousa da
Camara, Boletim da Sociedade Broteniana,
Coimbra 24:164. 1908/1909. Nom. illegit.
Art. 64.1. =Leptosphaeria dracaenae M. de
Sousa da Camara. On dead leaves of
Dracaena draco (L.) L., Agavaceae.
Portugal.

218

convallariae (L. Fuckel) P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Ilustrans 1:38. 1877. =Ple-
ospora convallariae L. Fuckel, Symbolae
Mycologicae, pp. 138-139. 1870. On dry
stems of Convallaria multiflora L., Liliaceae.
France.

convallariae J.V. d’ Almeida & M. de Sousa da
Camara forma dracaenae J.V. d’ Almeida &
M. de Sousa da Camara, Boletim da So-
ciedade Broteriana, Coimbra 24:164. 1908/
1909. On dead leaves of Dracaena draco (L.)
L., Agavaceae. Portugal.

convallariae J.V. d’ Almeida & M. de Sousa da
Camara forma rusci J.V. d’ Almeida & M. de
Sousa da Camara, Boletim da Sociedade
Broteriana, Coimbra 24:164. 1908/1909. On
dead leaves of Ruscus aculeatus L., Rus-
caceae. Portugal.

cookei R. Pirotta, Atti dell’ Istituto Botanico della
Universita e Laboratorio Crittogamica di
Pavia, Milano 2 & 3:162—163. (1877) 1879
[as cookii]. On cortex of living runners on
Vitis vinifera L., Vitaceae. Great Britain.

coorgica K.H. Anahosur, Sydowia. Annales
Mycologici, Horn, N.O. 24(1-6):286-287.
(1970) 1971. On living twigs of Pteridium
aquilinum (L.) Kuhn, Dennstaedtiaceae.
India.

corae N. Patouillard, Journal de Botanique, Paris
2:150. 1888. On thallus of Cora pavonia
(Swartz) Fries (=Dictyonema), Lichenes
(Thelephoraceae). Central America.

corallorhizae C.H. Peck, Report. New York State
Museum of Natural History, Albany 38:105.
1885. On dead stems of Corallorrhiza
multiflora Nutt., Orchidaceae. U.S.A.

cordylines (C.L. Spegazzini) P.A. Saccardo & A.
Trotter in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 22:231. 1913. [Incorrectly cited as a
comb. nov. based on Leptosphaerella
cordylines C.L. Spegazzini. Nom. inval. Art.
43.1.] On dry leaves of Cordyline dracaenoi-
des Kunth, Agavaceae. Argentina.

cornuta E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 7(1-4):273-274. 1953. On dead
stems of Laserpitium siler L., Umbelliferae.
Switzerland.

coronillae G. Moesz, Botanikai K6zlemények,
Budapest 28:164. 1931. On dead stems of
Coronilla sp. [as C. comata L.], Leguminosae.
Hungary.

corrugans H. Rehm, Osterreichische Botanische
Zeitschrift, Wien 54:84. 1904. On living
leaves of Cytisus alpinus Miller, Legumino-
sae. Yugoslavia.

corticola (L. Fuckel) P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 1:342. 1878; Fungi

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Italici autographice delineati (additis nonnullis
extra-italicis asterisco notatis), Patavii, Table
288. 1878. =Sphaeria corticola L. Fuckel,
Symbolae Mycologicae, p. 114. 1870.
=Metasphaeria corticola (L. Fuckel) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:166. 1883. =Pleosphaerulina corticola (L.
Fuckel) H. Rehm, Annales Mycologici, Berlin
10:538. 1912. =Clathridium corticola (L.
Fuckel) R.A. Shoemaker & E. Miiller,
Canadian Journal of Botany, Ottawa 42:404.
1964. =Griphosphaeria corticola (L. Fuckel)
F. v. Hohnel, Annales Mycologici, Berlin
16:87. 1918. =Discostroma corticola (L.
Fuckel) I. Brockmann, Sydowia. Annales
Mycologici, Horn, N.O. 28:313. (1975) 1976.
On cortex of branchlets, on dry stems of
Cornus sanguinea L., Lonicera caprifolium
L., Prunus baccata Borkh., Prunus domestica
L., Prunus spinosa L., Rosa sp., Caprifoli-
aceae, Rosaceae. Austria, Canada, Czecho-
slovakia, Germany, Hungary, Italy, Switzer-
land.

corvina (E. Rostrup) J. Lind, Danish Fungi as
Represented in the Herbarium of E. Rostrup,
Nordisk Forlag, Copenhagen, p. 217. 1913.
=Metasphaeria corvina E. Rostrup, Dansk
Botanisk Forening, Copenhagen, Meddelsher
2(4):90-91. 1888. On rotten feathers of
Corvus cornix [Animalia], Corvidae.
Denmark.

corynispora A.N. Berlese & G. Bresadola, Annuario
della societa Degli Alpinisti Tridentini
14:328-329. 1889; Micromycetes Tridentini
Contribuzione Allo Studio dei Funghi
Microscopici del Trentino, pp. 32-33. 1889.
On dead stems of Cirsium lanceolatum (L.)
Scop., Compositae. Italy.

cosmicola A. Chiplonkar, Sydowia. Annales
Mycologici, Horn, N.O. 22(1-4):274-275.
(1968) 1969 [as cosmosicola}]. On dried stems
of Cosmos sulphureus Cav., Compositae.
India.

coumarounae R. Gonzalez Fragoso & R. Ciferri,
Boletin de la R. Sociedad Espanola de

_ Historia Natural, Madrid 27:326—327. 1927.

Nom. nud. Art. 34.1. On living leaves of
Coumarouna punctata S.F. Blake, Legumino-
sae. Dominican Republic.

coumarounae R. Gonzalez Fragoso & R. Ciferri,
Publicaciones Estacion Agronémica de Moca,
Santa Domingo. Series B, Botanica, No.
11:22. 1928. Nom. inval. Art. 34.1. On living
leaves of Coumarouna punctata S.F. Blake,
Leguminosae. Dominican Republic.

crastophila (P.A. Saccardo) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:321.
1875. =Sphaerella crastophila P.A. Sac-

March 1991

cardo, Atti dell’ Accademia Scientifica
Veneto-Trentino-Istriana, Padova 2(2):142.
1873. On leaves of Festuca sp., Gramineae.
Italy.

crastophila P.A. Saccardo forma tofieldiae P.A.
Saccardo in G. Bresadola and P.A. Saccardo,
Malpighia. Rassegna Mensuale di Botanica.
Messina, Genova 11:289. 1897. On stems and
leaves of Tofieldia calyculata (L.) Wahlenb.,
Liliaceae. Italy.

crepini (G.D. Westendorp) G. de Notaris, Commen-
tario della Societa Crittogamologica Italiana,
Milan 2:486. 1867. =Sphaeria crepini G.D.
Westendorp, Bulletins de L’Académie Royale
des Sciences, des Lettres et des Beaux-Arts de
Belgique, Brussels, Series 2, 7:88. 1859. On
bracteoles of spikes on Lycopodium an-
notinum L., Lycopodiaceae. Belgium, Italy.

crozalsiana R.C. Maire, Annales Mycologici, Berlin
11:352. 1913. On dead branches of Tamarix
sp., Tamaricaceae. Algeria.

crozalsii L. Vouaux, Bulletin de la Société Mycolo-
gique de France, Paris 29:120. 1913 [as
crozalsi}. On thallus of Aspicilia calcarea
(L.) Korb [=Lecanora calcarea (L.)
Sommerf.], Placodium teicholytum (Ach.)
DC., Lichenes (Teloschistaceae). France.

crucheti E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):255-256. 1950. =Mera-
sphaeria cladii D. Cruchet, Bulletin de la
Société Vaudoise des Sciences Naturelles,
Lausanne 55:166—167. 1923. Non Lepto-
sphaeria cladii D. Cruchet, 1923. On
branches of Cladium mariscus (L.) Pohl,
Araceae. Switzerland.

cruenta P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:318-319. 1881. On dead stems of
Thalictrum flavum L., Ranunculaceae.
France.

crustacea J. Schroter in F.J. Cohn, Kryptogamen-
Flora Von Schlesien. Im Namen Der Schlesis-
chen Gesellschaft fiir vaterlandische Cultur
herausgegeben von Prof. Dr. Ferdinand Cohn,
Secretair der Botanischen Section, Breslau
3(2):365. 1894. On dead stems of Artemisia
sp., Compositae. Poland.

cryptica H. Sydow, Annales Mycologici, Berlin
28:85-86. 1930. Parasitic on stromata of
Phyllachora sp., Fungi (Phyllachoraceae).
Venezuela.

cucurbitae L. Montemartini, Rivista di Patologia
Vegetale, Padova 8:185. 1916. On leaves of
Cucurbita pepo L., Cucurbitaceae. Italy.

cucurbitarioides J.H. Fabre, Annales des Sciences
Naturelles, Paris, Botanique, Series 6, 15:50.
1883. On more or less dry stems of Do-
rycnium suffruticosum Vill., Leguminosae.
France.

Nomenclator of Leptosphaeria

219

culmicola (E.M. Fries:E.M. Fries) G.L. Rabenhorst,
Botanische Zeitung, Berlin & Leipzig 24:412.
1866. =Sphaeria culmicola E.M. Fries:E.M.
Fries, Systema Mycologicum Sistens
Fungorum 2:430. 1823. On grass culms of
unknown host of Gramineae, Gramineae.
Italy.

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten,
Mycologia Fennica Pars 2, Pyrenomycetes,
p. 108. 1873. Nom. illegit. Art. 64.1.
=Sphaeria culmicola E.M. Fries:E.M. Fries,
Systema Mycologicum Fungorum 2:430.
1823. On dried culms of unknown host of
Gramineae, Gramineae. Italy.

culmicola (E.M. Fries) P.A. Karsten forma major
P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:323. 1875. On stems of
grass, Gramineae. Italy.

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten
forma melicae F. Fautrey in C. Roumeguére,
Revue Mycologique, Toulouse 17:176. 1895;
Fungi Selecti Gallici Exsiccati, Century 69,
No. 6847. Anno 1895. On sheaths of dry
straw of Melica altissima L., Gramineae.
France?

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten var.
aquatica P.A. Saccardo, Michelia Commen-
tarium Mycologicum Fungos in Primis
Italicos Illustrans 2:319. 1881. On reeds of
Scirpus lacustris L., Cyperaceae. France.

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten var.
hispalensis R. Gonzalez Fragoso, Trabajos del
Museo Nacional de Ciencias Naturales,
Madrid, Series Botanica 10:87. 1916. On
leaves and sheaths of Phragmites communis
Trin., Gramineae. Spain.

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten var.
minor P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:322—323. 1875.
On stems of grass, Gramineae. Italy.

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten var.
nigrans (M.R. Roberge) P.A. Karsten,
Mycologia Fennica Pars 2, Pyrenomycetes,
pp. 108-109. 1873. =Sphaeria (Caulicola)
nigrans M.R. Roberge in J. Desmaziéres,
Annales des Sciences Naturelles, Paris,
Botanique, Series 3, 6:79. 1846. =Lepto-
sphaeria nigrans (M.R. Roberge) V. Cesati &
G. de Notaris. On sheath of Dactylis
glomerata L., Gramineae. Finland, France.

culmicola (E.M. Fries:E.M. Fries) P.A. Karsten var.
rhizomatum P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 2:319. 1881. On
rhizome of Phragmites sp., Gramineae.
France.

culmifida P.A. Karsten, Fungi Fenniae Exsiccati,
Century 10, No. 961. Anno 1870; Mycologia
Fennica 2:103. 1873. =Metasphaeria

220

culmifida (P.A. Karsten) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:174.
1883. =Trichometasphaeria culmifida (P.A.
Karsten) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):140. 1957. On
culms of Phleum pratense L., Gramineae.
Finland.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:235. 1863.
=Sphaeria culmifraga E.M. Fries:E.M. Fries,
Systema Mycologicum Sistens Fungorum
2:510. 1823. =Pleospora culmifraga (E.M.
Fries:E.M. Fries) L. Fuckel, Symbolae
Mycologicae, p. 137. 1870. On culm of grass
of unknown host of Gramineae, Gramineae.
Italy, Sweden.

culmifraga P.A. Saccardo, Mycotheca Veneta,
sistens Fungos Venetos Exsiccatos, Century 1,
No. 77. Anno 1875. Nom. nud. Art. 32.1 [as
(Fr.) Ces. et DN.]. =Leptosphaeria amphi-
bola P.A. Saccardo. On culms of Sorghum
vulgare Pers., Gramineae. Italy.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris forma majuscula H. Rehm,
Ascomyceten, Fascicle 4, No. 195. Anno
1873. Non vidi. Hedwigia, Dresden 24:67.
1885. On fallen culms of unknown host,
Gramineae. Unknown country.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris forma minuscula H. Rehm,
Hedwigia, Dresden 24:66—67. 1885.
=Phaeosphaeria minuscula (H. Rehm) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1545. 1989. On unknown substrate
of unknown host, unknown family. Canada,
Sweden.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris forma poae F. Fautrey in C.
Roumeguére, Fungi Selecti Gallici Exsiccati,
Century 58, No. 5755. Anno 1891; Revue
Mycologique, Toulouse 13:129. 1891. On
rachis of panicles of old straw of Poa sudetica
Haenke, Gramineae. France.

culmifraga (E.M. Fries:E.M. Fries) var. alpestris H.
Rehm, Osterreichische Botanische Zeitschrift,
Wien 56:296. 1906. Nom. nud. Art. 32.1. On
grass blade of unknown host, Gramineae.
Austria.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris var. bromicola G. Bresadola,
Revue Mycologique, Toulouse 12:184. 1890.
On stems of Bromus asper Murray, Grami-
neae. Hungary.

culmifraga (E.M. Fries:E.M. Fries) P.A. Karsten var.
herpotrichoides (G. de Notaris) P.A. Karsten,
Mycologia Fennica 2:108. 1873; Bidrag till
Kannedom om Finlands Natur och Folk.
Utgifna af Finska Vetenskaps-Societeten,

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Helsingfors 23:108. 1873. =Leptosphaeria
herpotrichoides G. de Notaris.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris var. linearis P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 2:319.
1881. On stems of Brachypodium sp.,
Gramineae. France.

culmifraga (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris var. propingua P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 2:319.
1881. On leaves of Poa aquatica L.,
Gramineae. France.

culmorum B. Auerswald in H. Rehm, Bericht des
Naturwissenschaftlichen Vereins fiir Schwa-
ben und Neuberg, Augsburg 26:60. 1881.
=Scleropleella culmorum (B. Auerswald) F. v.
Hohnel, Berichte der Deutschen Botanischen
Gesellschaft, Berlin 36:135—140. 1918.
[Falsely so cited.] =Nodulosphaeria
culmorum (B. Auerswald) G. Moesz, Magyar
tudomanyos akadémia Balkan-kutatasainak
Tudomanyos Eredményei, Budapest 3:140.
1926. =Phaeosphaeria microscopica (P.A.
Karsten) O. Eriksson var. culmorum (B.
Auerswald) O. Eriksson, Arkiv for Botanik.
Uppsala, Stockholm, Series 2, 6:427. 1967.
=Phaeosphaeria culmorum (B. Auerswald) A.
Leuchtmann, Sydowia. Annales Mycologici,
Horn, N.O. 37:113. 1984. On sheaths and
leaves of unknown host of grass, Gramineae.
Canada, Czechoslovakia, Denmark, Finland,
Germany, Italy, Sweden, Switzerland, U.S.A.,
WESSERE

culmorum B. Auerswald forma apogon (P.A.
Saccardo & C.L. Spegazzini) H. Rehm,
Ascomyceten, Fascicle 15, No. 734. Anno
1883; Hedwigia, Dresden 23:71—72. 1884.
On unknown substrate on unknown host,
unknown family. Unknown country.

culmorum B. Auerswald forma epigeii A.I. Lobik,
Materialy po Floristicheskim i Faunistich-
eskim Obsledovaniyam Terskogo Okruga
{Data from Investigations on the Flora and
Fauna of the Ter Region], p. 23. 1928. On
leaves of Calamogrostis epigejos (L.) Roth,
Gramineae. U.S.S.R.

culmorum B. Auerswald forma /ungarica H. Rehm,
Ascomyceten, Fascicle 19, No. 941. Anno
1888; Hedwigia, Dresden 27:173. 1888. On
dry culms of Luzula albida (Hoffm.) DC.,
Juncaceae. Hungary.

culmorum B. Auerswald forma phragmitis A.1.
Lobik, Material po Floristicheskim i
Faunisticheskim Obsledovaniyam Terskogo
Okruga [Data from Investigations on the Flora
and Fauna of the Ter Region], p. 24. 1928.
On leaves of Phragmites communis Trin.,
Gramineae. U.S.S.R.

March 1991

culmorum B. Auerswald var. flavobrunnea J.
Feltgen, Vorstudien zu einer Pilz-Flora des

Grossherzothums, Luxemburg 1(3):205. 1903.

On dry culms of Glyceria spectabilis Mert. &
Koch (or Phalaris sp.?), Gramineae. Luxem-
bourg.

culmorum B. Auerswald var. paleicola P.C.
Hennings, Verhandlungen des Botanischen
Vereins Der Provinz Brandenburg, Berlin
44:177-178. 1903. On culms of Carex
leporina L., Cyperaceae. Germany.

cumana P.A. Saccardo & C.L. Spegazzini in P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
1:394-395. 1878. =Metasphaeria cumana
(P.A. Saccardo & C.L. Spegazzini) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:177. 1883. On dead leaves of Carex sp.,
Cyperaceae. Italy.

cumulata W. Kirschstein, Verhandlungen des
Botanischen Vereins Der Provinz Bran-
denburg, Berlin 48:56. (1906) 1907. On
fallen culms of Phragmites sp., Gramineae.
Germany.

curta P.A. Saccardo & Abbé Flageolet in P.A.
Saccardo, Bulletin de la Société Mycologique
de France, Paris 12:66. 1896. On dead stems
of Rhus typhina L., Anacardiaceae. France.

cycadis M. Svréek, Ceska Mykologie, Praha 34:177.
1980. On upper surface of living leaves of
Cycas revoluta Thunb., Cycadaceae. Iran.

cylindrospora B. Auerswald & G. Niessl v. Mayer-
dorf in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:37. 1883. On dead stems of
Epilobium angustifolium L., Onagraceae.
Czechoslovakia.

cylindrospora A.M. Saccas, Etude de la Flore
Cryptogamique des Caféiers en Afrique
Centrale. Bulletin de Institut Frangais du Café
du Cacao et d’ Autres Plantes Stimulantes
(Bulletin IFCC No. 16), pp. 237-239. 1981.
Nom. inval. Art. 37.1. On dead branches of
Coffea robusta L. Linden (=Coffea canephora
Pierre ex Froehn.), Rubiaceae. Central
African Republic.

cylindrostoma K. Starback, Arkiv for Botanik,
Uppsala, Stockholm 5(7):23. 1905. On
unidentified ?stems of unknown host,
unknown family. Argentina.

cynaracearum B. Auerswald & G. Niessl v.
Mayendorf in G. Niessl v. Mayendorf, Ver-
handlungen des Naturforschenden Vereins in
Briinn 10:174. (1871) 1872. =Metasphaeria
cynaracearum (B. Auerswald & G. Niessl v.
Mayendorf) P.A. Saccardo, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:170. 1883. On dry
leaves of Carlina acaulis L., Compositae.
Czechoslovakia.

Nomenclator of Leptosphaeria 22

cynodontis-dactyli E. Marchal & R.L. Steyaert,
Bulletin. Société Royale de Botanique de
Belgique, Bruxelles 61(Series 2, 12):161.
1929. On living leaves of Cynodon dactylon
(L.) Pers., Gramineae. Zaire.

cynoglossi L. Hollos, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 5:455.
1907. On dried stems of Cynoglossum
officinale L., Boraginaceae. Hungary.

cynops J.H.C. Fabre, Annales des Sciences
Naturelles, Paris, Botanique, Series 6,
15:50-51. 1883. On dead stems of Plantago
cynops L., Plantaginaceae. France.

cynosuri L.M. Unamuno Yrigoyen, Anales del
Jardin Botanico de Madrid 4:152. 1944. On
culms of Cynosurus cristatus L., Gramineae.
Spain.

cyperi L. Hollés, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:31. 1926. On dry
leaves of Cyperus flavescens L., Cyperaceae.
Hungary.

cypericola L. Hollos, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:32. 1926. On dry
leaves of Cyperus flavescens L., Cyperaceae.
Hungary.

cyperina G. Passerini, Erbario Crittogamico Italiano,
Series 2, Fascicle 22, No. 1074. Anno 1881.
=Phaeosphaeria cyperina (G. Passerini) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1540. 1989. On stubble of
Cyperus monti L., Cyperaceae. Italy.

dactylina G. Passerini, Rendiconti della Sedute della
R. Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:91. 1887. On dry culms of
Dactylis glomerata L., Gramineae. Italy.

danica A.N. Berlese, cones Fungorum Omnium
Hucusque Cognitorum ad usum Sylloges
Saccardianae Adcommodatae 1:87. 1892.
=Leptosphaeria marina E. Rostrup. Non
Leptosphaeria marina J.B. Ellis & B.M.
Everhart. =Didymosphaeria danica (A.N.
Berlese) ILM. Wilson & J.M. Knoyle,
Transactions of the British Mycological
Society, London 44:57. 1961. =Lautitia
danica (A.N. Berlese) S. Schatz, Canadian
Journal of Botany, Ottawa 62:31. 1984.
Parasitic on Chondrus crispus J. Stackhouse,
Algae (Gigartinaceae). Denmark.

daphnes M.R. de Sousa Dias & M. de Sousa da
Camara, Agronomia Lusitana, Sacavém
15:23-24. 1953. On branchlets of Daphne
gnidium L., Thymelaeaceae. Portugal.

daphniphylli C.T. Dzhalagoniya, Trudy Sukhum-
skogo Botanicheskogo Sada, Sukhum 15:118.
1964 [as daphniphylliae]. On living leaves of
Daphniphyllum macropodium Mig., Daph-
niphyllaceae. U.S.S.R.

darkeri R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2699-2700. 1984. On

222 Illinois Natural History Survey Bulletin

stems of Agastache urticifolia (Benth.) Ktze.,
Labiatae. U.S.A.

dasylirii (G.L. Rabenhorst) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:67-68. 1883.
=Sphaeria dasylirii G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbaril vivi
Mycologici Continuatio Edita Nova, Series 2,
Century 27, No. 2655. Anno 1881; Hedwigia,
Dresden 21:9. 1882. On leaves of Dasylirion
junceum Zucc., Agavaceae. Italy.

davidii S.A. Gucevicz, Trudy Gosudarstvennogo
Nikitskogo Botanicheskogo Sada, Yalta
29:191. 1959. On dried stems of Buddleja
davidii Franch., Loganiaceae. U.S.S.R.

daviesiae F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 8(1-6):197. 1954. On dry
branches of Daviesia latifolia R. Br.,
Leguminosae. Australia.

davisiana F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 7(1-4):20. 1953. On dry stems of
Scutellaria brevibracteata Stapf, Labiatae.
Turkey.

dearnessti R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2700. 1984. On stems of
Asclepias syriaca L., Asclepiadaceae. Canada.

debeauxii C. Roumeguére & P.A. Saccardo in C.
Roumeguére, Revue Mycologique, Toulouse
2:188. 1880. =Metasphaeria debeauxii (P.A.
Saccardo) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:182. 1883. On costa of leaves of
Chamaerops humilis L., Palmae. Algeria.

decaisneana (L. Crié) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:50. 1883. =Depazea
caisneana L. Crié, Annales des Sciences
Naturelles, Paris, Botanique, Series 6, 7:48.
1878. On dry leaves of Populus sp., Sali-
caceae. France.

deficiens F. Tassi, Bolletino del R. Orto Botanico,
Siena 3:121. 1900. On stems of Tellima
grandiflora (Pursh) Douglas ex Lindley,
Saxifragaceae. China.

defodiens (J.B. Ellis) J.B. Ellis in N.L. Britton,
Catalogue of Plants Found in New Jersey.
Geological Survey of New Jersey, Final
Report of the State Geologist, Trenton
2(1):525. 1889. =Sphaeria (Leptosphaeria)
defodiens J.B. Ellis, Bulletin of the Torrey
Botanical Club (and Torreya), New York
8:90. 1881. On dead plant of Juncus effusus
L., Juncaceae. U.S.A. (New York).

delawayi N.T. Patouillard, Revue Mycologique,
Toulouse 8:82. 1886. On dead stems of
Primula sikkimensis Hook., Primulaceae.
China.

dematiicola W. Kirschstein, Annales Mycologici,
Berlin 34:188. 1936. On old fruiting stem of
Typha angustifolia L., Typhaceae. Germany.

dematium F.A. Hazslinszky, Matematikai és
Természettudomanyi K6zlemenyek Von-

Vol. 34 Art. 3

atkozolag a Hazai Viszonyokra, Budapest
25(2):43. 1892; Magyarorszag s
Tarsorszagainak Sphaeriai, Budapest, p. 107.
1892. On dead stems of Hypericum perfora-
tum L., Guttiferae. Hungary.

demissa G. Niessl v. Mayendorf, Instituto. Revista
Scientifica e Litteraria, Coimbra 31:89-90.
1883. On dry stems of Vinca media
Hoffmanns. & Link, Apocynaceae. Portugal.

dennisiana (A. Leuchtmann) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1578.
1989. =Phaeosphaeria dennisiana A.
Leuchtmann, Sydowia. Annales Mycologici,
Horn, N.O. 37:155-156. 1984. On dry leaves
of Minuartia sedoides (L.) Hiern., Caryophyl-
laceae. Great Britain.

densa J. Bresadola, Hedwigia, Dresden 35:199.
1896. On leaves of Acorus calamus L.,
Araceae. Germany.

depressa (L. Fuckel) H.G. Winter, Dr. L.
Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):473. 1885. =Sphaeria
depressa L. Fuckel, Symbolae Mycologicae,
p. 115. 1870. =Metasphaeria depressa (L.
Fuckel) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:166. 1883. On dry stems of
Carpinus sp., Betulaceae. Austria.

depressa L.E. Wehmeyer & S. Ahmad, Biologia.
Biological Society of Pakistan, Lahore 10:12.
1964. Nom. illegit. Art. 64.1. [A later
homonym of Leptosphaeria depressa T.
Petch. 1926.] On stems of Euphorbia sp. (as
E. oxyroidea), Euphorbiaceae. Pakistan.

depressa T. Petch, Ceylon Journal of Science,
Colombo, Section A (Botany) 10:135. 1926.
On stems of Camellia theifera Griff.,
Theaceae. Sri Lanka.

derasa (M.J. Berkeley & C.E. Broome) B. Auers-
wald, Botanischer Tauschverein in Wien, p. 4.
1886. Non vidi. =Sphaeria derasa M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 9:328. 1852. =Nodulosphaeria derasa
(M.J. Berkeley & C.E. Broome) L. Holm,
Symbolae Botanicae Upsalienses 14(3):89.
1957. On unknown substrate of Senecio
Jacobaea L., Compositae. Great Britain.

derasa (M.J. Berkeley & C.E. Broome) F. v.
Thiimen, Mycotheca Universalis, Century 3,
No. 269. Anno 1875. Nom. illegit. Art. 64.1.
=Sphaeria derasa M.J. Berkeley & C.E.
Broome, Annals and Magazine of Natural
History, London, Series 2, 9:328. 1852.
=Nodulosphaeria derasa (M.J. Berkeley &
C.E. Broome) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):89. 1957. On
unknown substrate of Senecio jacobaea L..
Compositae. Great Britain.

derasa (M.J. Berkeley & C.E. Broome) F. v.
Thiimen forma alpestris H. Rehm, Hedwigia,

March 1991

Dresden 24:235. 1885. On stems of unknown
host of composite family, Compositae. Italy.

derasa (M.J. Berkeley & C.E. Broome) F. v.
Thiimen forma macrospora F. Fautrey in C.
Roumeguére, Revue Mycologique, Toulouse
16:8. 1894; Fungi Selecti Gallici Exsiccati,
Century 65, No. 6436. Anno 1894. On dry
branches of Sambucus ebulus L., Caprifoli-
aceae. France?

(Pocosphaeria) derasa (M.J. Berkeley & C.E.
Broome) B. Auerswald forma robusta P.
Strasser, Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien 57:315.
1907. =Leptosphaeria robusta (P. Strasser)
E. Miller. =Nodulosphaeria robusta (P.
Strasser) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):90. 1957.
=Ophiobolus robustus (P. Strasser) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 54:2389. 1976. On dry stems of
Senecio nemorensis L., Compositae. Austria.

derasa (M.J. Berkeley & C.E. Broome) B. Auers-
wald var. franconica F. Petrak, Kryptoga-
mische Forschungen, Miinchen 2:162—163.
1931. =Leptosphaeria franconica (F. Petrak)
E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):298-299. 1950. On dry
stems of /nula salicina L., Compositae.
Romania.

desciscens C.A. Oudemans, Beihefte zum Botani-
schen Zentralblatt, Cassel 11:527. 1902. On
stems of unknown host, unknown family.
Netherlands.

desmonci H. Sydow & P. Sydow, Hedwigia,
Dresden 49:79. 1909. On branches of
Desmoncus sp., Palmae. Brazil.

(Leptosphaerella) diana P.A. Saccardo & C.L.
Spegazzini in P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Ilustrans 1:398—399. 1878.
=Sphaerulina (Leptosphaerella) diana (P.A.
Saccardo & C.L. Spegazzini) M.C. Cooke,
Grevillea, London 18:80. 1890. =Mycotodea
diana (P.A. Saccardo & C.L. Spegazzini) W.
Kirschstein in O.C. Schmidt, Krypto-
gamenflora der Mark Brandenburg und
Angrenzender Gebiete herausgegeben von
dem Botanishen Verein der Provinz Bran-
denburg, Leipzig 7:437. 1938. On rotten
leaves of Acer negundo L., Aceraceae. Italy.

dianthi (E. Rostrup) J. Lind, Danish Fungi as
Represented in the Herbarium of E. Rostrup,

=Metasphaeria dianthi E. Rostrup, Botanisk
Tidsskrift, Kjobenhavn 26:311. 1905. On
stems of Dianthus superbus L., Caryophyl-
laceae. Denmark.

dianthi A. Christow, Bulgaria. Ministerstvo na
zemledielieto i durzhuvnitie imoti. Zemlesto-
panska biblioteka (Ministerium fiir Land-
wirtschaft und Staatsdomanen Land-

Nomenclator of Leptosphaeria

tN
ho
Ww

wirtschaftsbiblothek) 43:4—6. 1931. Nom.
illegit. Art. 64.1. On unknown substrate of
Dianthus sp., Caryophyllaceae. Bulgaria.

dianthi L. Hollés, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkoz6lag a Hazai
Viszonyokra (Maguar tudomanyos akadémia
Budapest. Matematikai és természettudo-
manyi bizottsag), Budapest 35:32. 1926.
Nom. illegit. Art. 64.1. On dry stems of
Dianthus caryophyllus L., Caryophyllaceae.
Hungary.

dianthi N. Golovina, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 12:156. 1959. Nom.
illegit. Art. 64.1. On dead stems of Dianthus
sp., Caryophyllaceae. U.S.S.R. (Uzbek).

diaporthoides H.G. Winter, Boletim da Sociedade
Broteriana, Coimbra 3:56. (1884) 1885. On
dried stems of unknown host of Umbelliferae,
Umbelliferae. Brazil, Portugal.

dichosciadii F. Petrak, Sydowia. Annales Mycol-
ogici, Horn, N.O. 9(1-6):561. 1955. On
decaying leaves of Dichosciadium ranuncu-
laceum (F. v. Miiller) Domin, Hydrocot-
ylaceae. Australia.

dichroa G. Passerini, Rendiconti della Sedute della
R. Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:90. 1887. =Passeriniella
dichroa (G. Passerini) A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:51. 1890. On dry branches of Deutzia
scabra Thunb., Saxifragaceae. Italy.

didymellae-vincetoxici E. Miiller, Sydowia. Annales
Mycologici, Horn, N.O. 5(3—6):51-52. 1951.
On fruiting bodies on Didymella vincetoxici
(de Not.) Sacc., Fungi (Phaeosphaeriaceae).
Switzerland.

digitalis (P.L. Crouan & H.M. Crouan) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:85. 1883. =Sphaeria digitalis P.L. Crouan
& H.M. Crouan, Florule Du Finistére
Contenant Les Descriptions De 360 Espeéces
Nouvelles De Sporogames, De Nombreuses
Observations et une synonymie des plantes
Cellulaires et Vasculaires Qui Croissent
Spontanément Dans ce Département, p. 28.
1867. On dead stems of Digitalis sp.,
Scrophulariaceae. France.

dioica (E.M. Fries:E.M. Fries) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:18.
1883 [as (Moug.) Sacc.]. =Sphaeria dioica
E.M. Fries:E.M. Fries, Elenchus Fungorum
Sistens Commentarium in Systema Mycol-
ogicum 2:82. 1828. On branches of Acer
pseudoplatanus L., Spartium sp., Aceraceae,
Leguminosae. France, Great Britain.

224 Illinois Natural History Survey Bulletin

discors P.A. Saccardo & J.B. Ellis in P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 2:567.
1882. =Metasphaeria discors (P.A. Saccardo
& J.B. Ellis) P.A. Saccardo, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:173. 1883. =Pas-
seriniella discors (P.A. Saccardo & J.B. Ellis)
A.E. Apinis & C.G.C. Chesters, Transactions
of the British Mycological Society, London
47:432. 1964. On culms of Spartina sp.,
Gramineae. U.S.A. (New Jersey).

disseminata G. de Notaris, Commentario della
Societa Crittogamologica Italiana, Milan
2:486. 1867. On unknown substrate of
unknown host, Gramineae. Italy.

dissiliens (M.C. Cooke & J.B. Ellis) J.B. Ellis in
N.L. Britton, Catalogue of Plants Found in
New Jersey. Geological Survey of New
Jersey, Final Report of the State Geologist,
Trenton 2(1):525. 1889. =Sphaeria (Caulico-
lae) dissiliens M.C. Cooke & J.B. Ellis,
Grevillea, London 5:51. 1876. On dead stems
of Desmodium sp., Leguminosae. U.S.A.
(New Jersey).

distributa (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:18.
1883. =Sphaeria (Obtectae) distributa M.C.
Cooke & J.B. Ellis, Grevillea, London 7:41.
1878. On small twigs of Desmodium sp.,
Leguminosae. U.S.A. (New Jersey).

dobrogica T. Savulescu & C. Sandu-Ville, Hed-
wigia, Dresden 75:170. 1935. On dead
branches of Smilax excelsa L., Liliaceae.
Romania.

dodonaeae A. Canonaco, Bollettino di Studi e
Informazioni. R. Giardino Botanico (Coloni-
ale) Palermo 14:extr. p. 12. 1936; 14:20.
1937. [Note extract was published in 1936,
which is date of publication for the species. ]
On branches of Dodonaea viscosa (L.) Jacq.,
Sapindaceae. Turkey.

dolioloides (B. Auerswald) P.A. Karsten, Mycologia
Fennica Pars 2, Pyrenomycetes, pp. 106-107.
1873. =Nodulosphaeria dolioloides B.
Auerswald in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi
Mycologici Continuatio Edita Nova, Series
Secunda, Century 6, No. 547. 1863. =Ple-
ospora dolioloides (B. Auerswald) L. Fuckel,
Symbolae Mycologicae, p. 138. 1870. On
stems of Achillea millefolium auct., Anthemis
tinctoria L., Chrysanthemum vulgare (L.)
Bernh., Senecio vulgaris L., Tanacetum
vulgare L., Compositae. Finland, France,
Germany, Portugal, Spain.

dolioloides (B. Auerswald) B. Auerswald, Botanis-
cher Tauschvereinin Wien, p. 4. 1866. Non
vidi. =Nodulosphaeria dolioloides B.

Vol. 34 Art. 3

Auerswald in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi
Mycologici Continuatio, Edition 3, Century 6,
No. 547. Anno 1863. =Pleospora dolioloides
(B. Auerswald) L. Fuckel, Symbolae
Mycologicae, p. 138. 1870.

dolioloides (B. Auerswald) P.A. Karsten, Fungi
Fenniae Exsiccati, Century 10, No. 978. Anno
1870. =Nodulosphaeria dolioloides B.
Auerswald in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi
Mycologici Continuatio, Edition 3, Century 6,
No. 547. Anno 1863. =Pleospora dolioloides
(B. Auerswald) L. Fuckel, Symbolae
Mycologicae, p. 138. 1870. On dry stems of
Achillea millefolium auct., Anthemis tinctoria
L., Chrysanthemum vulgare (L.) Bernh.,
Senecio vulgaris L., Tanacetum vulgare L..,
Compositae. Finland, Germany.

dolioloides (B. Auerswald) P.A. Karsten var. cirsii
P.A. Karsten, Hedwigia, Dresden 23:4. 1884.
On dead stems of Cirsium sp., Compositae.
Finland.

dolioloides (B. Auerswald) P.A. Karsten var. inops
P.A. Karsten, Hedwigia, Dresden 23:4. 1884.
On dead stems of Anthriscus sylvestris (L.)
Hoffm., Celastraceae. Finland.

dolioloides (B. Auerswald) P.A. Karsten var. lathyri
J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(2):164—165.
1901. On dry stems of Lathyrus niger (L.)
Bernh., Leguminosae. Luxembourg.

dolioloides (B. Auerswald) P.A. Karsten var.
rhinanthi J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(2):165. 1901. On dry stems of Rhinanthus
sp., Scrophulariaceae. Luxembourg.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:234—235.
1863. =Sphaeria doliolum C.H. Persoon,
Icones et Descriptiones Fungorum Minus
Cognitorum, p. 39. 1800; E.M. Fries, Systema
Mycologicum Sistens Fungorum 2:509. 1823.
=Bilimbiospora doliolum B. Auerswald in
G.L. Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio, Ausgabe III, Century 3, No. 261. Anno
1860. Nom. rejicem. [Bilimbiospora
doliolum is the same as Leptosphaeria
doliolum but is based on different types.]
=Pleospora doliolum (C.H. Persoon:E.M.
Fries) L. Tulasne & C. Tulasne, Selecta
Fungorum Carpologica 2:276. 1863.
=Cryptosphaeria doliolum (C.H. Persoon)
R.K. Greville, Scottish Cryptogamic Flora 4,
Table 239. 1826. On stems of Achillea sp..
Ambrosia sp., Anaphalis sp., Arctium sp.,
Aster sp., Cirsium sp., Erigeron sp., Eupato-
rium sp., Helianthus sp., Lactuca sp..

March 1991

Polymmis sp., Senecio sp., Solidago sp.,
Sonchus sp. (Compositae); Aralia sp.
(Araliaceae); Apocynum sp., Asclepias sp.
(Asclepiadaceae); Potentilla sp., Rubus sp.
(Rosaceae); Dasystoma sp., Gerardia sp.,
Veronica sp. (Scrophulariaceae); Smilax sp.
(Smilacaceae); Heracleum sp., Pastinaca sp.
(Umbelliferae); Urtica sp. (Urticaceae),
Araliaceae, Asclepiadaceae, Compositae,
Rosaceae, Scrophulariaceae, Smilacaceae,
Umbelliferae, Urticaceae. Belgium, Canada,
Finland, France, Germany, Great Britain,
Italy, Sweden, U.S.A.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris forma carlinae-vulgaris C. Sandu-
Ville, Academia Republicii Populare Romine.
Filiala Iasi. Studii si Cercetari Stiintifice.
Biologie si Stiinte Agricole 9:258. 1964;
Studii §i Cercetari de Biologie, Bucuresti,
Series Botanica 16:296. 1964. On dead stems
of Carlina vulgaris L., Compositae.
Romania.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris forma carlinae-vulgaris C. Sandu-
Ville, Ciuperci Pyrenomycetes-Sphaeriales

Din Romania, p. 275. 1971. Nom. illegit. Art.

64.1. On dead stems of Carlina vulgaris L.,
Compositae. Romania.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris forma syndoliola H. Rehm,
Ascomycetes Lojkani Lecti in Hungaria,
Transylvania et Galicia, Berlin, pp. 55-56.
1882. On dry stems of Peucedanum longifo-
lium Waldst. & Kit., Umbelliferae. Hungary.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. angustispora N.T. Patouillard,
Revue Mycologique, Toulouse 8:180. 1886.
On dead stems of Pedicularis debilis Franch.
ex Maxim., Scrophulariaceae. China.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. cacaliae N. Naumov,
Ural’skoe Obshchestvo Estestvoznaniia, v
Ekeaterinburgie Zadipiski 35(11—12):21.
1916. On dry stems of Cacalia hastata L.,
Compositae. U.S.S.R.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. conoidea V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:235. 1863.
=Leptosphaeria conoidea (V. Cesati & G. de
Notaris) P.A. Saccardo.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. dissimilis H. Rehm, Ascomy-
ceten, Fascicle 18, No. 888. Anno 1886;
Hedwigia, Dresden 26:94. 1887. [Reprint is
page 14 at FH.] On decayed stems of
Heracleum palmatum Baumg., Umbelliferae.
Romania.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. /eonuri C. Sandu-Ville,

Nomenclator of Leptosphaeria 225

Academia Republicii Populare Romine.
Filiala lasi. Studii si Cercetari Stiintifice.
Biologie si Stiinte Agricole 10 (Fascicle
2):213-214. 1959. On dead stems of
Leonurus cardiaca L., Labiatae. Romania.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. pachyspora P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 2:318.
1881. On stems of Adonis pyrenaica DC.,
Ranunculaceae. France.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. pinquicula P.A. Saccardo,
Revue Mycologique, Toulouse 3:44. 1881;
Reliquiae Mycologicae Libertianae, Series
Altera Reviserunt C. Roumegueére & P.A.
Saccardo, Toulouse, No. 121. 1881; Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 2:598. 1882. On
stems of Foeniculum sp., Umbelliferae.
France.

doliolum (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris var. subdisticha K. Starback &
A.Y. Grevillus in K. Starback, Bihang till K.
Svenska Vetenskapsakademiens Handlingar,
Stockholm 16(3), No. 3, pp. 6—7. 1890;
Bidrag till Kannedomen om Sveriges
Ascomycetflora, pp. 6-7. 1890. On dry stems
of Artemisia vulgaris L., Compositae.
Sweden.

donacina P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):155. 1873. =Phaeosphaeria donacina
(P.A. Saccardo) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1524. 1989.
[Anamorph: Ascochyta donacina P.A.
Saccardo.] On dead, decaying culms of
Arundo donax L., Gramineae. Netherlands.

drabae (W. Nylander) P.A. Karsten, Mycologia
Fennica Pars 2, Pyrenomycetes, p. 102. 1873.
=Sphaeria drabae W. Nylander in W.
Nylander and T. Saelan, Herbarium Musei
Fennici Forteckning Ofver Finska Musei Vaxt
samling, Utgifven Af Sallskapet Pro Fauna et
Flora Fennica Och Uppgjord af W. Nylander
Och Th. Saelan. Med en Karta, p. 112. 1859.
On stems of Draba alpina L., Cruciferae.
U.S.S.R.

dracaenae M. de Sousa da Camara, Revista
Agronomica, Lisbon 1:23. 1903. On dead
leaves of Dracaena draco (L.) L., Agavaceae.
Portugal.

dracaenae M. de Sousa da Camara forma rusci J.V.
d’Almeida & M. de Sousa da Camara. [An
error for Leptosphaeria convallarieae J.V.
d’Almeida & M. de Sousa da Camara forma
rusci J.V. d’ Almeida & M. de Sousa da
Camara. |

draconis M.J. de Urries y Azara, Anales del
Instituto Botanico A.J. Cavanillo, Madrid

226 Illinois Natural History Survey Bulletin

14:158-159. (1955) 1956. On leaves of
Dracaena draco (L.) L., Agavaceae. Spain.

drechsleri (R.A. Shoemaker) M.E. Barr in M.E.
Barr, C.T. Rogerson, S.J. Smith, and J.H.
Haines, Bulletin of the New York State
Museum, Albany 459:23. 1986. =Ophiobolus
drechsleri R.A. Shoemaker, Canadian Journal
of Botany, Ottawa 54:2376-2377. 1976. On
unknown substrate of Ambrosia trifida L.,
Helianthus annuus L., Helianthus grosserra-
tus Martens, Heliopsis sp., Verbesina
virginica L., Compositae. Canada, U.S.A.

dryadea P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:311. 1875. On dried leaves
of Quercus pedunculata Ehrh., Fagaceae.
Italy.

(Leptosphaerella) dryadea P.A. Saccardo subsp.
lussoniensis P.A. Saccardo, Annales Mycol-
ogici, Berlin 12:304—305. 1914. On dying
stems and leaves of Kigelia pinnata DC.,
Bignoniaceae. Philippines.

dryadis E. Rostrup, Botanisk Tidsskrift, Kjoben-
havn 25:305. 1903. On decorticated wood
stems and fruits of Dryas octopetala L.,
Rosaceae. Iceland.

dryadis E. Rostrup, Norske Ascomyceter 1 Chris-
tiania Universitetets Botaniske Museum.
Christiania [Oslo] (I. Kommission Hos Jacob
Dybwad), p. 24. 1904. [Issued in Skrifter
Udg. af Videnskabsselskabet i Christiania.
Mathematisk-naturvidenskabelig klasse.
Christiania (Oslo) No. 274.] Non vidi. On
upper surface of dried leaves of Dryas
octopetala L., Rosaceae. Norway.

dryophila (M.C. Cooke & H.W. Harkness) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:51. 1883. =Sphaerella dryophila M.C.
Cooke & H.W. Harkness, Grevillea, London
9:86. 1881. On leaves of Quercus sp.,
Fagaceae. U.S.A.

dubia P.A. Saccardo & J. Paoletti, Bulletin. Societe
R. de Botanique de Belgique, Bruxelles
28:90-91. 1889. On stems of Valeriana dubia
Turez., Valerianaceae. U.S.S.R.

dubiosa (V. Mouton) C.A. Oudemans, Enumeratio
Systematica Fungorum 1:981. 1919. =Lepto-
sphaeria nardi (E.M. Fries) V. Cesati & G. de
Notaris var. dubiosa V. Mouton.

dumetorum G. Niessl v. Mayendorf, Verhandlungen
des Naturforschenden Vereins in Briinn
10:176-177. (1871) 1872. On dried stems of
Humulus lupulus L., Urticaceae. Czechoslo-
vakia.

dumetorum G, Niessl v. Mayendort forma ebuli H.
Rehm ex H. Rehm, Ascomyceten, Fascicle 14,
No. 687. Anno 1882. Nom. nud. Art. 32.1;
Hedwigia, Dresden 22:56. 1883. On dry

Vol. 34 Art. 3

stems of Sambucus ebulus L., Caprifoliaceae.
Czechoslovakia.

dumetorum G. Niessl v. Mayendorf forma meliloti
H. Rehm, Hedwigia, Dresden 22:56. 1883.
On preceding year’s stems of Melilotus alba
Medicus, Leguminosae. Germany.

dumetorum G. Niessl v. Mayendorf var. coniformis
J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(2):159—-160.
1901. On dry stems of Senecio fuchsii C.C.
Gmelin, Compositae. Luxembourg.

dumetorum G. Niessl v. Mayendorf var. coronillae
G. Moesz, Arbeiten des Ungarischen
Biologischen Forschungs-Institutes, Tihany
3:96. 1930. On dead stems of Coronilla
coronata L., Leguminosae. Hungary.

dumetorum G. Niessl v. Mayendorf var. doli-
chospora J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(3):210-211. 1903. On corticated branches
of Sambucus racemosa L., Caprifoliaceae.
Luxembourg.

dumetorum G. Niessl v. Mayendorf var. galii-
borealis W.H. Trail, Transactions and
Proceedings of the Botanical Society of
Edinburgh 17:491. 1889. On dead stems of
Galium boreale L., Rubiaceae. Norway.

dumetorum G. Niessl v. Mayendorf var. marrubii
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:15. 1883. On stems of Marrubium vulgare
L., Labiatae. France.

dumetorum G. Niessl v. Mayendorf var. symphyti J.
Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):211.
1903. On dry stems of Symphytum cauca-
sicum Bieb., Boraginaceae. Luxembourg.

duplex (M.J. Sowerby:E.M. Fries) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:87.
1883. =Sphaeria duplex M.J. Sowerby:E.M.
Fries, Coloured Figures of English Fungi or
Mushrooms 3, Table 375. 1803; Systema
Mycologicum Sistens Fungorum 2:520. 1823.
On leaves of Sagittaria sp., Spartina sp..
Typha sp., Alismataceae, Gramineae,
Typhaceae. Germany, Great Britain, Sweden.

eburnea G. Niessl v. Mayendorf, Hedwigia, Dresden
20:98. 1881. =Metasphaeria eburnea (G.
Niessl v. Mayendorf ) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:162. 1883. On dry
stems of Chondrilla juncea L., Compositae.
Unknown country.

echiella J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg 1(3):219.
1903. On stems of Echium vulgare L.,
Boraginaceae. Luxembourg.

echii J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(4):28.

March 199]

1905. On barked stems of Echium vulgare L.,
Boraginaceae. Luxembourg.

echinella (M.C. Cooke) F. v. Thiimen, Mycotheca
Universalis, Century 3, No. 266. Anno 1875.
=Sphaeria echinella M.C. Cooke, Handbook
of British Fungi, p. 906. 1871. On rotten
stems of Atriplex sp., Chenopodiaceae. Great
Britain.

echinops (F.A. Hazslinszky) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:32. 1883. =Ple-
ospsora echinops F.A. Hazslinszky, Matema-
tikai és Természettudomanyi K6zlemenyek
Vonatkozolag a Hazai Viszonyokra, Budapest
10:47. 1872. On stems of Verbascum
phlomoides L., Scrophulariaceae. Hungary.

eichhorniae R. Gonzalez Fragoso & R. Cifferi,
Boletin de la R. Sociedad Espanola de
Historia Natural, Madrid 26:473-474. 1926.
Nom. inval. Art. 34.1. On dry leaves of
Eichhornia crassipes (C.F.P. Mart.) Solms-
Laub., Pontederiaceae. Dominican Republic.

elaeidicola K.A. Pirozynski, Mycological Papers.
Commonwealth Mycological Institute, Kew
129:19-20. 1972. On dead fronds of Elaeis
guineensis Jacq., Palmae. Tanzania.

elaeidis C. Booth & J.S. Robertson, Transactions of
the British Mycological Society, London
44:26. 1961. On leaves of Elaeis guineensis
Jacq., Palmae. Nigeria.

elaeospora (P.A. Saccardo) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:65. 1883. =Lepto-
sphaeria graminum P.A. Saccardo var.
elaeospora P.A. Saccardo. On dead leaves of
Phragmites communis Trin., Gramineae.
France.

elaoudi P. Rieuf, Al-Awamia; Revue de la Re-
cherche Agronomique Marocaine 16:51.
1965. On branches of Pelargonium capitatum
(L.) L’Her., Geraniaceae. Morocco.

ellisiana A.N. Berlese, cones Fungorum Omnium
Hucusque Cognitorum ad usum Sylloges
Saccardianae Adcommodatae 1:68-69. 1892.
=Leptosphaeria subconica J.B. Ellis & B.M.
Everhart, non Leptosphaeria subconica (G.W.
Clinton & C.H. Peck) P.A. Saccardo [as
Sphaeria (Leptosphaeria) subconica G.W.
Clinton & C.H. Peck in J.B. Ellis, North
American Fungi, Series I, Century 7, No. 697.
Anno 1881]. On dead stems of Oenothera
biennis L., Onagraceae. U.S.A.

elongata L.E. Wehmeyer, Mycologia, Lancaster,
Pennsylvania 44:633. 1952. =Phaeosphaeria
elongata (L.E. Wehmeyer) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1540.
1989. On dry stems of Elymus glaucus
Buckley, Gramineae. Canada, Finland,
U.S.A.

elymi P. Larsen in L. Kolderup Rosenvinge and E.
Warming, The Botany of Iceland 2(Part 3):

Nomenclator of Leptosphaeria

i)
tN
~

474-475. (1931) 1932. Nom. illegit. Art.
64.1. =Leptosphaeria larseniana A. Munk.
On dead plant of Elymus arenarius L.,
Gramineae. Iceland.

elymi G.F. Atkinson, Bulletin. Cornell University
Agricultural Experiment Station, Ithaca, New
York 3(1):7. 1897. On dead portions of
leaves of Elymus sp., Gramineae. U.S.A.

emiliana J.H. Fabre, Annales des Sciences
Naturelles, Paris, Botanique, Series 6, 9:90.
1878. On dry branches of Jasminum fruticans
L., Pistacia terebinthus L., Oleaceae,
Anacardiaceae. France.

empetri (A.P. de Candolle:E.M. Fries) H.G. Winter,
Dr. L. Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):487. 1885 [as (Fuckel)].
=Hyphoderma sphaerioides (J.B. Albertini &
L.D. v. Schweinitz) A.P. de Candolle var.
empetri A.P. de Candolle, Flore Frangaise
6:165. 1815. =Xyloma empetri C.H. Persoon
ined. =Hysterium sphaerioides J.B. Albertini
& L.D. v. Schweinitz, Conspectus Fungorum
in Lusatiae Superioris Agro Niskiensi
Crescentium e Methodo Persoonia, p. 57, No.
167. 1805. =Sphaeria empetri (A.P. de
Candolle) E.M. Fries:E.M. Fries, Systema
Mycologicum Sistens Fungorum 2:522. 1823.
=Sphaeria empetri (E.M. Fries?) L. Fuckel,
Symbolae Mycologicae Dritter Nachtrag, p.
18. 1875. Nom. illegit. Art. 64.1. =Metas-
phaeria empetri (A.P. de Candolle:E.M.
Fries) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:171. 1883. On dry leaves of
Empetrum nigrum L., Empetraceae. France.

endiusae (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:57. 1883. =Ple-
ospora endiusae L. Fuckel, Symbolae
Mycologicae, pp. 136-137. 1870. In
association with dried stems and pods of
Endiusa hirsuta Alef. (=Vicia), Leguminosae.
Germany.

endophaena E. Bommer, M. Rousseau & P.A.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:781. 1891. On dead branches of
Ailanthus glandulosa Desf., Simaroubaceae.
Belgium.

ephedrae A. Maublanc, Bulletin de la Société
Mycologique de France, Paris 21:88. 1905.
On dead branches of Ephedra distachya L.,
Ephedraceae. France.

epicalamia (H. Riess) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:236. 1863. =Sphaeria
epicalamia H. Riess in G.L. Rabenhorst,
Klotzschii Herbarium Vivum Mycologicum, I.
Ausgabe, Century 19, No. 1828. Anno 1854.
=Pleospora epicalamia L. Fuckel, Symbolae

228

Mycologicae, p. 139. 1870. =Phaeosphaeria
epicalmia (H. Riess) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):114.
1957. [Illustrations as Sphaeria epicolamia
H. Riess, Hedwigia, Dresden 1(6), Table IV,
Figure 6.] On dry culms of Luzula lutea (All.)
DC., Luzula nemorosa (Pollich)
E.H.F. Meyer, Luzula sylvatica (Hudson)
Gaudin, Juncaceae. Germany, Switzerland.

epicalamia V. Cesati & G. de Notaris var. pleospo-
roides J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(2):155. 1901. On dry culms of Luzula
albida (Hoffm.) DC., Juncaceae. Luxem-
bourg.

epicarecta (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:65. 1883.
=Sphaeria epicarecta M.C. Cooke, Grevillea,
London 5:120. 1877. On leaves of Carex sp.,
Cyperaceae. Great Britain.

epilobii E. Miller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):303-304. 1950. =Lepto-
sphaeria multiseptata H.G. Winter var. alpina
H. Rehm, Hedwigia, Dresden 24:235. 1885.
Non Leptosphaeria alpina A. Maublanc.
=Nodulosphaeria epilobii (E. Miiller) L.
Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):91. 1957. On dry stems of
Epilobium fleischeri Hochst., Epilobium
montanum L., Epilobium trigonum Schrank,
Onagraceae. Italy, Switzerland.

equiseti P.A. Karsten, Ofversigt af K. Vetenskap-
sakademiens Forhandlinger, Stockholm
29(2):101—102. 1872. =Phaeosphaeria
equiseti (P.A. Karsten) L. Holm & K. Holm,
Nordic Journal of Botany, Copenhagen
[Opera Botanica] 1:113. 1981. On aged stipes
of Equisetum variegatum Schleicher ex
Weber & Mohr, Equisetaceae. Canada,
U.S.S.R.

equiseticola L. Hollos, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:33. 1926. On dry
stems of Equisetum variegatum Schleicher ex
Weber & Mohr, Equisetaceae. Hungary.

eranthemi N.T. Patouillard, Revue Mycologique,
Toulouse 8:181. 1886. On peduncles of
Eranthemum sp., Acanthaceae. China.

eremophila (C.L. Spegazzini) P.A. Saccardo & A.
Trotter in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.

Saccardo 22:232. 1913. [Incorrectly cited as a

comb. nov. based on Leptosphaerella
eremophila C.L. Spegazzini.]| Nom. inval.
Art. 43.1.

ericae (E.M. Fries:E.M. Fries) A. Malbranche in
P.A. Saccardo, Syllogue Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
9:780-781. 1891. =Sphaeria ericae E.M.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Fries:E.M. Fries, Elenchus Fungorum Sistens
Commentarium in Systema Mycologicum
2:101. 1828. On trunks of Calluna vulgaris
(L.) Hull, Ericaceae. France.

erigerontis F.E. Clements & E.S. Clements,
Cryptogamae Formationum Coloradensium,
Century 1, No. 28. Anno 1906. Nom. inval.
Art. 32.1. On aged stems of Erigeron viscidus
Rydb., Compositae. U.S.A.

erigerontis (A.N. Berlese) A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:80-81. 1892. =Leptosphaeria agnita P.A.
Saccardo var. erigerontis A.N. Berlese. On
aged stems of Erigeron viscidus Rydb.,
Compositae. U.S.A. (Colorado).

eriobotryae H. Sydow, P. Sydow, & E. Butler,
Annales Mycologici, Berlin 9:409. 1911. On
leaves of Eriobotrya japonica (Thunb.)
Lindley, Rosaceae. India.

eriophora (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:87-88. 1883.
=Sphaeria (Caulicolae) eriophora M.C.
Cooke, Grevillea, London 5:153. 1877.
=Pocosphaeria eriophora (M.C. Cooke) A.N.
Berlese, Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:89. 1892. On stems of
Amaranthus sp., Chenopodium sp., Lappa sp.
(=Arctium sp.), Amaranthaceae, Chenopodi-
aceae, Compositae. U.S.A.

eryngit H. Fabre, Annales des Sciences Naturelles,
Paris, Botanique, Series 6, 15:49. 1883. On
somewhat dry stems and petioles of Eryngium
campestre L., Umbelliferae. France.

erythrinae H. Sydow, Annales Mycologici, Berlin
37:376. 1939. On dead deciduous leaves of
Erythrina sp., Leguminosae. Philippines.

espeletiae E. Miiller in E. Miiller and R.W.G.
Dennis, Kew Bulletin. Royal Botanic
Gardens, Kew 19:373, 384. 1965. On dead
inflorescence stalks of Espeletia (?Espeletia
schultzii Wedd.) (=Balsamorhiza), Composi-
tae. Venezuela.

ettalensis A. Allescher, Bericht der Bayerischen
Botanischen Gesellschaft zur Erforschung der
Heimischen Flora, Miinchen 5:13—14. 1897.
On dead stems of Laserpitium latifolium L.,
Umbelliferae. Germany.

eumorpha (M.J. Berkeley & M.A. Curtis) F.S. Earle,
Bulletin of the Torrey Botanical Club (and
Torreya), New York 25:361—362. 1898.
=Sphaeria eumorpha M.J. Berkeley & M.A.
Curtis, Grevillea, London 4:145. 1876.
=Didymella eumorpha (M.J. Berkeley &
M.A. Curtis) P.A. Saccardo, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 1:560. 1881. =Di-
dymosphaeria eumorpha (M.J. Berkeley &

March 1991

M.A. Curtis) G.F. Atkinson, Bulletin. Cornell
University Agricultural Experiment Station,
Ithaca, New York 3(1):6. 1889. On culms of
Arundinaria sp., Gramineae. U.S.A.
(Alabama, South Carolina).

euphorbiae G. Niess| v. Mayendorf in G.L. Raben-
horst, Fungi Europaei Exsiccati, Klotzschii
Herbarii vivi Mycologici Continuatio Edita
Nova, Series 2, Century 19, No. 1841. Anno
1874; Just’s Botanische Jarhesberichte, Berlin
2:319. 1876. On dried stems of Euphorbia
cyparissias L., Euphorbiaceae. Czechoslo-
vakia.

euphorbiae G. Niessl v. Mayendorf forma esulae J.
Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):212.
1903. On dry barked stems of Euphorbia
esula L., Euphorbiaceae. Luxembourg.

euphorbiaecola P. Brunaud, Bulletin de la Société
Botanique de France 34:244. 1887. On dead
stems of Euphorbia pilosa L., Euphorbiaceae.
France.

euphorbiicolla S. Ahmad, Monographs. Biological
Society of Pakistan, Lahore 8:87. 1978
(1979?) [as euphorbiaecola]. =Leptosphaeria
depressa L.E. Wehmeyer & S. Ahmad. 1964.
Nec Leptosphaeria depressa T. Petch. 1926.
On branches of Euphorbia osyridea Boiss.,
Euphorbiaceae. Pakistan.

eustoma (E.M. Fries:E.M. Fries) P.A. Saccardo, Atti
dell’ Accademia Scientifica Veneto-Trentino-
Istriana, Padova 2(2):258. 1873 [as Fr.?].
=Sphaeria eustoma E.M. Fries:E.M. Fries,
Elenchus Fungorum Sistens Commentarium
in Systema Mycologicum 2:109. 1828. On
pedicels of Sorghum vulgare Pers., Gramin-
eae. Italy.

eustoma (L. Fuckel) P.A. Saccardo, Fungi Italici
autographice delineati (additis nonnullis extra-
italicis asterisco notatis), Patavii, Table 497.
1879. Nom. illegit. Art. 64.1. =Pleospora
eustoma L. Fuckel, Symbolae Mycologicae, p.
139. 1870 [as Fr. El. II, p. 109, unter
Sphaeria(?); see discussion by L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):110-111. 1957]. =Phaeosphaeria
eustoma (L. Fuckel) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):109.
1957. On very rotten straw of unknown host
of Gramineae, Gramineae. Austria, Canada,
Switzerland, U.S.A.

eustoma (E.M. Fries:E.M. Fries) P.A. Saccardo
forma eustomoides (P.A. Saccardo) A.N.
Berlese, [cones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:56. 1890. [Vide Lepto-
sphaeria eustomoides P.A. Saccardo.]}

eustoma (E.M. Fries:E.M. Fries) P.A. Saccardo
forma /eguminosa C.E. Fairman, Annales
Mycologici, Berlin 4:327—328. 1906. On
inner surface of pods of Robinia pseudacacia
L., Leguminosae. U.S.A.

Nomenclator of Leptosphaeria 229

eustomella P.A, Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:251. 1881. On culms of Festuca
sp., Gramineae. Italy.

eustomoides P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:319-320. 1875.
=Leptosphaeria eustoma (E.M. Fries:E.M.
Fries) P.A. Saccardo forma eustomoides (P.A.
Saccardo) A.N. Berlese, Icones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardiannae Adcommodatae 1:56.
1890. =Phaeosphaeria eustomoides (P.A.
Saccardo) R.A. Shoemaker, Canadian Journal
of Botany, Ottawa 67:1526. 1989. On culms
of Andropogon ischaemum L., Gramineae.
Italy.

eustomoides P.A. Saccardo forma /olii F. Fautrey in
C. Roumeguére, Fungi Selecti Gallici
Exsiccati, Century 59, No. 5847. Anno 1891;
Revue Mycologique, Toulouse 13:167—168.
1891. On rachis of Lolium perenne L.,
Gramineae. France.

eustomoides P.A. Saccardo var. punctata A.C,
Batista, J.L. Bezerra & E.A.F. da Matta,
Memorias da Sociedade Broteriana, Coimbra
14:36-37. 1961. On leaves of Jambosa
malaccensis DC. (=Syzygium), Myrtaceae.
Brazil.

eutypoides C.H. Peck, Report. New York State
Museum of Natural History, Albany 38:105.
1885. On dead stems of Chenopodium album
L., Chenopodiaceae. U.S.A.

excelsa A.M. Saccas, Etude de la Flore Cryptoga-
mique des Caféiers en Afrique Centrale.
Bulletin de Institut Frangais du Café du Cacao
et d’Autres Plantes Stimulantes (Bulletin
IFCC No. 16), pp. 240-242. 1981. Nom.
inval. Art. 37.1. On dead branches of Coffea
excelsa Cheval., Rubiaceae. Central African
Republic.

exocarpogena C.E. Fairman, Proceedings of the
Rochester Academy of Science, Rochester,
New York 6:98. 1921. Ona shuck of hickory
nut tree of Hicoria Raf. (=Carya Nutt.),
Juglandaceae. U.S.A.

fagaricola (C.L. Spegazzini) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 24:1000. 1928.
[Incorrectly cited as a comb. nov. based on
Leptosphaerella fagaricola C.L. Spegazzini.]
Nom, inval. Art. 43.1.

faginea G., Passerini, Atti dell’ Reale Accademia
Nazionale dei Lincei. Rendiconti, Rome,
Series 4, 7(2):44. 1891. On dead stems of
Fagus sp., Fagaceae. Italy.

fallaciosa A.N. Berlese, Bulletin Trimestriel de la
Société Mycologique de France, Paris 5:43.
1889. On dead stems of Satureja hortensis L.,
Labiatae. Italy.

230

fallax A.N. Berlese, Fungi Moricolae Iconographia e
Descrizione dei Funghi Parassiti del Gelso,
Fascicle IV, No. 3. Anno 1887. On dead
stems of Morus alba L., Moraceae. Italy.

faulii G.D. Darker, Canadian Journal of Botany,
Ottawa 42:1006—1008. 1964. On dead
needles of Abies balsamea Miller, Pinaceae.
Canada, U.S.A.

feijoae G.V. Artemieff, Sovetskie Subtropiki,
Sukhum 7:62. 1935. On living leaves of
Feijoa sp. [as F. feijoicola] (=Acca), Myrta-
ceae. U.S.S.R.

feltgeni P.A. Saccardo & P. Sydow in P.A. Saccardo,

Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 16:513.
1902. Nom. illegit. Art. 64.1. =Lepto-
sphaeria hemerocallidis J. Feltgen.
ferruginea M. Chochrjakov, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 7:144. 1951. On
living leaves and stems of Ipomoea batatas
(L.) Lam., Convolvulaceae. U.S.S.R.
ferulicola E.N. Koschkelova in E.N. Koschkelova,
I.P. Frolov, and Z. Dzhuraeva, Mikoflora
Badkhyza, Karabilya i Yuzhnoi Chasti
Murgabskogo, Oazisa (Mikromitsety) [The
Mycoflora of Badkhyz, Karabil and the
Southern Part of the Murgab Oasis (Micromy-
cetes)], p. 87. 1970. Nom. nud. Art. 32.1. On
dry stems and stalks of Ferula badra-kema K.
Pol., Umbelliferae. U.S.S.R. (Karabilia).
fibrincola F. v. Hohnel & H. Rehm in H. Rehm,
Annales Mycologici, Berlin 3:228. 1905;
Ascomyceten, Fascicle 34, No. 1591. Anno
1905. On rotten paper. Germany.
fici-elasticae F. Petrak, Annales Mycologici, Berlin
14:164. 1916. On dead leaves of Ficus
elastica Roxb., Moraceae. Czechoslovakia.
fiedlaeri (G. Niessl v. Mayendorf) P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 1:39.
1877. =Cryptospora fiedlaeri G. Niessl v.
Mayendorf, Hedwigia, Dresden 13:42-43.
1874. =Metasphaeria fiedlaerii P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:165. 1883. On branches of Cornus
sanguinea L., Cornaceae. Italy.
filamentosa J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 4:76. 1888. On
dead pieces of living leaves of Yucca
filamentosa L., Agavaceae. U.S.A.
filiformis L.E. Wehmeyer, Mycologia, Lancaster,
Pennsylvania 44:641. 1952. =Ophiobolus
filiformis (L.E. Wehmeyer) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa
54:2378-2379. 1976. On stem of unknown
host of composite, Compositae. U.S.A.
fimbriata J.B. Ellis & B.M. Everhart, The North
American Pyrenomycetes. A Contribution to

Illinois Natural History Survey Bulletin

Mycologic Botany, p. 357. 1892. On
herbaceous stems of unknown host, unknown
family. U.S.A.

fimiseda H.G. Winter, Hedwigia, Dresden 10:163.
1871. On smoked hare (Lepus europaeus,
Leporidae) in Harth Woods. Germany.

fiumana F. Hazslinszky, Matematikai és Természet-
tudomanyi K6zlemenyek Vonatkozolag a
Hazai Viszonyokra, Budapest 25(2):143.
1892. On stems of unknown host of Labiatae,
Labiatae. Hungary.

flotoviae C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cordoba 25:70-71.
1921. On dead and dry clump of Flotovia
excelsa DC. (=Dasyphyllum), Compositae.
Chile.

fluviatilis (W. Phillips & C.B. Plowright) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:84. 1883. =Sphaeria fluviatilis W. Phillips
& C.B. Plowright, Grevillea, London 10:73.
1881. On unknown substrate of Lemanea
fluviatilis C. Aq., Algae (Lemaneaceae).
Great Britain.

foeniculacea H. Fabre, Annales des Sciences
Naturelles, Paris, Botanique, Series 6, 15:51.
1883. On dead stems of Foeniculum vulgare
Miller, Umbelliferae. France.

foeniculacea H. Fabre subsp. lupina P.A. Saccardo
& G. Scalia in P.A. Saccardo, C.H. Peck, and
W. Trelease, Harriman Alaska Expedition
5:29-30. 1904. On decaying stems of
Lupinus sp., Leguminosae. U.S.A. (Alaska).

foeniculi R. Gonzalez Fragoso, Boletim da So-
ciedade Broteriana, Coimbra, Series 2, 2:23.
1923. On dry stems of Foeniculum vulgare
Miller, Umbelliferae. Portugal.

foliicola N.A. Naumov, Ural’skoe Obshchestvo
Estestvoznaniia, v Ekeaterinburgie Zadipiski
35(11-12):21. 1916. On living leaves of
Libanotis montana Crantz, Umbelliferae.
U.S.S.R.

folliculata J.B. Ellis & B.M. Everhart, Proceedings
of the Academy of Natural Sciences of
Philadelphia 1890:237. 1890. On leaves of
Carex folliculata L., Cyperaceae. Canada
(Ontario).

folliculata J.B. Ellis & B.M. Everhart var. oxyspora
J.J. Davis, Transactions of the Wisconsin
Academy of Science, Arts and Letters,
Madison 18:87. 1915. =Didymella oxyspora
(J.J. Davis) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1576. 1989. On
unknown substrate of Carex gracillima
Schwein., Cyperaceae. U.S.A.

francoae (C.L. Spegazzini) P.A. Saccardo & A.
Trotter, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
22:221. 1913. =Leptosphaerella francoae
C.L. Spegazzini, Fungi Chilenses, p. 77. 1910;

Vol. 34 An. 3

:

March 1991

Revista de La Facultad de Agronomia Y
Veterinaria, Universidad Nacional de La
Plata, Series 2, 6:77. 1910. On year-old dry
scapes of Francoa sonchifoliae Cav.,
Saxifragaceae. Chile.

franconica (F. Petrak) E. Miller, Sydowia. Annales
Mycologici, Horn, N.O. 4(1—6):298-299.
1950. =Leptosphaeria derasa (M.J. Berkeley
& C.E. Broome) B. Auerswald var. fran-
conica F, Petrak, Kryptogamische
Forschungen, Miinchen 2:162—163. 1931.
=Nodulosphaeria franconica (F. Petrak) L.
Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):84. 1957. On unknown
substrate of /nula salicina L., Compositae.
Switzerland.

fraserae J.B. Ellis & B.M. Everhart, Bulletin of the
Torrey Botanical Club (and Torreya), New
York 27:52-53. 1900. On dead stems of
Frasera speciosa Dougl. ex Griseb., Gen-
tianaceae. U.S.A.

fraxini J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 3:44. 1887. On
living leaves of Fraxinus americana L.,
Oleaceae. U.S.A.

frigida L.N. Vasil’eva, Pirenomitsety i
Lokuloaskomitsety Severa Dal’nego Vostoka
(Leningrad:“Nauka”), p. 117. 1987 [as
frigidus|. On dead leaves of Luzula sp.,
Juncaceae. U.S.S.R.

frondis W. Kirschstein, Annales Mycologici, Berlin
33:212. 1935. On young living stems of
Lonicera tatarica L., Caprifoliaceae. Ger-
many.

fuckelii G. Niess| v. Mayendorf in W. Voss, Oster-
reichische Botanische Zeitschrift, Wien
32:357-358. 1882. =Phaeosphaeria fuckelii
(G. Niessl) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):123. 1957. On
dead stems of Calamagrostis montana Host,
Gramineae. Canada, Germany, Romania,
Sweden, Switzerland, U.S.A., Yugoslavia.

fuckelii G. Niess| v. Mayendorf forma filamentifera
P.A. Karsten, Hedwigia, Dresden 27:261.
1888. On sheath of Phalaris arundinacea L.,
Gramineae. Finland.

fuegiana C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cordoba 11:219.
1887; Fungi Fuegiani, No. 231. Anno 1887.
On leaves and stems of Hierochloé antarctica
R. Br., Gramineae. Argentina.

fulgida (M.C. Cooke & C.A. Peck) M.E. Barr in
M.E. Barr, C.T. Rogerson, S.J. Smith, and
J.H. Haines, Bulletin of the New York State
Museum, Albany 459:23. 1986. =Sphaeria
(Caulicolae) fulgida M.C. Cooke & C.A. Peck
in M.C. Cooke & J.B. Ellis, Grevillea,
London 6:15. 1877 [as G.W. Clinton & C.A.
Peck in C.A. Peck, Report. New York State
Museum of Natural History, Albany 29:62.

Nomenclator of Leptosphaeria 2

Ww

(1875) 1878]. =Ophiobolus fulgidus (M.C.
Cooke & C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:346. 1883.
=Rhaphidospora fulgida (M.C. Cooke & C.H.
Peck) M.C. Cooke, Grevillea, London 18:16.
1889. On stems of Ambrosia trifida L.,
Compositae. U.S.A. (New York).

fungicola H.G. Winter, Hedwigia, Dresden 25:101.

1886. On aged hymenium of Stereum
subpileatum Berkeley & Broome, Fungi
(Stereaceae). Sao Tome.

fuscella (M.J. Berkeley & C.E. Broome) V. Cesati &

G. de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:236. 1863.
=Sphaeria (Obtectae) fuscella M.J. Berkeley
& C.E. Broome, Annals and Magazine of
Natural History, London, Series 2, 9:325.
1882. On dead twigs of Rosa sp., Rosaceae.
Great Britain.

fuscella (M.J. Berkeley & C.E. Broome) P.A.

Saccardo forma microspora C. Roumeguere,
Revue Mycologique, Toulouse 12:163. 1890;
Fungi Selecti Gallici Exsiccati, Century 55,
No. 5438. Anno 1890. On dry branches of
Sarothamnus scoparius (L.) Wimmer ex
Koch, Leguminosae. France.

fuscella V. Cesati & G. de Notaris var. hippophaes J.

Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):212.
1903. On barky branches of Hippophaé
rhamnoides L., Elaeagnaceae. Luxembourg.

fuscella (M.J. Berkeley & C.E. Broome) V. Cesati &

G. de Notaris var. sydowiana P.A. Saccardo in
H. Sydow, Mycotheca Germanica, Fascicle
10-11, No. 485. Anno 1906; Annales
Mycologici, Berlin 4:484. 1907. On branches
of Hippophaé rhamnoides L., Elaeagnaceae.
Germany.

fuscidula G. Passerini, Rendiconti della Sedute della

R. Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:91. 1887. On dry leaves of
Melica altissima L., Gramineae. Italy.

fuscidula G, Passerini forma magnolii R. Gonzalez
Fragoso, Trabajos del Museo Nacional de
Ciencias Naturales, Madrid, Series Botanica
10:88. 1916. On wilted or dry leaves of
Melica magnolii Gren. & Godron, Gramineae.
Spain.

fusispora G. Niessl vy. Mayendorf, Verhandlungen
des Naturforschenden Vereins in Briinn
10:177—178. (1871) 1872. On fallen
branchlets of Genista tinctoria L., Legumino-
sae. Austria (near Gratz).

fusispora G. Niessl v. Mayendorf forma erysimi C.
Sandu-Ville, Ciuperci Pyrenomycetes-
Sphaeriales Din Romania, pp. 277-278. 1971.
On branches of Erysimum hieracifolium L.,
Cruciferae. Romania.

232

galeobdolonis J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(3):221. 1903. On dry stems of Galeobdolon
luteum Hudson, Labiatae. Luxembourg.

galeopsidicola F. Petrak, Annales Mycologici,
Berlin 25:289. 1927. On stems of Galeopsis
speciosa Miller, Labiatae. Czechoslovakia.

galii (G.H. Otth) P.A. Saccardo, Hedwigia Beiblatt,
Dresden 35:XXIX. 1896; Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 11:XXIX. 1896. =Pleospora galii
G.H. Otth, Mitteilungen der Naturforschenden
Gesellschaft in Bern 1867(660):56. 1868. On
stems of Galium sp., Rubiaceae. Switzerland.

galii-silvatici W. Kirschstein, Verhandlungen des
Botanischen Vereins der Provinz Bran-
denburg, Berlin 48:56. (1906) 1907. On dry
stems of Galium sylvaticum L., Rubiaceae.
Germany.

galiicola P.A. Saccardo, Atti dell” Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):152-153. 1873. On fallen rotten stems of
Galium mollugo L., Rubiaceae. Italy?,
Netherlands.

galiicola P.A. Saccardo var. brachyspora P.A.
Saccardo, Nuovo Giornale Botanico Italiano e
Bolletino della Societa Botanica Italiana,
Firenze 7:315. 1875. On stems of Centran-
thus sp., Hesperis sp., Lappa sp., Valeri-
anaceae, Cruciferae, Compositae. Italy.

galiorum (M.R. Roberge) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:235. 1863. =Sphaeria
(Caulicola) galiorum M.R. Roberge in J.
Desmazieres, Annales des Sciences
Naturelles, Paris, Botanique, Series 3, 6:77.
1846. =Metasphaeria galiorum (M.R.
Roberge) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:160. 1883. On old stems of
Galium sp., Rubiaceae. France.

(Metasphaeria) galiorum (M.R. Roberge) H.G.
Winter, Hedwigia, Dresden 26:57—58. 1887.
Nom. illegit. Art. 64.1. On old stems of
Galium sp., Rubiaceae. France.

galiorum P.A. Saccardo forma cirsiorum P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo

Scop., Compositae. France.

galiorum P.A. Saccardo forma gentianae P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:22. 1883. On dead stems of Gentiana lutea
L., Gentianaceae. France.

galiorum P.A. Saccardo subsp. antirrhini P.A.
Saccardo, Nuovo Giornale Botanico Italiano e
Bolletino della Societa Botanica Italiana,
Firenze, New Series 22:38—39. 1915. On
stems of Antirrhinum siculum Miller,
Scrophulariaceae. Yugoslavia.

Illinois Natural History Survey Bulletin

Vol. 34 An. 3

galiorum P.A. Saccardo var. gentianae P.A.
Saccardo, Atti del Istituto Veneto di Scienze,
Lettere ed Arti, Venezia, Series 6, 2:456-457.
(1883) 1884. On stems of Gentiana lutea L..,
Gentianaceae. Switzerland.

galiorum P.A. Saccardo var. lapsanae P.A. Saccardo
& P.A. Briard, Revue Mycologique, Toulouse
7:209. 1885. On dead stems of Lampsana
communis L. (=Lapsana communis L.),
Compositae. France.

galiorum P.A. Saccardo var. gnaphaliana C_E.
Fairman in C.F. Millspaugh and L.W. Nuttall,
Fieldiana:Botany, Chicago 5:352. 1923. On
stems of Gnaphalium sp., Compositae.
U.S.A. (California).

galligena K. Keissler, Beihefte zum Botanischen
Zentralblatt, Cassel, Abt. II, 37:274-275.
1920. On thallus of Parmelia [as P. atrata
Zahlbr.], Lichenes (Parmeliaceae). U.S.A.
(Hawaii).

gaubae F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 9(1-6):562. 1955. =Phaeo-
sphaeria gaubae (F. Petrak) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1514.
1989. On dry leaves of Danthonia frigida
Vickery, Gramineae. Australia.

gaultheriae J. Dearness, Mycologia, Lancaster,
Pennsylvania 9:349. 1917. On dead stems of
Gaultheria shallon Pursh, Ericaceae. Canada
(British Columbia).

geasteris L. Hollos, Annales Historico-Naturales
Musei Nationales Hungarici, Budapest 5:279.
1907. In internal layers of exoperidium of
Geaster pseudolimbatus Hollos, Fungi
(Geastraceae). Hungary.

genistae C.A. Oudemans, Verslagen van de Gewone
Vergadering der Wis-en Natuurkundige
Afdeeling. K. Academie van Wetenschappen
Te Amsterdam 9(1):141. 1900. On fruits of
Genista anglica L., Leguminosae. Nether-
lands.

genistae C.A. Oudemans var. microspora A.L.
Guyot, Revue de Mycologie, Paris 11:68.

1946. On dry stems of Genista tinctoria L.,
Leguminosae. France.

geographicola (F. Amold) P.A. Saccardo & D.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 17:731. 1905. =Phaeospora
geographicola F. Amold in F.W. Zopf,
Hedwigia, Dresden 35:356—357. 1896.
=Phaeospora geographicola F. Amold,
Verhandlungen der Zoologisch-Botanischen
Gesellschaft in Wien 46:139-140. 1896.
(Effectively published in Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wien
24:284. 1874.] On thalli of Rhizocarpon
geographicum (L.) DC., Lichenes (Rhizocar-
paceae). Germany.

March 199]

geographicola (F. Arnold) L. Vouaux, Bulletin de la
Société Mycologique de France, Paris 29:118.
1913. Nom. illegit. Art. 64.1. On thallus of
Rhizocarpon geographicum (L.) DC.,
Lichenes (Rhizocarpaceae). Italy.

georgius-fischeri R. Sprague, Mycologia, Lancaster,
Pennsylvania 50:815. 1958. On living leaves
of Muehlenbergia filiformis Rydb., Gramin-
eae. U.S.A.

gibelliana R. Pirotta, Atti dell” Istituto Botanico
della Universita e Laboratorio Crittogamica di
Pavia, Milano 2 & 3:164. (1877) 1879. On
branches of Vitis vinifera L., Vitaceae. Italy.

gigaspora A.M. Saccas, Etude de la Flore Cryptoga-
mique des Caféiers en Afrique Centrale.
Bulletin de Institut Frangais du Café du Cacao
et d’Autres Plantes Stimulantes (Bulletin
IFCC No. 16), pp. 248-250. 1981. Nom. nud.
Art. 37.1. On dead stems and branches of
Coffea robusta L. Linden (=Coffea canephora
Pierre ex Froehn.), Rubiaceae. Central
African Republic.

gigaspsora G. Niessl v. Mayendorf in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:65.
1883; G. Niessl v. Mayendorf in G.L. Raben-
horst, Fungi Europaei Exsiccati, Klotzschii
Herbarii vivi Mycologici Continuatio Edita
Nova, Series 2, Century 30, No. 2998. Anno
1883. On leaves of Carex paludosa Good.,
Cyperaceae. Czechoslovakia.

gillotiana P.A. Saccardo & C. Roumegueére, Revue
Mycologique, Toulouse 5:236. 1885. On
corticated branches of Salix sp., Salicaceae.
France.

ginkgo S.A. Gucevicz, Novosti Sistematiki Nizshikh
Rastenij, Novitates Systematicae Plantarum
Non Vascularium 7:160. 1970. On dry
branches of Ginkgo biloba L., Ginkgoaceae.
WS-S:R:

glandulosae A.J. Lobik, Bolezni Rastenii S.
Peterburg (Morbi Plantarum Scripta Sectionis
Phytopathologiae Horti Botanici Principalis)
17:161. 1928. On leaves of Ailanthus
glandulosa Desf., Simaroubaceae. U.S.S.R.

glaucopunctata (R.K. Greville) B. Auerswald,
Hedwigia, Dresden 7:185. 1868. =Crypto-
sphaeria glaucopunctata R.K. Greville, Flora
Edinensis or a Description of Plants Growing
Near Edinburgh, pp. 362-363. 1824.
=Sphaeria glaucopunctata (R.K. Greville) F.
Currey, Transactions of the Linnean Society
of London 22:333. 1859. On dead leaves of
Ruscus aculeatus L.,; Ruscaceae. France,
Great Britain, Italy.

gloeospora (M.J. Berkeley & F. Currey) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:25. 1883. =Sphaeria (Caulicolae)
gloeospora M.J. Berkeley & F. Currey in M.J.

Nomenclator of Leptosphaeria

Berkeley and C.E. Broome, Annals and
Magazine of Natural History, London, Series
3, 7:454. 1861. =Trichometasphaeria
gloeospora (M.J. Berkeley & F. Currey) L.
Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):144. 1957. On stems of
Artemisia absinthium L., Compositae. Great
Britain.

glyceriae L.M. Unamuno Yrigoyen, Revista de la R.
Academia de Ciencias exactas, fisicas y
naturales de Madrid 30:483-486. 1933. On
leaves of Glyceria fluitans (L.) R. Br.,
Gramineae. Spain.

glyceriae-plicatae T. Savulescu & C. Sandu-Ville,
Hedwigia, Dresden 73:74—75. 1933.
=Phaeosphaeria glyceriae-plicatae (T.
Savulescu & C. Sandu-Ville) R.A. Shoe-
maker, Canadian Journal of Botany, Ottawa
67:1527. 1989. On wilted leaves of Glyceria
plicata (Fries) Fries, Gramineae. Romania.

gnaphalii (G.D. Westendorp & L. Fuckel) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:34. 1883. =Pleospora gnaphalii G.D.
Westendorp? in L. Fuckel, Fungi Rhenani
Exsiccati A Leopoldo Fuckel Collecti
Supplement, Fascicle 7, Total Series Fascicle
22, No. 2153. Anno 1868; Symbolae
Mycologicae, p. 136. 1870. On dry stems of
Gnaphalium sp., Compositae. Austria,
Belgium.

gossypit N.N. Woronichin, Trudy Akademiia Nauk
SSSR Botanicheskii Muzei 21:125. 1927. On
spots, on leaves of Gossypium sp., Malvaceae.
U.S.S.R.

graminis (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:76-77. 1883.
=Pleospora graminis L. Fuckel, Symbolae
Mycologicae, p. 139. 1870. =Phaeosphaeria
graminis (L. Fuckel) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):118.
1957. On dry culms of Phragmites communis
Trin., Gramineae. Germany, Great Britain.

graminum P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:119. 1878; Fungi Italici autogra-
phice delineati (additis nonnullis extra-italicis
asterisco notatis), Patavii, Table 483. 1879.
=Metasphaeria graminum P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:174.
1883. On dead leaves of Calamagrostis sp.,
Gramineae. Germany.

graminum P.A, Saccardo var. elaeospora P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
2:320. 1881. SLeptosphaeria elaeospora
(P.A. Saccardo) P.A. Saccardo, Sylloge

234

Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:65. 1883.

grammodes (G. de Notaris) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:235. 1863.
=Sphaeria grammodes G. de Notaris,
Memorie della Accademia delle Scienze di
Torino, Series 2, 2:74-75. 1841. On dry
stems of Artemisia vulgaris L., Compositae.
Italy.

grandispora P.A. Saccardo, Michelia Commentar-
ium Mycologicum Fungos in Primis Italicos
Illustrans 1:341. 1878. =Metasphaeria
grandispora (P.A. Saccardo) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:181.
1883. =Massariosphaeria grandispora (P.A.
Saccardo) A. Leuchtmann, Sydowia. Annales
Mycologici, Horn, N.O. 37:172. 1984.
=Lophiotrema grandispora (P.A. Saccardo)
R.A. Shoemaker, Canadian Journal of Botany,
Ottawa 67:1580. 1989. On decaying leaves of
Typha latifolia L., Typhaceae. Italy.

gratissima P. Rieuf & G. Teasca, Al-Awamia; Revue
de la Recherche Agronomique Marocaine
34:54. 1970 [as gratissimae]. On wood and
leaves of Persea americana P. Mill., Lau-
raceae. Morocco.

gratissima P. Rieuf & G. Teasca var. longispora P.
Rieuf & G.Teasca, Al-Awamia; Revue de la
Recherche Agronomique Marocaine 34:56.
1970 [as gratissimae]. On branches, on
irregular, marginal spots of leaves of Persea
americana P. Mill., Lauraceae. Morocco.

grignonnensis A.L. Guyot, Revue de Mycologie,
Paris 11:66—68. 1946. On dry runners of
Clematis vitalba L., Ranunculaceae. France.

grisea G. Passerini, Atti della R. Accademia dei
Lincei Memorie, Rome, Series 4, 6:459.
(1889) 1890. On wilted culms of Sorghum
vulgare Pers., Gramineae. Italy.

grossulariae Z. Girzitska, Universytet Botanichnyi
Sad Visnik Izvestiia Kiev. 1(Livr. V—VI):167.
1927. On living branches of Ribes grossu-
laria L., Saxifragaceae. Spain.

guazimae R. Gonzalez Fragosa & R. Ciferri, Boletin
de la R. Sociedad Espanola de Historia
Natural, Madrid 26:474. 1926. Nom. nud.
Art. 34.1. On living leaves of Guazuma sp.,
Sterculiaceae. Dominican Republic.

guazumae R. Gonzalez Fragoso & R. Ciferri,
Publicaciones Estacion Agronomica de Moca,
Santa Domingo. Series B, Botanica, No.
8:22—23. 1927. Nom. inval. Art. 34.1. On
living leaves of Guazuma sp., Sterculiaceae.
Dominican Republic.

gynerii C.L. Spegazzini, Anales del Museo Nacional
de Historia Natural de Buenos Aires
6:281—282. 1898; Fungi Argentini Novi vel
Critici, No. 627. Anno 1899. On rotting dead

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

leaves of Gynerium argenteum Nees,
Gramineae. Argentina.

gypsophilae L. Hollés, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 5:45.
1907. On dry stems of Gypsophila paniculata
L., Caryophyllaceae. Hungary.

haematites (M.R. Roberge) G. Niessl v. Mayendorf
in G.L. Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio Edita Nova, Series 2, Century 28, No.
2761. Anno 1882. =Sphaeria (Caulicola)
haematites M.R. Roberge in J. Desmaziéres,
Annales des Sciences Naturelles, Paris,
Botanique, Series 3, 16:311. 1851. =Didy-
mella haematites (M.R. Roberge) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
1:553. 1882. =Sphaerella haematites (MLR.
Roberge) M.C. Cooke, Journal of Botany,
British and Foreign, London 21:109. 1883.
On dry stems of Clematis vitalba L., Ranun-
culaceae. France.

haematites (M.R. Roberge) H.G. Winter, Hedwigia,
Dresden 22:10. 1883. Nom. illegit. Art. 64.1.
=Sphaeria (Caulicola) haematites M.R.
Roberge in J. Desmaziéres, Annales des
Sciences Naturelles, Paris, Botanique, Series
3, 16:311. 1851. =Didymella haematites
(MLR. Roberge) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 1:553. 1882 [as M.R.
Roberge et J. Desmazieres]. =Sphaerella
haematites (M.R. Roberge) M.C. Cooke,
Journal of Botany, British and Foreign,
London 21:109. 1883. On runners of
Clematis vitalba L., Ranunculaceae. Bel-
gium, France, Germany.

haematites (M.R. Roberge) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:36. 1883. Nom.
illegit. Art. 64.1. =Sphaeria (Caulicola)
haematites M.R. Roberge in J. Desmaziéres,
Annales des Sciences Naturelles, Paris,
Botanique, Series 3, 16:311. 1851. On dry
stems of Clematis vitalba L., Ranunculaceae.
France.

halima T. Johnson, Mycologia, Lancaster, Pennsyl-
vania 48:502. 1956. =Phaeosphaeria halima
(T. Johnson) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1514. 1989. On
test panel of Liriodendron tulipifera L.
submerged in salt water, Magnoliaceae.
U.S.A.

haloxyli B. Kravtzev, Trudy Instituta Botaniki
Akademiya Nauk Zazakhskoi SSR Alma-Ata
2:146—147. 1955. On decorticated, rotten
wood of Haloxylon aphyllum (Minkw.) Iljin,
Chenopodiaceae. U.S.S.R.

hamamelidis C.E. Fairman, Proceedings of the
Rochester Academy of Science, Rochester,

March 1991

New York 6:127. 1929. On dead plant of
Hamamelis virginiana L., Hamamelidaceae.
U.S.A.

hardenbergiae C.G. Hansford, Proceedings of the
Linnean Society of New South Wales, Sydney
79:120-121. 1954. On leaves of Hardenber-
gia violacea (Schneev.) Stearn, Apocynaceae.
Australia.

harknessianna J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 1:91. 1885. On
dead stems of Frasera? sp., Gentianaceae.
U.S.A.

hausmanniana B. Auerswald, Hedwigia, Dresden
7:185. 1868; Osterreichische Botanische
Zeitschrift, Wien 18:241. 1868. On upper
surface of dead leaves of Silene acaulis (L.)
Jacq., Caryophyllaceae. Italy.

hausmanniana B. Auerswald var. cherleriae P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:47. 1883. [Based on Terry specimen #6179,
published as Leptosphaeria hausmanniana B.
Auerswald in P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 2:599. 1882.] On
underside of dead leaves of Silene acaulis (L.)
Jacq., Caryophyllaceae. France.

hazslinskyana A.N. Berlese, Icones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:78.
1892 [as Leptosphaeria hanzslinskyana]. On
culm of an unknown host of grass, Gramin-
eae. Germany.

hazslinszkii P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:28. 1883. [Based on Ditopella sp.,
published by F.A. Hazslinszky, Verhand-
lungen der Zoologisch-Botanischen
Gesellschaft in Wien 20:217. 1870.] On dead
branches of Rosa sp., Rosaceae. Romania.

hederae (M.J. Sowerby?) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:313.
1875. =Sphaeria hederae P.A. Saccardo, Atti
dell’ Accademia Scientifica Veneto-Trentino-
Istriana, Padova 2(1):144—145. 1873 [as
Sphaeria hedarae M.J. Sowerby?, Sphaeria
hederae M.C. Cooke?]. =Sphaeria hederae
M.J. Sowerby, Coloured Figures of English
Fungi or Mushrooms 3, Table 371, Figure 5.
1803. =Metasphaeria hederae (M.J.
Sowerby?) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:169. 1883. On lower surface of
rotting leaves of Hedera helix L., Araliaceae.
Great Britain?, Italy.

hedericola (J. Desmaziéres) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:51. 1883. =Sphaeria
(Depazea) hedericola J. Desmaziéres,

Nomenclator of Leptosphaeria

Annales des Sciences Naturelles, Paris,
Botanique, Series 3, 14:115. 1850 [as
hederoecola\. On wilted and dead leaves of
Hedera helix L., Araliaceae. France.

helianthemi B. Auerswald in W. Gonnermann and
G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Sammtlicher Pilze Europa’s,
Hefte 5 und 6, Synopsis Pyrenomycetum
Europaeorum, Table 12, Figure 166. 1869.
Nom. nud. Art. 32.1. =Metasphaeria
helianthemi (B. Auerswald) ex P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:160.
1883. On stems of Helianthemum sp.,
Compositae. Germany.

helianthi J.B. Ellis & B.M. Everhart in J.F. Brenckle,
Mycologia, Lancaster, Pennsylvania 9:284.
1917. Nom. nud. Art. 32.1. On unknown
substrate of Helianthus giganteus L.,
Compositae. U.S.A. (North Dakota).

helianthi S.T. Tilak in T.S. Viswanathan and S.T.
Tilak, Mycopathologia et Mycologia
Applicata, Den Haag 13:241. 1960. On dry
culms of Helianthus annuus L., Compositae.
India.

helichrysi M.C. Cooke [falsely so cited in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
9:783. 1891]. =Leptosphaerella helichrysi
M.C. Cooke, Grevillea, London 19:7. 1890.

helicicola (J. Desmazieres) G. Niess] v. Mayendorf,
Verhandlungen des Naturforschenden Vereins
in Briinn 10:175. (1871) 1872. =Sphaeria
(Folticola) helicicola J. Desmaziéres, Annales
des Sciences Naturelles, Paris, Botanique,
Series 3, 11:356—357. 1849. =Metasphaeria
helicicola (J. Desmaziéres) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:169.
1883. On dried leaves of Hedera helix L.,
Araliaceae. France.

heliopsidis (L.D. vy. Schweinitz:E.M. Fries) L. Holm,
Svensk Botanisk Tidskrift, Stockholm 62:230.
1968. =Sphaeria heliopsidis L.D. v. Schwein-
itz, Synopsis Fungorum Carolinae Superioris,
pp. 34-35. 1822. =Dothidea heliopsidis (L.D.
v. Schweinitz:E.M. Fries) L.D. v. Schweinitz,
Synopsis Fungorum in America Boreali
Media Degentium. Secundum Observationes
Ludovici Davidis de Schweinitz, p. 231. 1832;
E.M. Fries, Systema Mycologicum Sistens
Fungorum 2:556. 1823. =Montagnella
heliopsidis (L.D. vy. Schweinitz:E.M. Fries)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:646. 1883. =Phaeoderris heliopsidis (L.D.
v. Schweinitz:E.M. Fries) F. v. Hohnel,
Sitzungsberichte der Akademie der Wissen-
schaften in Wien, Mathematisch-
naturwissenschaftliche Klasse, Abt. I,

236

118:876. 1909. =Rosenscheldia heliopsidis
(L.D. v. Schweinitz:E.M. Fries) F. v. Theissen
& H. Sydow, Annales Mycologici, Berlin
13:649. 1915. =Gibberidea heliopsidis (L.D.
v. Schweinitz:E.M. Fries) C.L. Shear, My-
cologia, Lancaster, Pennsylvania 29:361.
1937. =Syncarpella heliopsidis (L.D. v.
Schweinitz:E.M. Fries) M.E. Barr in M.E.
Barr and J.R. Boise, Memoirs of the New
York Botanical Garden, Bronx 49:300. 1989.
On stems of Aster sp., Helianthus sp.,
Heliopsis sp., Compositae. U.S.A.

helminthospora (V. Cesati) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:235. 1863.
=Sphaeria helminthospora V. Cesati in G.L.
Rabenhorst, Klotzschii Herbarium Vivum
Mycologicum Sistens Fungorum Per Totam
Germaniam Cresentium Collectionem
Perfectam, Dresden, Edition I, Century 18,
No. 1735. Anno 1853; Botanische Zeitung,
Berlin & Leipzig 11:236. 1853. On unknown
substrate of Inula crithmoides L., Compositae.
Germany.

helminthospora (V. Cesati) V. Cesati & G. de
Notaris forma crithmi-maritimi P.A. Sac-
cardo, Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:33.
1883. On dead stems of Crithmum maritimum
L., Umbelliferae. France.

heloniaefolia (M.C. Cooke & J.B. Ellis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:68. 1883. =Sphaeria (Foliicola) heloniaefo-
lia M.C. Cooke & J.B.Ellis, Grevillea,
London 8:16. 1879. On leaves of Helonias
bullata L., Liliaceae. U.S.A.

(Leptosphaerella) helvetica P.A. Saccardo & C.L.
Spegazzini in P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 1:399. 1878. On
upper surface of wilted leaves of Selaginella
helvetica (L.) Spring, Selaginellaceae. Italy.

helvetica P.A. Saccardo & C.L. Spegazzini forma
major H. Rehm, Ascomyceten, Fascicle 18,
No. 884. Anno 1886; Hedwigia, Dresden
26:93. 1887. On unknown substrate of
Selaginella helvetica (L.) Spring, Selaginel-
laceae. Italy.

hemerocallidis J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(2):155. 1901. [Validly published by
reference to an effectively published
description in Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg 1:252.
1899, as Leptosphaeria sp.| =Leptosphaeria
feltgeni P.A. Saccardo & P. Sydow, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 16:513. 1902. Nom.
illegit. Art. 64.1. On dry stems of Hemerocal-
lis sp., Liliaceae. Luxembourg.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

hemicrypta C.A. Oudemans, Nederlandsch kru-
idkundig Archief. Verslagen en Mededeelin-
gen der Nederlandsche Botanische Vereenig-
ing, Leiden, Series 2, 4:521—522. 1886. On
leaves of Carex sp., Cyperaceae. Nether-
lands.

(Clypeosphaeria) hendersoniae (J.B. Ellis) M.C.
Cooke, Grevillea, London 17:91. 1889.
=Sphaeria (Obtectae) hendersonia J.B. Ellis
in M.C. Cooke and J.B. Ellis, Grevillea,
London 6:14-15. 1877. =Clypeosphaeria
hendersoniae (J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:91.
1883. On canes of Rubus sp., Rosaceae.
U.S.A.

hendersoniae (L. Fuckel) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):26.
1957. Nom. illegit. Art. 64.1. Non Lepto-
sphaeria (Clypeosphaeria) hendersoniae
M.C. Cooke, Grevillea, London 17:91. 1889.
=Cucurbitaria hendersoniae L. Fuckel,
Symbolae Mycologicae, p. 172. 1870.
=Melanomma hendersoniae P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:109.
1883. On corticated or decorticated branches
of Salix sp., Salicaceae. Italy, Sweden.

hermodactyli T. Ferraris in T. Ferraris and C. Massa,
Annales Mycologici, Berlin 10:287. 1912. On
leaves of Hermodactylus tuberosus (L.)
Miller, Iridaceae. Italy.

herpotrichoides G. de Notaris, Sferiacei Italici, pp.
80-81. 1863. =Phaeosphaeria herpotrichoi-
des (G. de Notaris) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):115.
1957. On culms of Andropogon sp., Koeleria
cristata (L.) Pers., Gramineae. Finland, Italy.

hesperia M.E. Barr, Canadian Journal of Botany,
Ottawa 45:1044. 1967. =Linospora brunellae
J.B. Ellis & B.M. Everhart, Proceedings of the
Academy of Natural Sciences of Philadelphia
1894:337. 1894. [Anamorph: Phoma
brunellae (J.B. Ellis & B.M. Everhart) C.E.O.
Kuntze, Revisio Generum Plantarum 3:502.
1898.] =Ceuthocarpon brunellae (J.B. Ellis
& B.M. Everhart) A.N. Berlese, [cones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
2:149. 1899. On living and dying leaves of
Prunella vulgaris L., Labiatae. Canada,
U.S.A.

hesperidicola R. Picbauer, Bosanko-Khertsovachki
Musey Glasnik, Sarajevo 48:104. 1936. On
dead stems of Hesperis dinarica G. Beck,
Cruciferae. Yugoslavia.

heterospora (G. de Notaris) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:67. 1883 [as
Leptosphaeria heterospora (G. de Notaris) G.

March 1991

Niessl v. Mayendorf]. =Sphaeria heterospora
G. de Notaris, Sferiacei Italici, Century 1,
Fascicle 2, p. 65. 1863. =Byssothecium
heterosporum (G. de Notaris) F. v. Thiimen,
Mycotheca Universalis, Century 14, No.
1361. Anno 1879. On living, subterranean
rhizome of /ris germanica L., Iridaceae.
Czechoslovakia, Italy.

heterospora (G. de Notaris) G. Niessl v. Mayendorf,

Verhandlungen des Naturforschenden Vereins
in Briinn 10:173. 1872. =Sphaeria hetero-
spora G. de Notaris, Sferiacei Italici, p. 65.
1863. =Byssothecium heterosporum (G. de
Notaris) F. v. Thiimen, Mycotheca Univer-
salis, Century 14, No. 1361. Anno 1879.
=Trematosphaeria heterospora (G. de
Notaris) H.G. Winter, Dr. L. Rabenhorst’s
Kryptogaman-Flora von Deutschland,
Oesterreich und der Schweiz, Second edition,
1(2) Ascomyceten: Gymoasceen und Pyreno-
myceten, p. 277. 1885. =Phaeosphaeria
heterospora (G. de Notaris) J.R. Boise,
Mycologia, Lancaster, Pennsylvania 77:236.
1985. On living, subterranean rhizome of /ris
germanica L., Iridaceae. Czechoslovakia,
Italy.

heufleri (G. Niessl v. Mayendorf) P.A. Saccardo,

Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:83.
1883. =Sphaerella heufleri G. Niessl v.
Mayendorf, Verhandlungen des Naturforsch-
enden Vereins in Briinn 10:167. (1871) 1872.
=Mycotodea heufleri (G. Niess| v. Mayen-
dorf) W. Kirschstein, Annales Mycologici,
Berlin 34:201. 1936. On dry sporangia of
Polytrichum formosum Hedw., Musci (Poly-
trichaceae). Italy.

heveae A.M. Saccas, Agronomie Tropicale, Nogent-

sur-Marne 8:244. 1953. On living leaves of
Hevea brasiliensis Muell. Arg., Euphor-
biaceae. French Equatorial Africa.

hiemalis P.A. Saccardo & C.L. Spegazzini in P.A.

Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
1:395-396. 1878. =Phaeosphaeria hiemalis
(P.A. Saccardo & C.L. Spegazzini) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1527. 1989. On decaying stems of

Equisetum hyemale L. [as hiemalis|, Equiseta-

ceae. Canada, Italy, U.S.A.

hierochloae C.A. Oudemans, K. Akademie van

Wetenschappen Amsterdam, Afdeeling
Natuurkund Verslagen en Mededeelingen 3
Reeks, Deel 2:155. 1885 [as hierochloes).
=Phaeosphaeria hierochloae (C.A. Oude-
mans) O. Eriksson, Arkiv for Botanik,
Uppsala, Stockholm, New Series 6:424. 1967.
On unknown substrate of Hierochloé alpina
(Willd.) Roemer & Schultes, Gramineae.
U.S.S.R.

Nomenclator of Leptosphaeria 237

(Melanomma) hippophaes (J.H. Fabre) M.C. Cooke,

Grevillea, London 17:91. 1889. =Mela-
nomma hippophaes J.H. Fabre, Annales des
Sciences Naturelles, Paris, Botanique, Series
6, 9:92—93. 1878. On dry branches of
Hippophae rhamnoides L., Elaeagnaceae.
France.

hippophaes (A. Sollmann) A. Nannizzi, Atti dell’

Accademia dei Fisiocritici di Siena, Series 10,
5:385. 1931 [as hipphophaes). =Sphaeria
hippophaes A. Sollmann, Botanische Zeitung,
Berlin & Leipzig 20:379-380. 1862. =Cla-
dosphaeria hippophaes (A. Sollmann) T.R.J.
Nitschke in G.H. Otth, Mitteilungen der
Naturforschenden Gesellschaft in Bern
1870:105. 1871. =Massaria hippophaes (A.
Sollmann) A.L. Jaczewski, Bulletin de

l’ Herbier Boissier, Geneve and Bale 2:684.
1894. On dead branches of Hippophae
rhamnoides L., Elaeagnaceae. Germany,
Italy, Switzerland.

hippophaes (A. Sollmann) P.A. Saccardo, Sylloge

Fungorum Omnium Hucusque Cognitorum

hippopphaes]. Nom. illegit. Art. 63.1.
=Sphaeria hippophaes A. Sollmann, Botanis-
che Zeitung, Berlin & Leipzig 20:379-380.
1862. On dead corticated primocanes of
Hippophae rhamnoides L., Elaeagnaceae.
Germany.

hirta (E.M. Fries:E.M. Fries) V. Cesati & G. de

Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:236. 1863.
=Sphaeria hirta E.M. Fries:E.M. Fries,
Kungliga Svenska Vetenskapsakademiens
Handlingar, Uppsala and Stockholm
1818:105. 1818; Systema Mycologicum
Sistens Fungorum 2:483. 1823 [as Sphaeria
(Cyt.) hirta]. =Nodulosphaeria hirta (E.M.
Fries:E.M. Fries) G.L. Rabenhorst, Klotzschii
Herbarium Vivum Mycologicum Sistens
Fungorum Per Totam Germaniam Cresentium
Collectionem Perfectam, Dresden, Edition II,
Century 8, No. 725. Anno 1858. Nom. rejic.
On dead stems of Sambucus racemosa L.,
Caprifoliaceae. Germany, Sweden.

hirta (L. Fuckel) W. Kirschstein, Annales Mycol-

ogici, Berlin 34:188—189. 1936. Nom. illegit.
Art. 63.1. =Massaria hirta L. Fuckel,
Symbolae Mycologicae, p. 155. 1870. On
dead thin branches of Sambucus racemosa L.,
Caprifoliaceae. Germany.

hispanica J. Checa & G. Moreno, Canadian Journal

of Botany, Ottawa 65:2096-2097. 1987. On
twigs of Artemisia vulgaris L., Compositae.
Spain.

hollosiana L.E. Wehmeyer, Mycologia, Lancaster,

Pennsylvania 55:319. 1963. =Leptosphaeria
maritima L. Hollés. Non Leptosphaeria
maritima J.B. Ellis & B.M. Everhart. Non

i)
wo
oo

Leptosphaeria maritima P.A. Saccardo. On
stems of Biebersteinia emodii Jaub. & Spach.,
Potentilla argyrophylla Wallich ex Lehm.,
Potentilla fragarioides L., Triglochin
maritima L., Trisetum spicatum (L.) K.
Richter, Geraniaceae, Gramineae, Juncaceae,
Rosaceae. India.

hollosti G. Moesz, Magyar Botanikai Lapok,
Budapest 28:54. 1929. =Leptosphaeria
tamaricis L. Hollos, Botanikai K6zlemények,
Budapest 25:130. 1928. Non Leptosphaeria
tamaricis (R.K. Greville) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:26.
1883. On dried branches of Tamarix gallica
L., Tamaricaceae. Hungary.

holmii M.T. Lucas, Memorias da Sociedade
Broteriana, Coimbra 21:36. 1970. On dead
culms of Scirpus holoschoenus L., Cyper-
aceae. Portugal.

holmit (L.N. Vasil’eva) L.N. Vasil’eva, Pirenomit-
sety i Lokuloaskomitsety Severa Dal nego
Vostoka (Leningrad:“Nauka”), p. 122. 1987.
Nom. illegit. Art. 64.1. =Nodulosphaeria
holmii L.N. Vasil’eva, Mikologiia i Fitopa-
tologiia Akademiia Nauk SSSR, Leningrad
13:277-278. 1979. On stems of Lagotis
minor (Willd.) Standley, Scrophulariaceae.
WESISERE

honiaraensis T. Matsushima, Microfungi of the
Solomon Islands and Papua-New Guinea, pp.
73-74. 1975. On decayed leaf of a broadleaf
tree of unknown tree host, unknown family.
Solomon Islands.

hordei A.A. Domaschova in A.A. Domaschova and
N.A. Gamalitzkaja, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 15:75-76. 1962. On
unknown substrate of Hordeum turkestanicum
Nevski, Gramineae. U.S.S.R.

hottai K. Hara, Byéchu-gai Zasschi (Journal of Plant
Protection), Tokyo 6(4):37. 1918. Nom. nud.
Art. 32.1. On trunks of Thea sinensis L.,
Theaceae. Japan.

hottai K. Hara ex K. Hara, Chagyokai (Tea Journal)
14(9):14-15. 1919; Mycologia, Lancaster,
Pennsylvania 13:24. 1921. Parasitic on trunks
of Thea sinensis L., Theaceae. Japan.

houseana P.A. Saccardo, Annales Mycologici,
Berlin 13:116. 1915. On dead stems of
Thalictrum dioicum L., Ranunculaceae.
U.S.A.

hrubyana F., Petrak, Annales Mycologici, Berlin
25:290-291. 1927. On dry stems of Linum
tenuifolium L., Linaceae. Czechoslovakia.

hurae N.T. Patouillard, Bulletin de la Société
Mycologique de France, Paris 16:185—186.
1900. Ona stem of Hura crepitans L.
(=Manitia), Euphorbiaceae. Guadeloupe
(West Indies).

Illinois Natural History Survey Bulletin

Vol. 34 An. 3

huthiana R. Staritz, Verhandlungen des Botanischen
Vereins Der Provinz Brandenburg, Berlin
55:79-80. 1913. In leaf tissue of Sparganium
ramosum Hudson, Sparganiaceae. Germany.

hyalina K.S. Panwar & S.J. Kaur, Kavaka, Madras
3:68. 1975. On dead twigs of unknown host,
unknown family. India.

hyalospora P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:323. 1875. =Metasphaeria
hyalospora (P.A. Saccardo) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:179.
1883. On decayed stems of Zea mays L.,
Gramineae. Italy.

hydrangeae S.A. Gucevicz, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 7:161. 1970. On
dry branches of Hydrangea arborescens L..,
Saxifragaceae. U.S.S.R.

hydrophila P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:320—321. 1875. On canes
of Juncus effusus L., Juncaceae. Italy.

hyparrheniae C.G. Hansford, Proceedings of the
Linnean Society of London 153:24. 1941. On
dead stems of Hyparrhenia sp., Gramineae.
Uganda.

hyperborea (L. Fuckel) A.N. Berlese & P. Voglino,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo Addi-
tamenta Ad Volumina I-IV, p. 140. 1886.
=Pleosora hyperborea L. Fuckel, Pilze der
Zweiten Deutschen Nordpolfahrt 2:92. 1874.
=Mycotodea hyperborea (L. Fuckel) W.
Kirschstein, Annales Mycologici, Berlin
34:201. 1936. =Leptosphaerulina hyperborea
(L. Fuckel) M.E. Barr, Contributions from the
University of Michigan Herbarium, Ann
Arbor 9:542. 1972. On upper surface of dried
leaves of Andromeda tetragona L., Ericaceae.
Greenland, Norway.

hyperici H.G. Winter, Hedwigia, Dresden 11:147.
1872. On dry stems of Hypericum perforatum
L., Guttiferae. Germany.

hypericola S.A. Gucevicz, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 13:183—184. 1960.
On dried perennial parts of Hypericum
alpestris Stev. ex Ledeb., Guttiferae.
U.S.S.R. (Tauria).

hysterioides J.B. Ellis & B.M. Everhart, Bulletin of
the Washburn Laboratory of Natural History
[Washburn College], Topeka 1:4. 1884. On
dead leaves of Xerophyllum tenax (Pursh)
Nutt., Liliaceae. U.S.A.

icositana R.C. Maire, Bulletin de la Societe
d’Histoire Naturelle de |’Afrique du Nord,
Alger 8:170. 1917. On dry stems of Rubia
peregrina L., Rubiaceae. Algeria.

ae

March 1991

immunda P.A. Karsten, Mycologia Fennica Pars 2,
Pyrenomycetes, p. 105. 1873; Bidrag till
Kannedom om Findlands Natur och Folk.
Utgifna af Finska Vetenskaps-Societeten,
Helsingfors 23:105. 1873. =Metasphaeria
immunda (P.A. Karsten) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:170.
1883. On leaves of Populus tremula L.,
Salicaceae. Finland.

impressa (C.G.T. Preuss) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:57. 1883.
=Sphaeria impressa C.G.T. Preuss, Linnaea.
Institut botanique de |’ Université de Genéve
26:713-714. 1853. On pods of Cheiranthus
annuus L., Polypodiaceae. Germany.

inarensis (E.A. Vainio) K. Keissler, Annalen des
Naturhistorischen Museums in Wien 42:99.
1928. =Trypethelium inarense E.A. Vainio,
Societas pro Fauna et Flora Fennica,
Helsingfors Meddelanden 10:197—198. 1883.
=Xenosphaeria inarensis (E.A. Vainio) E.A.
Vainio, Acta Societatis pro Fauna et Flora
Fennica, Helsingforsiae 49(2):142-143. 1921.
=Sphaeria inarensis (E.A. Vainio) W.

Nylander, Flora, Jena und Regensburg 68:300.

1885. Nom. inval. Art. 33.1. Parasitic on
unknown lichen, Lichenes. Scandinavia.

incarcerata (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:86. 1883. =Sphaeria incarcerata M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:152. 1876. =Heptameria
(Leptosphaeria) incarcerata (M.J. Berkeley &
M.A. Curtis) M.C. Cooke, Grevillea, London
17:33. 1889. =Passeriniella incarcerata
(M.J. Berkeley & M.A. Curtis) A.N. Berlese,
Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:51. 1892. On stems of
Spartina sp., Gramineae. U.S.A.

inconspicua H. Rehm, Osterreichische Botanische
Zeitschrift, Wien 56:296. 1906. Nom. nud.
Art. 32.1. On ashes of Berberis sp., Berberi-
daceae. Austria.

incruenta W. Kirschstein, Annales Mycologici,
Berlin 37:108—109. 1939. On dry stems of
Clematis vitalba L., Ranunculaceae. Ger-
many.

inculta P.A. Saccardo & A.F. Malbranche in P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Illustrans
2:598. 1882. On stems of Chenopodium
album L., Chenopodiaceae. France.

indeprensa (M.C. Durieu de Maisonneuve & J.P.
Montagne) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:85. 1883. =Sphaeria indeprensa

Nomenclator of Leptosphaeria 239

M.C. Durieu de Maisonneuve & J.P.
Montagne in M.C. Durieu de Maisonneuve,
Exploration Scientifique de L’ Algérie Publiée
Par Ordre du Gouvernement. Series
Naturelles. Botanique I, p. 538. (1848) 1849.
On dead and withered leaves of Silene
rosulata Soyer-Willem. & Godr., Caryophyl-
laceae. Algeria.

indica H. Sydow, P. Sydow, & E. Butler, Annales
Mycologici, Berlin 9:409. 1911. On leaves
and stems of Asparagus sp., Liliaceae. India.

inecola K. Hara, Diseases of the Rice Plant, p. 157.
1918. On unknown substrate of Oryza sativa
L., Gramineae. Japan.

infernalis G. Niessl v. Mayendorf, Instituto. Revista
Scientifica e Litteraria, Coimbra 31:89. 1883.
=Montagnula infernalis (G. Niessl v.
Mayendorf), A.N. Berlese, Icones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 2:69.
1896. On dead leaves of Furcraea longaeva
Karw. & Zucc., Agavaceae. Portugal.

inquinans C.H. Peck, Bulletin of the New York State
Museum, Albany 131:22—23. 1909. On bark
of Acer saccharum auct., Aceraceae. U.S.A.

insignis P.A. Karsten, Ofversigt af K. Vetenskap-
sakademiens Forhandlingar, Stockholm
29(2):100. 1872. =Phaeosphaeria insignis
(P.A. Karsten) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):120. 1957. On
leaves of Aira alpina L., Alopecurus ovatus
Knapp, Dupontia fisheri R. Br., Dupontia
psilosantha (Rupr.) Griseb., Gramineae.
Canada, Norway.

insignis P.A. Karsten forma airae-cespitosae C.
Roumeguere, Fungi Selecti Gallici Exsiccati,
Century 58, No. 5754. Anno 1891; Revue
Mycologique, Toulouse 13:129. 1891. On
leaves of Aira cespitosa L., Gramineae.
France.

inspersa (L.D. v. Schweinitz) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:27—28. 1883.
=Sphaeria inspersa L.D. v. Schweinitz in F.
Currey, Transactions of the Linnean Society
of London 22:328. 1869. Non Sphaeria
inspersa M.J. Berkeley. On bark of unknown
host [L.D. v. Schweinitz specimen in W.J.
Hooker’s herbarium], unknown host,
unknown family. U.S.A.

insulana P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze, New Series 22:39. 1915. On
dead stems of Antirrhinum siculum Miller,
Scrophulariaceae. Yugoslavia.

intermedia (G. Niessl v. Mayendorf), Hedwigia,
Dresden 26:46-48. 1887. On unknown
substrate of Calamagrostis sp., Gramineae.
Austria.

interspersa (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum

Digessit P.A. Saccardo 2:77. 1883.
=Sphaeria interspersa M.C. Cooke, Grevillea,
London 6:146. 1878. On sheaths of Zea mays
L., Gramineae. U.S.A. (Florida).

involucralis G, Passerini, Hedwigia, Dresden
12:140. 1873. [Falsely so cited by P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:172. 1883.] =Sphaeria involucralis G.
Passerini, Hedwigia, Dresden 12:140. 1873.
=Metasphaeria involucralis (G. Passerini)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:172. 1883. On involucre of Castanea sp.,
Fagaceae. Italy.

iridicola J.B. Lambotte & F. Fautrey, Revue
Mycologique, Toulouse 17:168. 1895. On
frozen leaves of /ris foetidissima L., Iridaceae.
France.

iridigena F. Fautrey in F. Fautrey and J.B. Lambotte,
Revue Mycologique, Toulouse 17:168—169.
1895. On dry leaves of /ris pseudacorus L.,
Iridaceae. France.

iridigena F. Fautrey var. typhae J. Feltgen, Vor-
studien zu einer Pilz-Flora des Grossher-
zogthums, Luxemburg 1(3):206—207. 1903.
On dry leaves of Typha angustifolia L.,
Typhaceae. Luxembourg.

iridis L. Hollés, Annales Historico-Naturales Musei
Nationalis Hungarici, Budapest 5:45—46.
1907. On dry leaves of /ris arenaria Waldst.
& Kit., Iridaceae. Hungary.

irrepta G. Niessl v. Mayendorf in G. Linhart, Fungi
Hungarici Exsiccati, Century 4, No. 368.
Anno 1885. On unknown substrate of Cycas
revoluta Thunb., Cycadaceae. Hungary.

isariphora (J. Desmaziéres) B. Auerswald in W.
Gonnermann and G.L. Rabenhorst, Mycologia
Europaea, Abbildungen Sammtlicher Pilze
Europa’s, Hefte 5 und 6, Synopsis Pyrenomy-
cetum Europaeorum, Table 12, Figure 170.
1869. =Sphaeria isariphora J. Desmazieres,
Annales des Sciences Naturelles, Paris,
Botanique, Series 2, 19:358. 1843; Plantes
Cryptogames Du Nord De La France, Edition
1, Fascicle 26, No. 1291. Anno 1843.
=Sphaerella isariphora (J. Desmaziéres) G.
de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:237. 1863.
On dry leaves of Stellaria graminea L.,
Caryophyllaceae. France, Germany.

ischaemi G. Passerini, Erbario Crittogamico Italiano,
Series 2, Fascicle 22, No. 1073. Anno 1881.
On dry culms of Andropogon ischaemum L.,
Gramineae. Italy.

isocellula K.S. Panwar & S.J. Kaur, Kavaka, Madras
5:41. 1977. On dead wood of Lantana
camara L., Verbenaceae. India (Mt. Abu,
Rajasthan).

iwamotoi I. Miyake, Journal of the College of
Agriculture, Imperial University of Tokyo

Illinois Natural History Survey Bulletin

Vol. 34 Ar. 3

2:249. 1910. On leaves of Oryza sativa L.,
Gramineae. Korea.

jaceae L. Holm, Svensk Botanisk Tidskrift,
Stockholm 42:35-36. 1952. =Nodulosphaeria
Jaceae (L. Holm) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):86.
1957. On dead stems of Centaurea jacea L.,
Compositae. Sweden.

Jacksonensis R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2704-2705. 1984.
=Leptosphaeria tetonensis L.E. Wehmeyer.
On old stems of Pedicularis bracteosa Benth.
ex Hook., Scrophulariaceae. U.S.A.

Jacksonti R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2705. 1984. On stems of
Eupatorium maculatum L. and E. purpureum
L., Compositae. Canada.

Jahnii C.E. Chardon, Boletin de la Sociedad
Venezolana de Ciencias Naturales, Caracas
5(40):263-264. 1939. [Extract pp. 353—354.]
=Phaeophragmocauma jahnii (C.E. Chardon)
E. Miller in E. Miiller & R.W.G. Dennis,
Kew Bulletin. Royal Botanic Gardens, Kew
19(3):376. 1965. On living leaves of Espeletia
neriifolia Sch. Bip. ex Wedd., Compositae.
Venezuela.

Janus (M.J. Berkeley & M.A. Curtis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:85.
1883. =Sphaeria janus M.J. Berkeley & M.A.
Curtis, Grevillea, London 4:154. 1876.
=Metasphaeria janus (M.J. Berkeley & M.A.
Curtis) A.N. Berlese, Icones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:146.
1894 [as Metasphaeria junci (Oud.) Sacc.].
=Dothidella janus (M.J. Berkeley & M.A.
Curtis) F. v. HOhnel, Sitzungsberichte der
Akademie der Wissenschaften in Wien,
Mathematisch-naturwissenschaftliche Klasse,
Abt. I, 124:68. 1915. =Mycosphaerella janus
(M.J. Berkeley & M.A. Curtis) F. Petrak,
Sydowia. Annales Mycologici, Horn, N.O.
11:340. (1957) 1958. On leaves of Quercus
virens Ait. (=Q. virginiana Mill.), Fagaceae.
U.S.A.

Japonica (I. Miyake) K. Hara, Report of the
Sericultural Association of Japan, Tokyo
27(314):226. 1918. =Massaria japonica I.
Miyake, Sangyo Shikenjo Hokoku (Bulletin of
the Sericultural Experiment Station) 1(5):319.
1915. On Morus sp., Moraceae. Japan.

Jasmini, Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 15:191.
1901. [Attributed to L. Fuckel by P.A.
Saccardo in Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
3:309. 1884, cited L. Fuckel, Symbolae
Mycologicae Zweiter Nachtrag, p. 24, 1873.
The fungus listed there is Pleospora jasmini
(J.L. Castagne) L. Fuckel.]

March 1991 Nomenclator of Leptosphaeria 241

johansonii E. Miller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):238. 1950 [as johansoni).
=Metasphaeria arabidis C.J. Johanson,
Hedwigia, Dresden 25:122. 1886. Non
Leptosphaeria arabidis A. Allescher. On dry
stems of Arabis alpina L., Cruciferae.
Switzerland.
jubaeae C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cordoba 25:71—72.
1921. On dead or decaying leaves of Jubaea
spectabilis Humb., Bonpl., & Kunth, Palmae.
Chile.
juncaginearum (J. Schroter) A. Munk, Dansk
Botanisk Arkiv, Kj@benhavn 17(1):377. 1957.
[Basionym not completely cited.] =Myco-
sphaerella juncaginearum J. Schroter in F.J.
Cohn, Kryptogamen-Flora Von Schlesien. Im
Namen Der Schlesischen Gesellschaft fiir
vaterlandische Cultur herausgegeben von
Prof. Dr. Ferdinand Cohn, Secretair der
Botanischen Section, Breslau 3(2):342. 1894.
=Phaeosphaerella juncaginearum (W.G.
Lasch & J. Schroéter) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 11:312. 1895. On
dead stems of Triglochin maritima L.,
Triglochin palustris L., Juncaginaceae.
Germany.
junci (C.A. Oudemans) J. Lind, Danish Fungi as
Represented in the Herbarium of E. Rostrup,
Nordisk Forlag, Copenhagen, p. 217. 1913.
=Sphaeria junci C.A. Oudemans, Archives
Néerlandaises des Sciences Exactes et
Naturelles, Haarlem 8:405. 1873 [as Sphaeria
junci Fr.(?)). =Metasphaeria junci P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:177. 1883. On leaves and culms of Juncus
glaucus Sibth., Juncaceae. Netherlands.
junci J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(2):156.
1901. On culms of Juncus effusus L.,
Juncaceae. Luxembourg.
juncicola H. Rehm in H. Rehm, Ascomyceten,
Fascicle 11, No. 533. Anno 1879. Nom. nud.
Art. 32.1. On dry culms of Juncus trifidus L.,
Juncaceae. Italy.
juncicola H. Rehm ex H.G. Winter in H.G. Winter,
Hedwigia, Dresden 19:167. 1880. =Lepto-
sphaeria juncicola H. Rehm, Ascomyceten,
Fascicle 11, No. 533. 1879. Nom. nud. Art.
32.1. =Phaeosphaeria juncicola (H. Rehm ex
H.G. Winter) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):129. 1957. On
dried culms of Juncus hostii Tausch, Juncus
trifidus L., Juncaceae. Austria, Czechoslova-
kia, Italy, Switzerland.
juncicola H. Rehm ex H. Rehm, Bericht des
Naturwissenschaftlichen Vereins fiir Schwa-
ben und Neuberg, Augsburg 26:128. 1881.
On dried culms of Juncus hostii Tausch,
: Juncaceae. Italy.

juncina (B. Auerswald) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:66. 1883.
=Sphaeria juncina B. Auerswald in G.L.
Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio Edita Nova, Series Secunda, Century 8,
No. 748. Anno 1865. =Sphaerella juncina (B.
Auerswald) B. Auerswald in W. Gonnermann
and G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Sammtlicher Pilze Europa’s,
Hefte 5 und 6, Synopsis Pyrenomycetum
Europaeorum, p. 18. 1869. =Mycotodea
juncina W. Kirschstein, Annales Mycologici,
Berlin 37:134. 1939. =Phaeosphaeria
juncina (B. Auerswald) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):127.
1957. On dry culms of Juncus effusus L.,
Juncaceae. Czechoslovakia, Germany, Great
Britain, Portugal.

Juncina (B. Auerswald) P.A. Saccardo forma
macrospora G. Niessl v. Mayendorf in V.
Mouton, Bulletin. Société R. de Botanique de
Belgique, Bruxelles 39:43-44. 1900. On
veins of leaves of Luzula albida (Hoffm.)
DC., Juncaceae. Belgium.

junciseda P.A. Karsten, Ofversigt af K. Vetenskaps-
akademiens Forhandlingar, Stockholm
29(2):101. 1872. On dead plant of Juncus
biglumis L., Juncaceae. Norway (Spits-
bergen).

junci-acuti R. Gonzalez Fragoso, Asociacién
Espanola Para el Progresso de las Ciencias,
Congreso de Coimbra 1925:12. 1925. Nom.
inval. Art. 34.1. On dry culms of Juncus
acutus L., Juncaceae. Portugal.

junci-glauci L.M. Unamuno Yrigoyen, Boletin de la
R. Sociedad Espanola de Historia Natural,
Madrid 29:394. 1929. On dry stems of Juncus
glaucus Sibth., Juncaceae. Spain.

Juncorum (P.L. Crouan & H.M. Crouan) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:86. 1883. =Pleospora juncorum P.L.
Crouan & H.M. Crouan, Florule Du Finistere
Contenant Les Descriptions De 360 Espeéces
Nouvelles De Sporogames, De Nombreuses
Observations et une synonymie des plantes
Cellulaires et Vasculaires Qui Croissent
Spontanément Dans ce Département, p. 22.
1867. On dead stems of Juncus acutiflorus
Ehrh. ex Hoffm., Juncaceae. France.

Juniperi F. Fautrey, Revue Mycologique, Toulouse
17:70. 1895. On branchlets at ends of dry
branches of Juniperus communis L., Cupres-
saceae. France.

juniperina F.A. Hazslinszky, Matematikai és
Természettudomanyi K6zlemenyek Von-
atkozolag a Hazai Viszonyokra, Budapest
25(2):150. 1892. On leaves of Juniperus
nana Willd., Cupressaceae. Hungary.

242

kali J.H. Fabre, Annales des Sciences Naturelles,
Paris, Botanique, Series 6, 15:50. 1883.
=Metasphaeria kali (J.H. Fabre) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2(Addenda Ad Volumen Secundum):LXI.
1883. On dry stems of Salsola kali L.,
Chenopodiaceae. France.

kalmiae C.H. Peck, Report. New York State
Museum of Natural History, Albany 39:53.
(1885) 1886. =Gibberidea kalmiae (C.H.
Peck) M.E. Barr, Canadian Journal of Botany,
Ottawa 39:311. 1961. On dead stems of
Kalmia angustifolia L., Ericaceae. U.S.A.

kalmusti G. Niessl v. Mayendorf in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:44.
1883. On stems of Artemisia vulgaris L.,
Compositae. Czechoslovakia.

kerguelensis P.C. Hennings in E. v. Drygalski,
Deutsche Suedpolar Expedition 1901-1903,
8:11. 1906. On leaves of Festuca kerguelen-
sis Hook. f., Gramineae. Kerguelen Island
(near Antarctica).

kochiana E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 5(3-6):50. 1951. On dead reeds
of Carex baldensis L., Cyperaceae. Italy.

koerberi (B. Stein) H.G. Winter, Dr. L. Rabenhorst’s
Kryptogaman-Flora von Deutschland,
Oesterreich und der Schweiz 1(2):443. 1885.
=Leptorhaphis koerberi B. Stein in F. Cohn,
Kryptogamen-Flora Von Schlesien. Im
Namen Der Schlesischen Gesellschaft fiir
vaterlandische Cultur herausgegeben von
Prof. Dr. Ferdinand Cohn, Secretair der
Botanischen Section, Breslau 2(2):350. 1879.
=Ophiobolus koerberi (B. Stein) A.N. Berlese
& P. Voglino, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
Additamenta Ad Volumina I-IV, p. 190.
1886. On thallus of Koerberiella wimmeriana
(Koerber) B. Stein, Lichenes (Porpidiaceae).
Czechoslovakia, Germany.

korrae J.C. Walker & A.M. Smith, Transactions of
the British Mycological Society, London
58:461. 1972. =Ophiosphaerella korrae (J.C.
Walker & A.M. Smith) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1591.
1989. On stolons and sheaths of Avena sativa
L., Axonopsus compressus (Swartz) Beauv.,
Cynodon dactylon (L.) Pers., Eremochloa
ophiuroides (Munro) Hack., Oryza sativa L.,
Triticum aestivum L., Gramineae. Australia
(New South Wales), U.S.A.

kotschyana F. Petrak, Annalen des Naturhistorischen
Museums in Wien 50:433. 1939. On leaves of
Acantholimon acerosum Boiss., Acantholimon
melananthum Boiss., Plumbaginaceae. Iran.

kuangfuensis J. Yen & S.K. Sun, Mycotaxon. An
International Journal Designed to Expedite

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens, Ithaca,
New York 7:396-397. 1978. On living
branches of Saccharum officinarum L.,
Gramineae. Taiwan.

kunzeana A.N. Berlese, Icones Fungorum Omnium
Hucusque Cognitorum ad usum Sylloges
Saccardianae Adcommodatae 1:66. 1890.
=Leptosphaeria typharum G. Kunze. Nom. J.
Desmaziéres. On stems of Typha latifolia L.,
Typhaceae. Germany.

lacustris (L. Fuckel) H.G. Winter, Dr. L.
Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2), p. 451. 1885; Hedwigia,
Dresden 26:57. 1887. =Sphaeria lacustris L.
Fuckel, Fungi Rhenani Exsiccati A Leopoldo
Fuckel Collecti, Fascicle 25(10), No. 2436.
Anno 1872; Symbolae Mycologicae Zweiter
Nachtrag, p. 22. 1873. =Metasphaeria
lacustris (L. Fuckel) P.A. Saccardo, Syllogue
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:173. 1883. =Mas-
sarina lacustris (L. Fuckel) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:181. 1984. =Wertsteinina lacustris (L.
Fuckel) R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 67:1595. 1989. On leaves
and stems of Schoenoplectus lacustris (L.)
Palla, Typha angustifolia L., Typha latifolia
L., Cyperaceae, Typhaceae. Germany,
Switzerland.

ladina E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):293. 1950. =Nodu-
losphaeria ladina (E. Miller) L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):83. 1957. On dead stems of Laser-
pitium halleri Crantz, Umbelliferae. Switzer-
land.

lagenoides C.L. Spegazzini, Anales del Museo
Nacional de Historia Natural de Buenos Aires
23:57. 1912; Mycetes Argentinenses, Series 6,
No. 1377. 1912. On dry decaying sheaths of
Elymus? barbatus F. Kurtz., Gramineae.
Argentina.

lamprocarpi (G. Passerini) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:66. 1883. =Sphae-
rella lamprocarpi G. Passerini, Erbario
Crittogamico Italiano, Series 2, Fascicle 8,
No. 392. Anno 1871. On florets and bracts of
Juncus lampocarpus Ehbrh. ex Hoffm.,
Juncaceae. Italy.

lankeana P.C. Hennings, Verhandlungen des
Botanischen Vereins Der Provinz Bran-
denburg, Berlin 47:220-221. 1906. On last
year’s stems of Hypericum perforatum L.,
Guttiferae. Sri Lanka.

larseniana A. Munk, Dansk Botanisk Arkiv,
Kjobenhavn 14(7):27. 1952. =Leptosphaeria

March 1991

elymi P. Larsen. Non G.F. Atkinson, 1897.
=Phaeosphaeria larseniana (A. Munk) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1568. 1989. On stems and leaves
of Baldingera arundinacea (L.) Dumortt.,
Elymus arenarius L., Iris pseudacorus L.,
Phragmites communis Trin., Gramineae,
Iridaceae. Iceland, U.S.A.

larvalis P.A. Saccardo, Boletim da Sociedade
Broteriana, Coimbra 21:213. 1905. On dead
culms of Equisetum [as E. pallidum],
Equisetaceae. Cape Verde Islands.

lasioderma J.B. Ellis & B.M. Everhart, Proceedings
of the Academy of Natural Sciences of

Philadelphia 1893:135. 1893. =Dothiodotthia
lasioderma (J.B. Ellis & B.M. Everhart) M.E.

Barr, Mycotaxon. An International Journal

Designed to Expedite Publication of Research

on Taxonomy & Nomenclature of Fungi &
Lichens, New York 34:523. 1989. On living

leaves of Artemisia tridentata Nutt., Composi-

tae. U.S.A.

lasiosphaerioides K. Starbaéck & A.Y. Grevillus in
K. Starback, Bihang till K. Svenska
Vetenskapsakademiens Handlingar, Stock-
holm 16(3):7. 1890; Bidrag till Kannedomen
om Sveriges Ascomycetflora, p. 7. 1890. On
dry stems of Aconitum lycoctonum L.,
Ranunculaceae. Sweden.

lassenensis R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 67:1578. 1989. On stems of
Allium validum S. Wats., Liliaceae. U.S.A.
(California).

latebrosa J.B. Ellis in N.L. Britton, Catalogue of
Plants Found in New Jersey. Geological
Survey of New Jersey, Final Report of the
State Geologist, Trenton 2(1):524. 1889.
Nom. illegit. Art. 64.1. =Leptosphaeria
latebrosa (J.B. Ellis) P.A. Saccardo.

latebrosa (J.B. Ellis) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:86. 1883.

=Sphaeria latebrosa J.B. Ellis, Bulletin of the

Torrey Botanical Club (and Torreya), New
York 8:90. 1881. On basal sheaths of
Andropogon sp., Gramineae. U.S.A.

lathonia P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:310. 1875. On leaves of
Helleborus viridis L., Ranunculaceae. Italy.

lathonia P.A. Saccardo var. hellebori-foetidi C.B.
Massalongo, Atti del Istituto Veneto di
Scienze, Lettere ed Arti, Venezia 74:254.
1914. On leaves of Helleborus foetidus L.,
Ranunculaceae. Italy.

lathyri F. Fautrey, Fungi Selecti Gallici Exsiccati,
Century 56, No. 5537. Anno 1891; Revue

| Mycologique, Toulouse 13:8. 1891. On stems

of Lathyrus sylvestris L., Leguminosae.
France.

Nomenclator of Leptosphaeria 243

lathyrina P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):153—154. 1873. On dead stems of
Lathyrus latifolius L., Leguminosae. Italy.

lathyri P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):154. 1873. =Metasphaeria lathyri (P.A.
Saccardo) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:159. 1883. On fallen dead stems
of Lathyrus sylvestris L., Leguminosae. Italy.

lauri A. Maublanc, Bulletin de la Société Mycolo-
gique de France, Paris 21:89. 1905. On leaves
of Laurus nobilis L., Lauraceae. France.

lavandulae M. de Sousa da Camara, Agronomia
Lusitana, Sacavém 13:126. 1951. On
branchlets of Lavandula stoechas L.,
Labiatae. Portugal.

lecanora H. Fabre, Annales des Sciences Naturelles,
Paris, Botanique, Series 6, 15:49. 1883. On
dry stems of Salsola kali L., Chenopodiaceae.
France.

leersiae G. Passerini, Atti della Societa Crittogamol-
ogica Italiana, Milano, Series 2, 2:43-44.
1879. =Leptosphaeria leersiana P.A.
Saccardo nec (ut videtur) Sphaerella (Lepto-
sphaerella) leersiae G. Passerini. On dried
leaves of Leersia oryzoides (L.) Swartz,
Gramineae. Italy.

leersiana P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:60. 1883. =Leptosphaeria
leersiae G. Passerini. Non Sphaerella
(Leptosphaerella) leersiae G. Passerini. On
dried leaves of Leersia oryzoides (L.) Swartz,
Gramineae. Italy.

leiostega (J.B. Ellis) J.B. Ellis, Catalogue of Plants
Found in New Jersey. Geological Survey of
New Jersey, Final Report of the State
Geologist, Trenton 2(1):525. 1889.
=Sphaeria (Leptosphaeria) leiostega J.B.
Ellis, Bulletin of the Torrey Botanical Club
(and Torreya), New York 8:91. 1881. On
various dead twigs and limbs of Carya sp.,
Rosa sp., Vaccinium sp., Juglandaceae,
Rosaceae, Ericaceae. U.S.A. (New Jersey).

lejostega (J.B. Ellis) F. Hazslinszky, Matematikai és
Természettudomanyi K6zlemenyek Von-
atkozdlag a Hazai Viszonyokra, Budapest
25(2):148. 1892. =Sphaeria (Leptosphaeria)
leiostega J.B. Ellis, Bulletin of the Torrey
Botanical Club (and Torreya), New York
8:91. 1881. =Metasphaeria lejostega (J.B.
Ellis) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:164. 1883. On branches of Carya
sp., Rosa sp., Vaccinium sp., Juglandaceae,
Rosaceae, Ericaceae. U.S.A. (New Jersey).

lelebae |. Hino & K. Katumoto, Bulletin of the
Faculty of Agriculture, Yamaguchi Univer-

244

sity, Shimonoseki 9:904. 1958. On dead
culms of Leleba sp. [as L. simplex], Gramin-
eae. Japan.

lemaneae (F. Cohn) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:84. 1883.
=Sphaeria lemaneae F.J. Cohn in A. de Bary
and M.S. Woronin, Beitrag zur Morphologie
und Physiologie der Pilze 3:1—7. 1870. On
thalli of Lemanea fluviatilis C. Aq., Algae
(Rhodophyceae-Lemaneaceae). Germany.

lemoinii C. Richon, Catalogue Raisonné des
Champignons Qui Croissent Dans le Départe-
ment de la Marne, p. 300. 1889. On an
unknown monocotyledon host, unknown
family. France.

leptogiophila (Minks) H.G. Winter, Dr. L.
Rabenhorst’s Krypytogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):443. 1885 [as
Leptosphaeria?|. =Leptorhaphis leptogio-
phila Minks in H.G. Winter, Flora, Jena und
Regensburg 60:211. 1877. =?Metasphaeria
leptogiophila (Minks) A.N. Berlese & P.
Voglino, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
Additamenta Ad Volumina I-IV, p. 158.
1886. On thalli of Leptorhaphis leptogiophila
Minks ex Wint., Lichenes (Arthropyreni-
aceae). Germany.

leptospora (G. de Notaris) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:16. 1883. =Cucur-
bitaria leptospora G. de Notaris, Sferiacei
Italici, Century I, Fascicle 2, pp. 62-63. 1863.
On stems of Dianthus atrorubens All.,
Caryophyllaceae. Italy.

lespedezae Ziling, Trudy Botanicheskogo Instituta
Akademiy Nauk SSSR, Leningrad, Moskva,
Series 2, 3:680. 1936. On dead stems of
Lespedeza bicolor Turcz., Leguminosae.
U.S.S.R.

lethalis J.B. Ellis & B.M. Everhart, Proceedings of
the Academy of Natural Sciences of Philadel-
phia 1895:423. 1895. On dead stems of
Penstemon confertus Doug}. ex Lindl.,
Scrophulariaceae. U.S.A.

leucomelaria (W. Mudd) L. Vouaux, Bulletin de la
Société Mycologique de France, Paris 29:121.
1913. =Sphaeria leucomelaria W. Mudd, A
Manual of British Lichens, Containing
Descriptions of All Species and Varieties, and
Five Plates With Figures of the Spores of One
Hundred and Thirty Species Illustrative of the
Genera, p. 105. 1861. Nom. inval. Art. 34.1b.
=Tichothecium leucomelarium (W. Mudd)
A.N. Berlese & P. Voglino, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo Additamenta Ad
Volumina I-IV, p. 121. 1866. Parasitic on

Illinois Natural History Survey Bulletin

thallus of Borrera leucomela (L.) Mudd
[=Anaptychia leucomela (L.) Mass.}, Lichenes
(Physciaceae). Great Britain.

leucoplaca (K.F.W. Wallroth) E.A. Vainio, Acta
Societatis pro Fauna et Flora Fennica,
Helsingforsiae 49(2):143-144. 1921.
=Verrucaria leucoplaca K.F.W. Wallroth,
Flora Cryptogamica Germaniae 1:299. 1831.
=Pyrenula leucoplaca (K.F.W. Wallroth)
G.W. Koerber, Systema Lichenum Germa-
niae, p. 361. 1855. =Verrucaria alba H.A.
Schrader var. leucoplaca (K.F.W. Wallroth)
L.E. Schaerer in Enumeratio Critica Lich-
enum Europaeorum, Quos Ex Nova Methodo
Digerit, p. 219. 1850. =Eopyrenula leuco-
placa (K.F.W. Wallroth) R.C. Harris, The
Michigan Botanist, Ann Arbor 12:19. 1973.
On bark of Acer sp., Alnus sp., Carya sp.,
Corylus sp., Fraxinus sp., Juglans sp.,
Populus sp., Quercus sp., Salix sp., Sorbus
sp., Tilia sp., Ulmus sp., Aceraceae, Juglan-
daceae, Betulaceae, Oliaceae, Salicaceae,
Fagaceae, Rosaceae, Tiliaceae, Ulmaceae.
Finland.

libanotis (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:16—-17. 1883.
=Pleospora libanotis L. Fuckel, Symbolae
Mycologicae Zweiter Nachtrag, p. 24. 1873.
On dry stems of Libanotis montana Crantz,
Umbelliferae. Belgium, France, Switzerland.

licatensis P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:70. 1883. =Leptosphaeria
typhicola G. Passerini & V. Beltrani. 1883.
Nec Leptosphaeria typhicola P.A. Karsten.
=Phaeosphaeria licatensis (P.A. Saccardo)
R.A. Shoemaker, Canadian Journal of Botany,
Ottawa 67:1532. 1989. On dry leaves of
Typha latifolia L., Typhaceae. Canada, Italy.

licatensis P.A. Saccardo forma rupefortensis G.
Passerini in G. Passerini and P. Brunaud,
Revue Mycologique, Toulouse 8:206. 1886.
On dead leaves of Typha latifolia L.,
Typhaceae. France.

lichenicola F.W. Zopf, Hedwigia, Dresden 35:385.
1896; Nova Acta Academiae Caesareae
Leopoldino Carolinae Germanicae Naturae
Curiosorum, Halle 70:160—163. 1897:
Untersuchungen tiber die durch Parasitische
Pilze Hervorgerufenen Krankheiten der
Flechten, pp. 64-67. 1897. On thallus and
apothecia of Solorina crocea (L.) Ach.,
Lichenes (Peltigeraceae). Italy.

lilicola K. Hara, Byéchu-gai Zasschi (Journal of
Plant Protection), Tokyo 5:885-886. 1918.
On stems of Lilium sp., Liliaceae. Japan.

lili J.B. Ellis & J. Dearness in J.B. Ellis and B.M.
Everhart, Proceedings of the Academy of
Natural Sciences of Philadelphia 1893:136.

Vol. 34 Art. 3

March 1991

1893. On leaves of Lilium superbum L.,
Liliaceae. Canada.

limitata (C.H. Persoon) M.C. Cooke, Grevillea,
London 17:91. 1889. =Sphaeria limitata C.H.
Persoon in K.F.W. Wallroth, Flora Crypto-
gamica Germaniae 2:783. 1833 [as Sphaeria
limitata C.H. Persoon ined. ex spec. Kunze!
sub. Sphaeria mamillana, in E.M. Fries,
Elenchus Fungorum Sistens Commentarium
in Systema Mycologicum 2:99, 1828].
=Clypeosphaeria limitata (C.H. Persoon) L.
Fuckel, Symbolae Mycologicae, p. 117. 1870.
On dry branches of Cornus alba L., Cornus
sanguinea L., Rhamnus sp., Cornaceae,
Rhamnaceae. Germany.

limosa F. Fautrey in C. Roumeguére, Revue
Mycologique, Toulouse 13:128. 1891; Fungi
Selecti Gallici Exsiccati, Century 58, No.
5751. Anno 1891. On dry stems of Equisetum
limosum L., Equisetaceae. France.

lindquistii M.J. Frezzi, Revista de Investigaciones
Agropecuarias, Buenos Aires, Series 5,
Pathologia Vegetal 5(7):73—79. 1968. [Latin
description appended as an addendum to the
offprints. May not constitute valid publica-
tion!] On fallen stems of Helianthus annuus
L., Compositae. Argentina.

linearis (P.A. Saccardo) E. Miiller, Sydowia.
Annales Mycologici, Horn, N.O. 4(1-6):258.
1950. =Leptosphaeria culmifraga (E.M.
Fries:E.M. Fries) G. de Notaris var. linearis
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:75. 1883. On culms of Agropyron repens
(L.) Beauv., Milium effusum L., Poa
nemoralis L., Gramineae. France, Switzer-
land.

lineolaris G. Niessl vy. Mayendorf in G. Linhart, Bo-
tanisches Zentralblatt, Jena & Dresden
26:119-120. 1886; Fungi Hungarici Exsiccati,
Century 5, No. 466. Anno 1886.
=Phaeosphaeria lineolaris (G. Niessl v.
Mayendorf) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1545. 1989. On
hard culms of Aira cespitosa L., Gramineae.
Hungary.

lingue (C.L. Spegazzini) P.A. Saccardo & A. Trotter,

Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 22:219.
1913. Leptosphaerella ?lingue C.L. Spegaz-
zini, Fungi Chilenses, p. 77. 1910; Revista de
La Facultad de Agronomia Y Veterinaria,
Universidad Nacional de La Plata, Series 2,
6:77. 1910. On decaying and wilted leaves of
Persea lingue Nees, Lauraceae. Chile.

lithophilae S.A. Gucevicz, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 13:196. 1960. On
dry stems of Sobolewskia lithophila Bieb.,
Cruciferae. U.S.S.R.

Nomenclator of Leptosphaeria 245

littoralis P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:38. 1877. =Phaeosphaeria
littoralis (P.A. Saccardo) L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):121. 1957. On culms of Scirpus sp.,
Cyperaceae. Italy.

littoralis P.A. Saccardo forma calamagrostidis-
arenariae P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:78. 1883. On culms and rhizomes
of Ammophila arenaria (L.) Link, Scirpus sp.,
Cyperaceae, Gramineae. Germany, Italy.

livida P. Voglino, Atti della R. Accademia
D’Agricoltura di Torino 53:352-353. 1911.
On somewhat living leaves of unknown host,
unknown family. Italy.

lobayensis A.M. Saccas, Etude de la Flore Cryptoga-
mique des Caféiers en Afrique Centrale.
Bulletin de Institut Francais du Café du Cacao
et d’Autres Plantes Stimulantes (Bulletin
IFCC No. 16), pp. 251-253. 1981. Nom.
inval. Art. 37.1. On dead stems of Coffea
excelsa Cheval., Rubiaceae. Central African
Republic.

lobeliae K.H. Anahosur, Sydowia. Annales Mycol-
ogici, Horn, N.O. 24(1-6):286. (1970) 1971.
On living leaves of Lobelia excelsa Lesch.,
Campanulaceae. India.

lolti H. Sydow & P. Sydow, Hedwigia, Dresden
39:(1). 1900. On culms of Lolium perenne L.,
Gramineae. Germany.

longchampsi P.A. Saccardo, Revue Mycologique,
Toulouse 7:145. 1885 [as (West.) Sacc.; based
on Sphaeria longchampsi G.D. Westendorp in
E. Marchal, Cryptogames de la Belgique].
Nom. inval. Art. 29.1. [E. Marchal used the
above title in 1882 for the general distribution
of exchange specimens (D.H. Pfister,
Mycotaxon. An International Journal
Designed to Expedite Publication of Research
on Taxonomy & Nomenclature of Fungi &
Lichens, Ithaca, New York 23:123. 1885).]
On dry stems of Libanotis sp., Umbelliferae.
Belgium.

longipedicellata J.H. Miller & G. Burton, Mycolo-
gia, Lancaster, Pennsylvania 34:2—3. 1942.
On dead stems of herbaceous plants of
Daucus carota L., Smallanthus uvedalia (L.)
Mackenzie, Solidago caesia L., Compositae,
Umbelliferae. U.S.A. (Georgia).

longispora A.M. Saccas, Etude de la Flore Cryptoga-
mique des Caféiers en Afrique Centrale.
Bulletin de Institut Frangais du Café du Cacao
et d’Autres Plantes Stimulantes (Bulletin
IFCC No. 16), pp. 244-256. 1981. Nom.
inval. Art. 37.1. On stems and branches of
Coffea excelsa Cheval., Rubiaceae. Central
African Republic.

longispora J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg 1(2):159.

246

1901. On dry stems of /ris pseudacorus L.,
Iridaceae. Luxembourg.

lonicerae S.A. Gucevicz, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 7:161—162.
1970. Nom. illegit. Art. 64.1. On dry
branches of Lonicera canadensis Bartr.,
Caprifoliaceae. U.S.S.R.

lonicerae (F. Fautrey) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):46. 1957.
=Metasphaeria lonicerae F. Fautrey in C.
Roumegueére, Fungi Selecti Gallici Exsiccati,
Century 54, No. 5359. 1890. Parasitic on
branches of Caprifolium sp., Lonicera
xylosteum L., Caprifoliaceae. France,
Germany.

lonicerina (P.A. Karsten) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):22.
1957. =Sphaeria lonicerina P.A. Karsten,
Fungi Fenniae Exsiccati, Century 10, No. 971.
Anno 1870. =Melanomma lonicerinum (P.A.
Karsten) P.A. Saccardo; Michelia Commen-
tarlum Mycologicum Fungos in Primis
Italicos Illustrans 1:344. 1878. On stems of
Alnus sp., Salix sp., Betulaceae, Salicaceae.
Germany.

lophanthi (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:84-85. 1883. =Sphaeria lophanthi M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:152. 1876. On stems of
Lophanthus sp., Labiatae. U.S.A.

lucilla P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:310—311. 1875. Indis-
criminately on wilted leaves of Pyrus
communis L., Rosaceae. Italy.

lucina P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:311. 1875. On dead leaves
of Cytisus laburnum L., Leguminosae. Italy.

luctuosa G. Niessl v. Mayendorf in P.A. Saccardo,
Nuovo Giornale Botanico Italiano e Bolletino
della Societa Botanica Italiana, Firenze 7:321.
1875. =Phaeosphaeria luctuosa (G. Niessl v.
Mayendorf) Y. Otani & T. Mikawa, Memoirs
of the National Science Museum, Tokyo,
Japan [Kokuritsu Kagaku Hakubutsukan
Senho Memoirs] 4:78. 1971. On rotten culms
of Zea mays L., Gramineae. Canada, Italy,
U.S.A.

lunariae (M.J, Berkeley & C.E. Broome) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:57. 1883. =Sphaeria (Caulicolae) lunariae
M.J. Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
3, 3:374. 1859. On dried pods of Lunaria
rediviva L., Cruciferae. Great Britain.

lupini (F.S. Earle) J.R. Boise, On Trematosphaeria
(Loculoascomycetes, Fungi) and Disposition

Illinois Natural History Survey Bulletin

of Heterogenous Elements, Ph.D. dissertation,
University of Massachusetts, Amherst, p. 47.
1984. Nom. provis. Art. 34.1b. =Tremato-
sphaeria lupini F.S. Earle in E.L. Greene,
Plantae Bakerianae 2(1):17. 1901. On stems
of Lupinus sp., Leguminosae. U.S.A.
(Colorado).

lupinicola F.S. Earle in E.L. Greene, Plantae
Bakerianae 2:20. 1901. On dead stems of
Lupinus sp., Leguminosae. U.S.A.

lusitanica F. vy. Thiimen, Instituto. Revista Scienti-
fica e Litteraria, Coimbra 28:29-30. (1880)
1881; Contributiones ad Floram Mycologicam
Lusitanicam, Series 3, pp. 29-30. 1881. On
dry stems of Spartium junceum L., Legumino-
sae. Portugal.

luxemburgensis P.A. Saccardo & D. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 17:723.
1905. [Superfluous name for Leptosphaeria
dumetorum J. Feltgen. Non G. Niessl v.
Mayendorf, fide P.A. Saccardo, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 17:723. 1905.
Leptosphaeria dumetorum was attributed to
G. Niessl v. Mayendorf; the variety symphyti
was published by J. Feltgen as Leptosphaeria
dumetorum G. Niessl vy. Mayendorf var.
symphyti J. Feltgen.]

luxemburgensis P.A. Saccardo & D. Saccardo var.
dolichospora (J. Feltgen) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 17:723.
1905. =Leptosphaeria dumetorum G. Niessl
v. Mayendorf var. dolichospora J. Feltgen,
Vorstudien zu einer Pilz-Flora des Grossher-
zogthums, Luxemburg, Nacht. 3, p. 210.
1903. On branches of Sambucus racemosa L.
[as S. nigra], Caprifoliaceae. Luxembourg.

luxemburgensis P.A. Saccardo & D. Saccardo var.
symphyti J.Feltgen [falsely so cited in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
17:723. 1905.] =Leptosphaeria dumetorum
G. Niessl v. Mayendorf var. symphyti J.
Feltgen.

luzulae H.G. Winter, Hedwigia, Dresden 11:146.
1872. On dry leaves of Luzula maxima
(Reichard) DC., Juncaceae. Italy.

lycii G. Passerini in P. Brunaud, Actes de la Société
Linnéenne de Bordeaux 41:175. 1887. On
dead roots of Prunus spinosa L., Solanaceae.
Guadeloupe (West Indies).

lyciophila C.E. Fairman, Proceedings of the
Rochester Academy of Science, Rochester,
New York 6:126—127. 1922. On dead stems
of Lycium vulgare Dunal., Solanaceae.
U.S.A. (New York).

lycopodiicola C.H. Peck, Report. New York State
Museum of Natural History, Albany 38:105.
1885. =Phaeosphaeria lycopodiicola (C.H.
Peck) R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 67:1563. 1989. On dead

Vol. 34 Art. 3

March 199]

peduncles of Lycopodium clavatum L.,
Lycopodiaceae. Canada, U.S.A.

lycopodiicola C.H. Peck var. major H. Rehm in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
9:796. 1891 [as Leptosphaeria helvetica P.A.
Saccardo & C.L. Spegazzini in H. Rehm,
Ascomyceten, Fascicle 18, No. 884. Anno
1886]. On unknown substrate of Selaginella
helvetica (L.) Spring, Selaginellaceae.
Argentina.

lycopodina (J.P. Montagne) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:81. 1883.
=Sphaeria (Conferta) lycopodina J.P.
Montagne, Annales des Sciences Naturelles,
Paris, Botanique, Series 3, 12:313. 1849;
Sylloge Generum Specierumque Cryptoga-
marum Quas in Variis Operibus Descriptas
Iconobusque Illustratas, Nunc Ad Diagnosim
Reductas, Nonnullasque Novas Interjectas,
Ordine Systematico Disposiut, pp. 240-241.
1856. =Phaeosphaeria lycopodina (J.P.
Montagne) A. Hedjaroude, Sydowia. Annales
Mycologici, Horn, N.O. 22:78. 1968. On
dorsal bracts of Lycopodium annotinum L.,
Lycopodiaceae. Canada, Czechoslovakia,
Finland, Poland, Sweden, Switzerland, U.S.A.

lyndonvillae C.E. Fairman, Annales Mycologici,
Berlin 4:326—327. 1906. On pods of Robinia
pseudacacia L., Leguminosae. U.S.A.

lythri C.H. Peck, Bulletin of the Torrey Botanical
Club (and Torreya), New York 33:220-221.
1906. On dead stems of Lythrum alatum
Pursh, Lythraceae. U.S.A.

maclurae J.B. Ellis & B.M. Everhart, Proceedings of
the Academy of Natural Sciences of Philadel-
phia 1890:236—237. 1890. =Metasphaeria
maclurae (J.B. Ellis & B.M. Everhart) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
9:838. 1891. On leaves of Maclura auran-
tiaca Nutt., Moraceae. U.S.A.

macrochloae L. Lacoste, Revue de Mycologie 22,
Paris (Supplement Colonial No. 1):10—12.
1957. On leaves of Stipa tenacissima L.,
Gramineae. Algeria, Tunisia.

macrorostra A.M. Saccas, Etude de la Flore
Cryptogamique des Caféiers en Afrique
Centrale. Bulletin de Institut Frangais du Café
du Cacao et d’ Autres Plantes Stimulantes
(Bulletin IFCC No. 16), pp. 254-256. 1981.
Nom. inval. Art. 37.1. On dead branches of
Coffea robusta L. Linden (=Coffea canephora
Pierre ex Froehn.), Rubiaceae. Central
African Republic.

macrospora (L. Fuckel) F. v. Thiimen, Mycotheca
Universalis, Century 14, No. 1359. Anno
1879; Hedwigia, Dresden 21:83. 1882.
=Pleospora macrospora L. Fuckel, Symbolae

Nomenclator of Leptosphaeria

247

Mycologicae, p. 138. 1870. =Metasphaeria
macrospora (L. Fuckel) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:158.
1883. =Nodulosphaeria macrospora P.
Morthier, sub. Leptosphaeria macrospora F.
v. Thiimen. Nom. nud. On dry stems of
Senecio nemorensis L., Compositae. Austria.

macrosporidium E.B. Gareth Jones, Transactions of
the British Mycological Society, London
45:103. 1962. =Phaeosphaeria macrospori-
dium (E.B. Gareth Jones) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1532.
1989, On decayed stems of Spartina sp.,
Gramineae. Great Britain.

macrotheca (E. Rostrup) L. Holm, Svensk Botanisk
Tidskrift, Stockholm 46:38. 1952. =Metas-
phaeria macrotheca E. Rostrup in J. Lange,
Conspectus Florae Groenlandicae 3(1):561.
1889; Meddelelser om Grognland, Kjobenhavn
3:561. 1889. =Massaria macrotheca (E.
Rostrup) J. Lind, Skrifter om Svalbard og
Ishavet, Oslo 13:31. 1928. =Wettsteinina
macrotheca (E. Rostrup) E. Miiller, Sydowia.
Annales Mycologici, Horn, N.O. 12:203.
(1958) 1959. On dry leaves of Carex
hyperborea Drejer, Carex rigida Good.,
Cyperaceae. Greenland, Sweden.

maculans (J. Desmazieres) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:235. 1863.
=Sphaeria maculans J. Desmaziéres, Annales
des Sciences Naturelles, Paris, Botanique,
Series 3, 6:77—78. 1846. =Pleospora
maculans (J. Desmaziéres) L.R. Tulasne & C.
Tulasne, Selecta Fungorum Carpologica
2:274. 1863. On dried stems of Brassica
campestris L., Cruciferae. Belgium, France,
Germany, Great Britain, Italy, Portugal.

maculans (M.J. Sowerby) P.A. Karsten, Mycologia
Fennica 2:99. 1873. =Sphaeria maculans
M.J. Sowerby, Coloured Figures of English
Fungi or Mushrooms 3, Table 394, Figure 9.
1803. Nec Sphaeria maculans J.
Desmazieres. =Sphaerella maculans (M.J.
Sowerby) B. Auerswald in W. Gonnermann
and G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Sammtlicher Pilze Europa’s,
Hefte 5 und 6, Synopsis Pyrenomycetum
Europaeorum, p. 18. 1869. =Pleospora
sowerby L. Fuckel, Symbolae Mycologicae,
Erster Nachtrag, p. 301. 1871. =Lepto-
sphaeria sowerby (L. Fuckel) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:78.
1883. On leaves of Scirpus lacustris L.,
Scirpus sp., Cyperaceae. Finland, Great
Britain, Switzerland.

maculans (J. Desmaziéres) V. Cesati & G. de
Notaris forma denudata F. Fautrey in C.

Roumeguére, Fungi Selecti Gallici Exsiccati,
Century 71, No. 7037. Anno 1896; Revue
Mycologique, Toulouse 19:150. 1897. On dry
roots of Brassica oleifera Moench, Crucif-
erae. France?

maculans (M.J. Sowerby) P.A. Karsten var.
typhicola P.A. Karsten, Mycologia Fennica
Pars 2, Pyrenomycetes, p. 100. 1873.
(Erroneously cited as Leptosphaeria typhicola
P.A. Karsten in P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:79. 1883.]
=Phaeosphaeria typhicola (P.A.Karsten) A.
Hedjaroude, Sydowia. Annales Mycologici,
Horn, N.O. 22:86. (1968) 1969. =Massari-
osphaeria typhicola (P.A. Karsten) A.
Leuchtmann, Sydowia. Annales Mycologici,
Horn, N.O. 37:168. 1984. On rotting reeds of
Typha latifolia L., Typhaceae. Finland.

maderensis F. Petrak, Beiblatt zu den Botanischen
Jahrbiichern 62(No. 142), Heft 3:110. 1928.
On dry stems of Vinca major L. Apocynaceae.
Madeira Archipelago (Portugal).

magnoliae N.N. Woronichin, Vestnik Tiflisskogo
Botanicheskogo Sada, Tiflis (Moniteur du
Jardin Botanique de Tiflis) 35:4. 1914. On
living leaves of Magnolia grandiflora L.,
Magnoliaceae. U.S.S.R. (Caucasus).

magnusiana A.N. Berlese & P.A. Saccardo in A.N.
Berlese and P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
414. 1886. On dry leaves of Chamaerops
humilis L., Palmae. Italy.

maheui R.G. Werner, Bulletin de la Société des
Sciences Naturelles du Maroc, Rabat
19(1):50-51. 1939. On apothecia of Rinodina
collectica (Flk.) Arn., Lichenes (Physciaceae).
Morocco.

major (H. Rehm) M.L. Farr & R.M. Horner, Nova
Hedwigia. Zeitschrift fiir Kryptogra-
menkunde, Weinheim 15:248—249. 1968.
=Leptosphaeria helvetica P.A. Saccardo &
C.L. Spegazzini forma major H. Rehm,
Hedwigia, Dresden 26:93. 1887. On leaves of
Selaginella helvetica (L.) Spring, Selaginel-
laceae. Italy, U.S.A. (Arizona).

malojensis W. Kirschstein, Annales Mycologici,
Berlin 37:107—108. 1939. Ona dry branch of
Salix sp., Salicaceae. Germany.

malyi R. Picbauer, Glasnik Zemaljskog Museja u
Bosni 1 Hercegovini Sarajevo 45:66. 1933.
On stems of Pedicularis brachyodonta
Schlosser & Vuk. var. fallacis (Beck) Hayek,
Scrophulariaceae. Yugoslavia.

mamillana (E.M. Fries:E.M. Fries) M.C. Cooke,
Grevillea, London 17:91. 1889. =Sphaeria
mamillana E.M. Fries:E.M. Fries, Kungliga
Svenska Vetenskapsakademiens Handlingar,
Uppsala and Stockholm 1818:103. 1818;

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Systema Mycologicum Sistens Fungorum
2:487. 1823. =Clypeosphaeria mamillana
(E.M. Fries:E.M. Fries) J.B. Lambotte, Flore
Mycologique de la Belgique, Comprenant la
description des espéces trouvées jusqu’a ce
Jour. 2:247. 1880. On branches of Cornus
sp., Cornaceae. Belgium.

mamillula M. Anzi, Atti della Societa Italiana di
Scienze Naturali e del Museo Civile di Storia
Naturale, Milano 11:180. 1868. =Epicymma-
tia mammillula (M. Anzi) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 1:571.
1882. On upper surface of thallus of
Peltigera canina (L.) Willd., Lichenes
(Peltigeraceae). Italy.

mandshurica M. Miura, Flora of Manchuria and East
Mongolia, Part III, Cryptogams, Fungi
(Contributions of the Southern Manchuria
Railway Company 27):175. 1928. On leaves
of Pyrus malus L. [as Malus domestica
Borkh.], Rosaceae. China.

marantae H. Sydow & P. Sydow, Annales Mycol-
ogici, Berlin 18:100. 1920. On dying leaves
of Maranta arundinacea L., Marantaceae.
Philippines.

marcyensis (C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:80. 1883.
=Sphaeria mareyensis C.H. Peck, Report.
New York State Museum of Natural History,
Albany 31:51. 1879 [as marciensis}.
=Phaeosphaeria marcyensis (C.H. Peck) L.
Holm & K. Holm, Karstenia. Suomen Sieni-
Seura, Helsinki 21:68. 1981. On leaves of
Lycopodium annotinum L., Lycopodium
selago L., Lycopodiaceae. Canada, U.S.A.

marginalis G. Passerini, Atti della Societa Crittoga-
mologica Italiana, Milano, Series 2, 2:33.
1879. =Leptosphaeria passerinii P.A.
Saccardo. Non P.A. Saccardo, 1877. On
wilted leaves of Achillea ptarmica L.,
Compositae. Italy.

(Leptosphaeriella) marginalis P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 1:244.
1878. On leaves of Albizia julibrissin Durz.,
Leguminosae. Italy.

marginata (K.F.W. Wallroth) G. Niessl v. Mayen-
dorf, Verhandlungen des Naturforschenden
Vereins in Briinn 10:171—172. (1871) 1872.
=?Sphaeria marginata K.F.W. Wallroth,
Flora Cryptogamica Germaniae 2:770. 1833.
On fallen leaves of Pyrola secunda L.,
Pyrolaceae. Austria, Czechoslovakia, Italy.

marina E. Rostrup, Botanisk Tidsskrift, Kjobenhavn
17:234. 1890. Nom. illegit. Art. 64.1.
=Leptosphaeria danica A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum ad

March 1991

usum Sylloges Saccardianae Adcommodatae
1:87. 1892. =Leptosphaeria chondri L.K.
Rosenvinge, Botanisk Tidsskrift, Kjobenhavn
27:XXXIII-XXXV. 1906. =Didymosphaeria
marina (E. Rustrup) J. Lind, Danish Fungi as
Represented in the Herbarium of E. Rostrup,
Nordisk Forlag, Copenhagen, p. 214. 1913.
=Didymosphaeria danica (A.N. Berlese) I.M.
Wilson & J.M. Knoyle, Transactions of the
British Mycological Society, London 44:57.
1961. =Lautitia danica (A.N. Berlese) S.
Schatz, Canadian Journal of Botany, Ottawa
63:31. 1984. Parasitic on Chondrus crispus J.
Stackhouse, Algae (Gigartinaceae). Denmark.

marina J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 1:43. 1885.
=Heptameria marina (J.B. Ellis & B.M.
Everhart) M.C. Cooke, Grevillea, London
18:32. 1889. =Metasphaeria marina (J.B.
Ellis & B.M. Everhart) A.N. Berlese, [cones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:140. 1894. =Wettsteinina marina (J.B. Ellis
& B.M. Everhart) R.A Shoemaker, Canadian
Journal of Botany, Ottawa 67:1596. 1989. On
dead culms of Spartina sp., Gramineae.
U.S.A.

maritima L. Hollés, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 5:46.
1907. Nom. illegit. Art. 64.1. On dry stems
and leaves of Triglochin maritima L.,
Juncaginaceae. Hungary.

maritima (M.C. Cooke & C.B. Plowright) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:73. 1883. =Sphaeria maritima M.C. Cooke
& C.B. Plowright in M.C. Cooke, Grevillea,
London 5:120. 1877. On culms of Juncus
maritimus Lam., Juncaceae. Great Britain.

marram (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:60. 1883.
=Sphaeria marram M.C. Cooke, Grevillea,
London 5:120. 1877. =Heptameria marram
(M.C. Cooke) M.C. Cooke, Grevillea, London
18:31. 1889. =Phaeosphaeria marram (M.C.
Cooke) O. Eriksson, Arkiv for Botanik,
Uppsala, Stockholm, Series 2, 6:425. 1967.
On stems of Ammophila sp., Gramineae.
Canada, Great Britain.

martagoni M. Losa, Anales del Jardin Botanico de
Madrid 8:309. (1947) 1948. On dead stems of
Lilium martagon L., Liliaceae. Spain.

(Melanomma) martinianum (W.L. Lindsay) M.C.
Cooke, Grevillea, London 17:91. 1889.
=Sphaeria maritiniana W.L. Lindsay,
Transactions of the Royal Society of Ed-
inburgh 24:427-428. 1867. =Melanomma
martinianum (W.L. Lindsay) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque

Nomenclator of Leptosphaeria

249

Cognitorum Digessit P.A. Saccardo 2(Ad-
denda Ad Volumen Secundum):LIX. 1883.
On cortex of not yet dead trunks of Sophora
tetraptera J, Mill., Leguminosae. New
Zealand.

massariella P.A. Saccardo & C.L. Spegazzini in
P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:393—394. 1878. On dead branches
of Ulmus campestris auct., Ulmaceae. Italy.

massariella P.A. Saccardo & C.L. Spegazzini var.
brasiliensis H. Rehm, Annales Mycologici,
Berlin 5:528. 1907. On branches of unknown
host, unknown family. Brazil.

massariella P.A. Saccardo & C.L. Spegazzini var.
disticha D, Saccardo, Malpighia. Rassegna
Mensuale di Botanica. Messina, Genova
12:208. 1898. On dead stems of Morus alba
L., Moraceae. Italy.

massarioides P.A. Saccardo & C.L. Spegazzini,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Ilustrans 1:394.
1878. =Heptameria (Leptosphaeria)
massarioides (P.A. Saccardo & C.L. Spegaz-
zini) M.C. Cooke, Grevillea, London 18:31.
1889. =Massariosphaeria massarioides (P.A.
Saccardo & C.L. Spegazzini) S.M. Huhndorf,
J.L. Crane, & C.A. Shearer, Mycotaxon. An
International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens, Ithaca,
New York 37:204-205. 1990. On decaying
stems of Heracleum sphondylium L.,
Umbelliferae. Italy.

matisiae P.C, Hennings, Hedwigia, Dresden
48:107—108. 1908 [as matiaiae]. On fronds of
Matisia paraensis Hub., Matoniaceae
(Filicopsida). Brazil.

matritensis R. Gonzalez Fragosa, Boletin de la R.
Sociedad Espanola de Historia Natural,
Madrid 18:368—369. 1919. On culms, leaves,
and sheaths of Stipa pennata L., Gramineae.
Spain.

maydis G.L. Stout, Mycologia, Lancaster, Pennsyl-
vania 22:275. 1930. On leaves of Zea mays
L., Gramineae. U.S.A. (Illinois).

media §.A. Gucevicz, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 10:183. 1955. On
dead stems and sheaths of Briza media L.,
Gramineae. U.S.S.R.

medicaginicola M.A. Karimov, Notulae Systemati-
cae e Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komarov Academiae Scientia-
rum U.R.S.S., Petropolis 11:122. 1956. On
decaying stems of Medicago sativa L.,
Leguminosae. U.S.S.R.

medicaginis P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova

250

2(2):153. 1873. =Leptosphaeria medicag-
inum P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:534. 1882. Non P.A. Saccardo,
1883. On stems of Medicago sp.; not stated
but presumed, Leguminosae. Austria,
Germany, Italy.

medicaginis (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:19. 1883. Nom.
illegit. Art. 64.1. =Pleospora medicaginis L.
Fuckel, Symbolae Mycologicae, p. 134. 1870.
On dried stems of Medicago sativa L.,
Leguminosae. Germany.

medicaginum P.A. Saccardo, Michelia Commentar-
ium Mycologicum Fungos in Primis Italicos
Illustrans 2:534. 1881. =Leptosphaeria
medicaginis P.A. Saccardo, Atti dell’
Accademia Scientifica Veneto-Trentino-
Istriana, Padova 2(2):153. 1873. On stems of
Medicago sp., Leguminosae. Austria,
Germany, Italy.

megalospora B. Auerswald & G. Niessl v. Mayen-
dorf, Verhandlungen des Naturforschenden
Vereins in Briinn 10:180. (1871) 1872.
=Nodulosphaeria megalospora (B. Auerswald
& G. Niessl v. Mayendorf) L. Holm, Svensk
Botanisk Tidskrift, Stockholm 55:74. 1961.
=Ophiobolus megalosporus (B. Auerswald &
G. Niessl v. Mayendorf) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 54:2384.
1976. On dead stems of Sambucus ebulus L.,
Caprifoliaceae. Czechoslovakia.

melanommoides F. Fautrey in C. Roumeguére,
Revue Mycologique, Toulouse 13:128—129.
1891; Fungi Selecti Gallici Exsiccati, Century
58, No. 5752. Anno 1891. On old rotting
stems of Petroselinum hortense auct.,
Umbelliferae. France.

melanommoides C.L. Spegazzini, Anales del Museo
Nacional de Historia Natural de Buenos Aires
6:282. 1899; Fungi Argentini Novi vel Critici,
No. 628. Anno 1899. Nom. illegit. Art. 64.1.
=Leptosphaeria subsuperficialis P.A.
Saccardo & P. Sydow. On fallen decaying
culms of Panicum grumosum Nees, Gramin-
eae. Chile.

melanommoides A.N. Berlese, cones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:54.
1890. =Leptosphaeria revocans P.A.
Saccardo. Nec F. Fautrey, 1891. On branches
of Buxus sempervirens L., Buxaceae.
Belgium.

melicae F. Bubak, Annales des K.K. Naturhistori-
schen Hofmuseums Wien 28:198. 1914.
Nom. illegit. Art. 64.1. =Massariosphaeria
melicae (F. Bubak) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67: 1582.
1989. On dead stems of Melica inaequiglumis
Boiss., Gramineae. Turkey.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

melicae G. Passerini, Atti della Societa Crittogamol-
ogica Italiana, Milano, Series 2, 2:44 45.
1879. On wilted leaves of Melica uniflora
Retz., Gramineae. Italy.

meliloti L. Hollés, Matematikai és Természettu-
domanyi Kozlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:33. 1926. On dry
stems of Melilotus alba Medicus, Legumino-
sae. Hungary.

mellispora M.L. Farr & H.T. Homer, Nova Hed-
wigia. Zeitschrift fiir Kryptogramenkunde,
Weinheim 15:251—252. 1968. On leaves of
Selaginella underwoodii Hieron., Selaginel-
laceae. U.S.A. (Arizona).

mendozana F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 1(4-6):294-295. 1947. On dried
leaves of Trisetum longiglume Hackel,
Gramineae. Argentina.

menthae F. Fautrey & J.B. Lambotte, Revue
Mycologique, Toulouse 17:169. 1895. On dry
stems of Mentha sylvestris L. [=Mentha
longifolia (L.) Huds.], Labiatae. France.

meridionalis D. Saccardo, Mycotheca Italica,
Century 4, No. 302. Anno 1899. Nom. nud.
Art. 32.1. Ex D. Saccardo in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo
14:563-564. 1899. On branches of Tamarix
gallica L., Tamaricaceae. Italy.

mertensiae (J.B. Ellis) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:34. 1883.
=Sphaeria (Leptosphaeria) mertensiae J.B.
Ellis, Bulletin of the Torrey Botanical Club
(and Torreya), New York 8:90-91. 1881. On
dead leaves of Mertensia sp., Boraginaceae.
U.S.A.

mesoedema (M.J. Berkeley & M.A. Curtis) J.B. Ellis
& B.M. Everhart, The North American
Pyrenomycetes. A Contribution to Mycologic
Botany, p. 365. 1892. =Sphaeria mesoedema
M.J. Berkeley & M.A. Curtis in M.J.
Berkeley, Grevillea, London 4:151. 1876.
=Heptameria mesoedema (M.J. Berkeley &
M.A. Curtis) P.A. Saccardo, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:89. 1883. On stems
of Cirsium muticum Michx., Eupatorium
coronopifolium Willd., Compositae. U.S.A.
(South Carolina).

metasequoiae S.A. Gucevicz, Trudy Gosudarstven-
nogo Nikitskogo Botanicheskogo Sada, Yalta
32:120. 1960. On dry axillary shoots and
short shoots of Metasequoia glyptostroboides
Hu & Cheng, Taxodiaceae. U.S.S.R.
(Crimea).

michotii (G.D. Westendorp) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:58-59. 1883.
=Sphaeria michotii G.D. Westendorp,

March 1991

Bulletins de L’ Académie Royale des
Sciences, des Lettres et des Beaux-Arts de
Belgique, Brussels, Series 2, 7:87. 1859.
=Sphaerella michotii (G.D. Westendorp) B.
Auerswald in W. Gonnermann and G.L.
Rabenhorst, Mycologia Europaea, Ab-
bildungen Sammtlicher Pilze Europa’s, Hefte
5 und 6, Synopsis Pyrenomycetum Euro-
paeorum, p. 18. 1869. =Paraphaeosphaeria
michotii (G.D. Westendorp) O. Eriksson,
Arkiv for Botanik, Uppsala, Stockholm,
Series 2, 6:406. 1967. =Scleropleella michotii
(G.D. Westendorp) F. v. Héhnel, Annales
Mycologici, Berlin 18:76. 1920. On dead
culms of Andropogon sp., Juncus sp., Scirpus
sp., Gramineae, Juncaceae, Cyperaceae.
Belgium, France, Germany, Great Britain,
Italy, Portugal.

micropogon P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:320. 1875. On
canes of Scirpus holoschoenus L., Cyper-
aceae. Italy.

microscopica P.A. Karsten, Ofversigt af K.
Vetenskapsakademiens Forhandlingar,
Stockholm 29(2):102. 1872. =Scleropleella
microscopica (P.A. Karsten) A. Munk, Dansk
Botanisk Arkiv, Kjgbenhavn 15(2):107. 1953.
=Phaeosphaeria microscopica(P.A.Karsten)
O. Eriksson, Arkiv for Botanik, Uppsala,
Stockholm, Series 2, 6:426. 1967.
=Phaeosphaeria microscopica(P.A. Karsten)
O. Eriksson var. microscopica, Arkiv for
Botanik, Uppsala, Stockholm, Series 2, 6:427.
1967. On rotting leaves of Alopecurus ovatus
Knapp, Poa alpina L., Poa colpodea T. Fries,
Trisetum subspicatum (L.) Beauy., Gramin-
eae. Canada, Denmark, Finland, Germany,
Italy, Sweden.

microscopica P.A. Karsten forma brachypodii E.
Niel in C. Roumeguere, Fungi Selecti Gallici
Exsiccati, Century 65, No. 6437. Anno 1894;
Revue Mycologique, Toulouse 16:8. 1894.
On culms of Brachypodium sylvaticum
(Hudson) Beauv., Gramineae. France.

microscopica P.A. Karsten forma glyceriae F.
Fautrey in C. Roumeguére, Fungi Selecti
Gallici Exsiccati, Century 57, No. 5651. Anno
1891; Revue Mycologique, Toulouse 13:78.
1891. On seeds and leaves of Glyceria
aquatica (L.) Wahlenb., Gramineae. France.

microscopica P.A. Karsten subsp. calami P.A.
Karsten, Hedwigia, Dresden 22:178. 1883;
Acta Societatis pro Fauna et Flora Fennica,
Helsingforsiae 2(6):52—53. (1884) 1885. On
rotten leaves of Acorus calamus L., Araceae.
Finland.

microscopica P.A. Karsten var. alpina T. Ferraris,
Malpighia. Rassegna Mensuale di Botanica.
Messina, Genova 16:453. 1902. On leaves of
Luzula lutea (All.) DC., Juncaceae. Italy.

Nomenclator of Leptosphaeria 251

microscopica P.A. Karsten var. caricis-vulpinae R.
Staritz in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 17:729. 1905. On unknown
substrate of Carex vulpina L., Cyperaceae.
Germany.

microscopica P.A. Karsten var. flavo-brunnea J.
Feltgen [falsely so cited in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 17:729.
1905.] =Leptosphaeria culmorum B.
Auerswald var. flavo-brunnea J. Feltgen.

microspora J.B. Ellis & B.M. Everhart, Bulletin of
the Torrey Botanical Club (and Torreya), New
York 24:461. 1897. On dead stems of
Lespedeza capitata Michx., Leguminosae.
Canada.

microthyrioides J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(2):154. 1901. On dry stems of /ris
pseudacorus L., Iridaceae. Luxembourg.

mikaniae A.C. Batista & G.E.P. Peres in A.C.
Batista, R.G.S. Falcao, and G.E.P. Peres, Atas
do Instituto de Micologia, Recife 4:45—46.
1967. On leaves of Mikania sp., Compositae.
Brazil.

millefolii (L. Fuckel) G. Niessl v. Mayendorf in G.L.
Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio Edita Nova, Series 2, Century 23, No.
2239. Anno 1876. =Pleospora millefolii L.
Fuckel, Symbolae Mycologicae Dritter
Nachtrag, p. 20. 1875. On dry stems of
Achillea millefolium auct., Compositae.
Czechoslovakia.

minima (J.E. Duby) ex P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 2:320. 1881.
=Sphaeria minima J.E. Duby in C.
Roumeguére, Fungi Selecti Gallici Exsiccati,
Century 7, No. 694. Anno 1880. Nom. nud.
=M ycotodea minima (J.E. Duby ex P.A.
Saccardo) W. Kirschstein in O.C. Schmidt,
Kryptogamenflora der Mark Brandenburg und
Angrenzender Gebiete herausgegeben von
dem Botanishen Verein der Provinz Bran-
denburg, Leipzig 7(3):432. 1938. On leaves
of Potentilla argentea L., Rosaceae. France.

minoensis K. Hara, Botanical Magazine, Tokyo
27(317):250. 1913. On unknown substrate of
Phyllostachys bambusoides Sieb. & Zucc. var.
bambusoides, Gramineae. Japan.

mirabilis G. Niessl v. Mayendorf, Hedwigia,
Dresden 20:97. 1881. =Pleospora mirabilis
(G. Niessl v. Mayendorf) F. Petrak, Annales
Mycologici, Berlin 25:207. 1927. =Wettstein-
ina mirabilis (G. Niessl vy. Mayendorf) F. v.
Hohnel, Sitzungsberichte der Akademie der
Wissenschaften in Wien, Mathematisch-
naturwissenschaftliche Klasse, Abt. 1,

116:635. 1907. On dry stems of Chondrilla
juncea L., Algae (Chondriellaceae). Un-
known country.

mirandae A. Caballero, Boletin de la R. Sociedad
Espanola de Historia Natural, Madrid
27:61—62. 1927. On living thallus of
Mesogloia leveillei (J. Ag.) Mengh., Algae
(Chordariaceae, Phaeophyceae). Italy.

miyakeana P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 24:999, 1928. =Phaeosphaeria
eriobotryae |. Miyake, Botanical Magazine,
Tokyo 27:41-42. 1913. =Trematosphaerella
erlobotryae (I. Miyake) F.L. Tai, Sylloge
Fungorum Sinicorum, Science Press.
Academia Sinica, Peking, p. 330. 1979. On
leaves of Eriobotrya japonica (Thunb.)
Lindley, Rosaceae. China.

modesta (J. Desmazieéres) B. Auerswald in G.L.
Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio Edita Nova, Edition 3 (Editio Nova,
Series Secunda), Century 10, No. 948. Anno
1866. =Sphaeria (Caulicola) modesta J.
Desmaziéres, Annales des Sciences
Naturelles, Paris, Botanique, Series 3, No. 8,
p. 173. 1847. =Nodulosphaeria modesta (J.
Desmazieres) A. Munk, Dansk Botanisk
Arkiv, Kjobenhavn 15(2):136. 1953.
=Nodulosphaeria modesta (J. Desmaziéres)
A. Munk ex L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):80. 1957. On dry
stems of Scabiosa columbaria L., Dip-
sacaceae. Finland, France.

modesta (J. Desmazieres) P.A. Karsten, Fungi
Fenniae Exsiccati, Century 9, No. 889. 1869;
Mycologia Fennica 2 (Pyrenomycetes): 106.
1873. =Sphaeria modesta J. Desmazieéres,
Annales des Sciences Naturelles, Paris,
Botanique, Series 3, No. 8, p. 173. 1847.
Nom. illegit. Art. 64.1.

modesta (J. Desmaziéres) P.A. Karsten forma dauci
F. Fautrey in C. Roumeguére, Fungi Selecti
Gallici Exsiccati, Century 66, No. 6553. Anno
1894; Revue Mycologique, Toulouse 16:112.
1894. On flowering apex of Daucus sp.,
Umbelliferae. France.

modesta (J. Desmaziéres) P.A. Karsten forma
digitalis-luteae F. Fautrey in C. Roumegueére,
Fungi Selecti Gallici Exsiccati, Century 56,
No. 5538. Anno 1891; Revue Mycologique,
Toulouse 13:8. 1891. On dry stems of
Digitalis lutea L., Scrophulariaceae. France.

modesta (J. Desmaziéres) P.A. Karsten forma
Jacobaeae F, Fautrey in C. Roumeguére,
Revue Mycologique, Toulouse 14:172. 1892;
Fungi Selecti Gallici Exsiccati, Century 62,
No. 6141. Anno 1892. On stems of Senecio
Jacobaea L.., Compositae. France.

modesta (J. Desmaziéres) P.A. Karsten forma /appae
E. Niel in C. Roumeguére, Fungi Selecti

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Gallici Exsiccati, Century 65, No. 6438. Anno
1894; Revue Mycologique, Toulouse 16:8.
1894. On dead stems of Lappa sp. (=Arctium
sp.), Compositae. France?

modesta (J. Desmaziéres) P.A. Karsten forma
succisae F. Fautrey in C. Roumeguére, Fungi
Selecti Gallici Exsiccati, Century 56, No.
5539. Anno 1891; Revue Mycologique,
Toulouse 13:8. 1891. On dry stems of
Succisa sp., Dipsacaceae. France.

modesta (J. Desmazieres) P.A. Karsten forma
sylvestris F. Fautrey in C. Roumeguére, Fungi
Selecti Gallici Exsiccati, Century 61, No.
6034. Anno 1892; Revue Mycologique,
Toulouse 14:107. 1892. On stems of
Dipsacus sylvestris Hudson, Dipsacaceae.
France.

modesta (J. Desmaziéres) P.A. Karsten var. cibostii
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:40. 1883. On stems of Cynanchum sp., and
on an unknown species of Umbelliferae,
Asclepiadaceae, Umbelliferae. Italy.

modesta (J. Desmazieres) B. Auerswald var.
rubellula J. Desmaziéres in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:40.
1883. =Leptosphaeria rubellula (J. Des-
mazieres) F. v. HOhnel. On stems of
Barkhousia taraxacifolia (Thuill.) DC.,
Compositae. France.

modestula F.E. Clements & E.S. Clements, Crypto-
gamae Formationum Coloradensium, Century
3, No. 237. Anno 1907. Nom. nud. Art. 36.1.
On dead stems of Geranium richardsonii
Fisch. & Trautv., Geraniaceae. U.S.A.

molleriana J. Verissimo d’ Almeida & M. de Souza
da Camara, Boletim da Sociedade Broteriana,
Coimbra 24:165. 1908/1909. On leaves of
Cocos romanzoffiana Cham. [=Arecastrum
romanzoffianum (Cham.) Beccari}, Palmae.
Portugal.

molleriana G. Niessl v. Mayendorf, Instituto.
Revista Scientifica e Litteraria, Coimbra
31:90. 1883. =Metasphaeria molleriana (G.
Niessl v. Mayendorf) A.N. Berlese & P.
Voglino, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
Additamenta Ad Volumina I-IV, p. 157.
1886. [Perhaps Sphaerella molleriana F. v.
Thiimen?, fide G. Niess] v. Mayendorf.] On
dead leaves of Eucalyptus globulus Labill.,
Myrtaceae. Portugal.

molluginis G. Passerini, Atti dell” Accademia
Nazionale dei Lincei. Memoire, Rome, Series
4, 6:459. (1889) 1890. On dry stems of
Galium mollugo L., Rubiaceae. Italy.

molybdina J.P. Montagne & J.B. Mougeot ex P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Ilustrans

March 1991

2:319. 1881. =Sphaeria molybdina J.P.
Montagne in C. Roumeguére, Fungi Selecti
Gallici Exsiccati, Century 6, No. 580. Anno
1879. Nom. nud. Art. 32.1. =Sphaeria
molybdina J.P. Montagne in J.B. Mougeot,
J.A. Mougeot, C.G. Nestler, and W.P.
Schimper, Stirpes Cryptogamae Vogeso-
Rhenanae Quas in Rheni Superioris Inferioris-
que, Nec Non Vogesorum Praefecturis,
Collegerunt, J.B. Mougeot et C.G. Nestler,
Fascicle 15, No. 1451. Anno 1860. [P.A.
Saccardo’s validated description cites only
Fungi Gallici Exsiccati No. 580.] On wilted
stems of Vincetoxicum officinale Moench,
Asclepiadaceae. France.

monilispora (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:79. 1883.
=Sphaeria monilispora L. Fuckel, Fungi
Rhenani Exsiccati A Leopoldo Fuckel
Collecti, Supplement Fascicle 3, No. 1777.
Anno 1866. =Pleospora monilispora L.
Fuckel, Symbolae Mycologicae, p. 138. 1870.
On dry stalks and sheaths of Juncus lam-
pocarpus Ehrh. ex Hoffm., Juncaceae.
Austria.

monilispora P.A. Saccardo forma triglochinis J.
Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):208.
1903. On stalks of Triglochin palustris L.,
Juncaginaceae. Luxembourg.

monotis H. Rehm, Ascomyceten, Fascicle 18, No.
887. Anno 1886; Hedwigia, Dresden 26:94.
1887. [Reprint is page 14 at FH.] On dead
tufts of Saxifraga sp., Saxifragaceae. Italy.

montana G.B. Traverso, Manipolo di Funghi Della
Valle Pellina, Aosta (Tipografia cattolica), p.
20. 1912; Bulletin de la Société de la Flore
Valdotaine Aosta No. 8:34. 1912. Nom. nud.
Art. 32.1c. On dry stems of Salvia glutinosa
L., Labiatae. Italy.

monticola J.B. Ellis & B.M. Everhart, Bulletin of the
Torrey Botanical Club (and Torreya), New
York 24:461-462. 1897. On dead leaves and
petioles of Trifolium kingii S. Wats., Legumi-
nosae. U.S.A.

montis-bardi F. Fautrey & J.B. Lambotte, Revue
Mycologique, Toulouse 18:69. 1896. On
stems of Seseli montanum L., Umbelliferae.
France.

morierae F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 16(1-6):338. (1962) 1963. On dry
stems of Moriera sp., Cruciferae. Afghani-
stan. ‘

morthieriana P.A. Saccardo, Atti del Istituto Veneto
di Scienze, Lettere ed Arti, Venezia, Series 6,
2:457. (1883) 1884. On rotten leaves of
Succisa sp., Dipsacaceae. Switzerland.

mosana V. Mouton, Bulletin. Société R. de Bo-
tanique de Belgique, Bruxelles 39:45. 1900.
=Massariosphaeria mosana (V. Mouton) A.

Nomenclator of Leptosphaeria

253

Leuchtmann, Sydowia. Annales Mycologici,
Horn, N.O. 37:170. 1984. On culms of
Phragmites sp., Gramineae. Belgium, Great
Britain.

moutan W. Siemaszko, Acta Societatis Botanicorum
Poloniae, Warszawa 1:22. 1923. On leaves of
Paeonia moutan Sims. (=Paeonia suffruticosa
Haw.), Paeoniaceae. U.S.S.R.

moutoniana P.A. Saccardo & P. Sydow in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
16:517. 1902. =Leptosphaeria punctiformis
V. Mouton. On grass culms of unknown host,
Gramineae. Belgium.

mucosa V. Mouton, Bulletin. Société R. de Bo-
tanique de Belgique, Bruxelles 39:43. 1900.
On leaves of Typha sp., Typhaceae. Belgium.

muehlenbeckiae B.K. Vaidehi, Mycopathologia et
Mycologia Applicata Den Haag 38:135-136.
1969. On living phylloclades of Muehlen-
beckia platyclados Meissn., Polygonaceae.
India.

muehlenbergiae H. Rehm, Annales Mycologici,
Berlin 13:5. 1915. On branches of Muehlen-
bergia racemosa (Michx.) Britton, Stern, &
Pogg., Gramineae. U.S.A.

muelleri (J.E. Duby) B. Auerswald in W. Gonner-
mann and G.L. Rabenhorst, Mycologia
Europaea, Abbildungen Sammtlicher Pilze
Europa’s, Hefte 5 und 6, Synopsis Pyrenomy-
cetum Europaeorum, Table 12, Figure 167.
1869. =Sphaeria muelleri J.E. Duby in G.L.
Rabenhorst, Klotzschii Herbarium Vivum
Mycologicum Sistens Fungorum Per Totam
Germaniam Cresentium Collectionem
Perfectam, Dresden, Edition 2, Century 7, No.
642. Anno 1858. On scaly bark of Acer sp.,
Aceraceae. France.

muhlenbergiae (J.B. Ellis) G. Niessl v. Mayendorf,
Hedwigia, Dresden 22:33. 1883. =Dothidea
muhlenbergiae J.B. Ellis, Bulletin of the
Torrey Botanical Club (and Torreya), New
York 8:74~75. 1881. =Phyllachora muhlen-
bergiae (J.B. Ellis) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:604. 1883. On culms
of Muhlenbergia sp., Gramineae. North
America.

muirensis R. Sprague, Mycologia, Lancaster,
Pennsylvania 47:249. 1955. On living leaves
of Poa annua L., Gramineae. U.S.A.
(Alaska).

multiseptata H.G. Winter, Hedwigia, Dresden
11:148. 1872. =Entodesmium multiseptum
(H.G. Winter) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):133. 1957. On
dry stems of Lathyrus sylvestris L., Legumi-
nosae. Germany.

multiseptata H.G. Winter forma alpina H. Rehm,
Hedwigia, Dresden 24:235. 1885. Nom.
illegit. Art. 64.1. SLeptosphaeria epilobii E.

254 Illinois Natural History Survey Bulletin

Miiller. On dry stems of Epilobium fleischeri
Hochst., Onagraceae. Italy.

mume K. Hara, Byéchu-gai Zasschi (Journal of Plant
Protection), Tokyo 5:884. 1918 [as mune in
original publication]. On dead branch of
Prunus mume Sieb. & Zucc., Rosaceae.
Japan.

muralis P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:64. 1880. On stems of Parietaria
officinalis L., Urticaceae. France.

muricata J.B. Ellis & B.M. Everhart, Proceedings of
the Academy of Natural Sciences of Philadel-
phia 1893:447-448. 1894. On leaves of
Andropogon muricatus Retz. [=Vetiveria
zizanioides (L.) Nash], Gramineae. U.S.A.

musae T. Lin & J. Yen, Bulletin de la Société
Mycologique de France, Paris 87:427-429.
1971. On living leaves of Musa cavendishii
Lamb. ex Paxton, Musaceae. Taiwan.

musarum P.A. Saccardo & A.N. Berlese, Revue
Mycologique, Toulouse 11:204. 1889. On
dead leaves of Musa sp., Musaceae. Sao
Tomé.

muscari L. Hollés, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 7:51.
1909. On dry stems of Muscari comosum (L.)
Miller, Liliaceae. Hungary.

musicola (C.L. Spegazzini) P.A. Saccardo & A.
Trotter in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 22:229-230. 1913. [Incorrectly
cited as a comb. nov. based on Leptosphae-
rella musicola C.L. Spegazzini.] Nom. inval.
Art. 43.1.

musigena T. Lin & J. Yen, Bulletin de la Société
Mycologique de France, Paris 87:429-431.
1971. On living leaves of Musa cavendishii
Lamb. ex Paxton, Musaceae. Taiwan.

myricae J. Dearness & H.D. House, Bulletin of the
New York State Museum, Albany 197:30-31.
1917. On dead twigs and branches of Myrica
gale L., Myricaceae. U.S.A. (New York).

myrti S.A. Gucevicz, Trudy Gosudarstvennogo
Nikitskogo Botanicheskogo Sada, Yalta
32:155-156. 1960. On dry branches of
Myrtus communis L., Myrtaceae. U.S.S.R.

myrticola S.A. Gucevicz, Trudy Gosudarstvennogo
Nikitskogo Botanicheskogo Sada, Yalta
32:156—157. 1960. On dry branches of
Myrtus communis L., Myrtaceae. U.S.S.R.

nanae R.A. Shoemaker, Canadian Journal of Botany,

Ottawa 62:2708. 1984. On stems of Achillea
nana L., Compositae. Switzerland.

nandinae K. Togashi & E. Tsukamoto, Annals of the
Phytopathological Society of Japan, Tokyo
17:121. 1953. On unknown substrate of
Nandina domestica Thunb., Nandinaceae.
Japan.

napelli E. Miller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):245. 1950. On dead stems

Vol. 34 Art. 3

of Aconitum napellus L., Ranunculaceae.
Switzerland.

napi (L. Fuckel) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:45. 1883. =Pleospora napai L.
Fuckel, Symbolae Mycologicae, p. 136. 1870.
On stems and pieces of Brassica napus L.,
Cruciferae. Austria.

nardi (E.M. Fries) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:236. 1863. =Sphaeria
duplex M.J. Sowerby:E.M. Fries var. nardi
E.M. Fries:E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:520. 1823. =Sphaeria
nardi (E.M. Fries) E.M. Fries, Summa
Vegetabilium Scandinaviae, Seu Enumeratio,
Systematica et Critica, Plantarum tum
Cotyledonearum, Quam Nemearum Inter
Mare Occidentale et Album, Inter Eidoram et
Nordkop, Hactenus Lectorum, una Cum
Singulae Distributione Geographica, p. 394.
1849. =Sphaeria nardi (E.M. Fries) G.L.
Rabenhorst, Hedwigia, Dresden 1:116. 1857.
=Pleospora nardi (E.M. Fries) L. Fuckel,
Symbolae Mycologicae, p. 137. 1870.
=Phaeosphaeria nardi (E.M. Fries) L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):124. 1957. On dry culms and leaves of
Nardus stricta L., Gramineae. Belgium,
France, Germany, Great Britain, Sweden,
Switzerland.

nardi (E.M. Fries) V. Cesati & G. de Notaris var.
dubiosa V. Mouton, Bulletin. Société R. de
Botanique de Belgique, Bruxelles 26:177.
1887. =Leptosphaeria dubiosa (V. Mouton)
C.A. Oudemans, Enumeratio Systematica
Fungorum 1:981. 1919. On leaves of Scirpus
sylvaticus L., Gramineae. Belgium.

narmari J. Walker & A.M. Smith, Transactions of
the British Mycological Society, London
58:459-460. 1972. =Phaeosphaeria narmari
(J. Walker & A.M. Smith) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1551.
1989. On stolons and sheaths of Cynodon
dactylon (L.) Pers., Cynodon transvaalensis
Burtt-Davy, Hordeum vulgare L., Oryza
Sativa L., Pennisetum clandestinum Hochst.
ex Chiov., Stenotaphrum secundatum (Walter)
O. Kuntze, Triticum aestivum L., Gramineae.
Australia (New South Wales).

nashi (K. Hara) P.A. Saccardo in A. Trotter, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 24:999. 1928.
=Leptosphaerella nashi K. Hara, Kajyu-
byogai-ron Nihon-kankitsu-Kai, Schiznoka
[Fruit Tree Diseases], pp. 142—145. 1916.
=Phaeospora nashi (K. Hara) K. Hara,
Jitsuyo-Sakumotsu-B yorigaku [Manual of
Plant Pathology], p. 270. 1925. On leaves of
Pyrus sinensis Hort., Rosaceae. Japan.

March 1991

nectrioides C.L. Spegazzini in P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Ilustrans 1:458.
1879. On decayed runners of Clematis vitalba
L., Ranunculaceae. Italy.

neglecta G. Niess] v. Mayendorf, Verhandlungen des
Naturforschenden Vereins in Briinn
10:175—176. (1871) 1872. =Metasphaeria
neglecta (G. Niessl v. Mayendorf) P.A. Sac-
cardo, Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:173.
1883. On dried grass leaves of unknown host,
Gramineae. Italy (near Bozen, Tirolia).

neomaritima R.V. Gessner & J. Kohlmeyer,
Canadian Journal of Botany, Ottawa 54:2032.
1976. =Sphaeria maritima M.C. Cooke &
C.B. Plowright in M.C. Cooke, Grevillea,
London 5:120. 1877. Non Sphaeria maritima
P.L. Crouan & H. Crouan, Florule Du
Finistére Contenant Les Descriptions De 360
Espeéces Nouvelles De Sporogames, De
Nombreuses Observations et une synonymie
des plantes Cellulaires et Vasculaires Qui
Croissent Spontanément Dans ce Départe-
ment, p. 27. 1867. =Leptosphaeria maritima
(M.C. Cooke & C.B. Plowright) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:73. 1883. Non Leptosphaeria maritima L.
Hollos. 1906. =Phaeosphaeria neomaritima
(R.V. Gessner & J. Kohlmeyer) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1572. 1989. On deteriorating leaf
sheaths and decaying culms of Juncus
maritimus Lam., Juncus roemerianus Scheele,
Spartina alterniflora Loisel., Spartina
townsendii H. Groves & J. Groves, Spartina
sp., Juncaceae, Gramineae. Canada, Great
Britain, U.S.A.

neottizans (W.A. Leighton) F.W. Zopf, Hedwigia,
Dresden 35:361. 1896. =Verrucaria neotti-
zans W.A. Leighton, The Lichen-Flora of
Great Britain, Ireland, and the Channel
Islands, p. 497. Third edition. 1879. Parasitic
on Baeomyces rufus (Huds.) Rebent.,
Lichenes (Baeomycetaceae). Great Britain.

nervisequa H.G. Winter, Boletim da Sociedade
Broteriana, Coimbra 2:42. (1883) 1884.
=Metasphaeria nervisequa (H.G. Winter)
A.N. Berlese & P. Voglino, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo Additamenta Ad
Volumina I-IV, p. 158. 1886. On leaves of
Smilax pseudo-china L., Liliaceae. Portugal.

nesodes (M.J. Berkeley & C.E. Broome) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:85. 1883. =Sphaeria (Depazea) nesodes
M.J. Berkeley & C.E. Broome, Journal of the
Linnean Society (Botany), London 14:129.
(1873) 1875. On leaves of Hydrocotyle
asiatica L., Hydrocotylaceae. Sri Lanka.

Nomenclator of Leptosphaeria

nN
mn
wn

nicolai F. Bubak, Sitzungsberichte der K. Bohmis-
chen Gesellschaft der Wissenschaften, Prag
Mathematisch-Naturwissenschaftliche Klasse
1903(12):10. 1904. On preceding year’s
stems of Salvia officinalis L., Labiatae.
Yugoslavia.

niessleana G.L. Rabenhorst, Fungi Europaei
Exiccati, Klotzschii Herbarii vivi Mycologici
Continuatio. II] Ausgabe (Editio nova, Series
Secunda), Century 13, No. 1252. Anno 1869.
Nom. nud. Art. 32.1. On dried stems and
leaves of Lathyrus latifolius L., Leguminosae.
Czechoslovakia.

niessleana G.L. Rabenhorst ex G. Niessl v. Mayen-
dorf, Verhandlungen des Naturforschenden
Vereins in Briinn 10:179. (1871) 1872. =En-
todesmium niessleanum (G.L. Rabenhorst ex
G. Niessl v. Mayendorf) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):133.
1957. On dried stems and leaves of Lathyrus
latifolius L., Leguminosae. Czechoslovakia.

niessleana G.L. Rabenhorst ex G. Niessl v. Mayen-
dorf forma viciae W.B. Grove, Journal of
Botany, British and Foreign, London 68:74.
1930. On dead stems of Vicia sativa L.,
Leguminosae. Great Britain.

niessleana G.L. Rabenhorst var. staritzii H. Rehm,
Hedwigia, Dresden 39:193. 1900; Ascomy-
ceten, Fascicle 27, No. 1335. Anno 1900. On
dry stems of Seseli sp., Umbelliferae.
Germany.

nigrans (M.R. Roberge) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:235. 1863. =Sphaeria
culmicola E.M. Fries:E.M. Fries var. nigrans
(M.R. Roberge) E.M. Fries, Summa Vegeta-
bilium Scandinaviae, Seu Enumeratio,
Systematica et Critica, Plantarum tum
Cotyledonearum, Quam Nemearum Inter
Mare Occidentale et Album, Inter Eidoram et
Nordkop, Hactenus Lectorum, una Cum
Singulae Distributione Geographica, p. 393.
1849. =Sphaeria (Caulicola) nigrans M.R.
Roberge in J. Desmazieres, Annales des
Sciences Naturelles, Paris, Botanique, Series
3, 6:79. 1846. =Phaeosphaeria nigrans (MLR.
Roberge) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):112. 1957. On
sheath of Dactylis glomerata L., Gramineae.
Austria, Canada, Finland, France, Germany,
Great Britain, India, Italy, Sweden, Switzer-
land, U.S.A.

nigrans (M.R. Roberge) V. Cesati & G. de Notaris
forma arundinis C. Roumeguére, Fungi
Selecti Gallici Exsiccati, Century 43, No.
4265. Anno 1887. On culms of Arundo
phragmites L. (=Phragmites communis Trin.),
Gramineae. France.

nigrella B. Auerswald in W. Gonnermann and G.L.
Rabenhorst, Mycologia Europaea, Ab-
bildungen Simmtlicher Pilze Europa’s, Hefte

256

5 und 6, Synopsis Pyrenomycetum Euro-
paeorum, Table 12, Figure 163. 1869. =Di-
aporthe nigrella (B. Auerswald) G. Niessl v.
Mayendorf, Verhandlungen des Naturforsch-
enden Vereins in Briinn 10:201—202. (1871)
1872. Nec Sphaeria nigrella E.M. Fries.
=Diaporthopsis nigrella (B. Auerswald) J.H.
Fabre, Annales des Sciences Naturelles, Paris,
Botanique, Series 6, 15:35—36. 1883. On
stems and branches of Eryngium campestre
L., Umbelliferae. Germany.

nigrella (G.L. Rabenhorst) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:21. 1883. =Cucur-
bitaria nigrella G.L. Rabenhorst, Hedwigia,
Dresden 12:140. 1873. On dead stems of
Brassica rapa L., Cruciferae. Germany.

nigricans P.A. Karsten, Mycologia Fennica Pars 2,
Pyrenomycetes, p. 107. 1873. On dry stems
of Chenopodium album L., Chenopodiaceae.
Finland.

nigricans P.A. Karsten var. grindeliae C.E. Fairman,
Mycologia, Lancaster, Pennsylvania 10:245.
1918. On unknown substrate of Grindelia?
sp., Compositae. U.S.A. (New Mexico).

nigrificans F. Bubak & A. Wroblewski in F. Bubak,
Hedwigia, Dresden 57:329. 1916.
Presumably Eudarluca caricis (E.M.
Fries:E.M. Fries) O. Eriksson, Botaniska
Notiser, Lund 119:49—50. 1966. Erroneously
cited as on the leaves and culms of Carex sp.
on rust pustules. France.

nigromaculata (H. Rehm) E. Miiller, Veroeffentli-
chungen des Geobotanischen Institutes,
Eidgenossiche Technische Hochschule Riibel
in Zurich 87:24. 1986. =Ophiobolus
(Plejobolus) nigromaculata H. Rehm,
Annales Mycologici, Berlin 10:393—394.
1912. On stems of Aconitum compactum
Rchb., Aconitum nappelus L., Aconitum
paniculatum Lam., Ranunculaceae. Germany
(Upper Bavaria).

nitschkei H. Rehm, Ascomyceten, Fascicle 1, No.
15. Anno 1870. Nom. nud. Art. 32.1; Flora,
Jena und Regensburg 55:510. 1872. On hard,
dry stems of Cacalia sp., Compositae.
Austria.

nitschkei H. Rehm forma adenostylidis H. Rehm,
Ascomyceten, Fascicle 11, No. 532. Anno
1879. Nom. nud. Art. 32.1. On fallen stems
of Adenostyles albifrons (L. fil.) Reichenb.,
Compositae. Germany.

nobilis P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:312. 1875. =Metasphaeria
nobilis (P.A. Saccardo) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:169.
1883. On leaves of Laurus nobilis L.,
Lauraceae. Italy.

nodorum E. Miiller, Phytopathologische Zeitschrift,
Berlin 19:409-410. 1952. =Phaeosphaeria

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

nodorum (E. Miller) A. Hedjaroude,
Sydowia. Annales Mycologici, Hom, N.O.
22:79. 1968. [Anamorph: Septoria nodorum
(M.J. Berkeley) M.J. Berkeley.) On dead
culms of Triticum vulgare Vill., Gramineae.
Canada, Kenya, Switzerland.

nodorum E. Miiller forma specialis hordei V.
Smedegord-Petersen, Friesia. Nordisk
Mykologisk Tidsskrift, Kobenhavn
10:262-263. 1974. On leaf lamina and
sheaths of Hordeum vulgare L., Gramineae.
Denmark.

norfolcia (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:73. 1883.
=Sphaeria norfolcia M.C. Cooke, Grevillea,
London 5:120-121. 1877. =Phaeosphaeria
norfolcia (M.C. Cooke) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:152. 1984. On stems of Eleocharis sp.,
Juncus sp., Gramineae, Juncaceae. France,
Great Britain.

norvegica E. Rostrup, Norske Ascomyceter i
Christiania Universitetets Botaniske Museum.
Christiania (Oslo) (I. Kommission Hos Jacob
Dybwad), p. 24. 1904. [Issued in Skrifter
Udg. af Videnskabsselskabet i Christiania.
Mathematisk-naturvidenskabelig klasse.
Christiania (Oslo) No. 274.] Non vidi. On
stems of Braya sp., Cruciferae. Poland.

(Clypeosphaeria) notarisii M.C. Cooke, Grevillea,
London 17:91. 1889. =Sphaeria clypeiformis
G. de Notaris, Memorie della Accademia delle
Scienze di Torino, Series 2, No. 7, p. 113.
1853; Micromycetes Italici Novi vel minus
Cogniti Decas 7, p. 113. 1845. Non Sphaeria
clypeiformis L.V. de Lacroix in G.L.
Rabenhorst. =Clypeosphaeria notarisii L.
Fuckel, Symbolae Mycologicae, p. 117. 1870.
On barky runners of Rubus fruticosus L.,
Rosaceae. Italy.

obesa (M.C. Durieu de Maisonneuve & J.P.
Montagne) P.A. Saccardo, Michelia Commen-
tarlum Mycologicum Fungos in Primis
Italicos Illustrans 1:38. 1877; Fungi Italici
autographice delineati (additis nonnullis extra-
italicis asterisco notatis), Patavii, Table 284.
1878. =Sphaeria obesa M.C. Durieu de
Maisonneuve & J.P. Montagne, Exploration
Scientifique de L’ Algérie Publiée Par Ordre
du Gouvernement. Series Naturelles.
Botanique I, pp. 526-527. 1868. =Hep-
tameria obesa (M.C. Durieu de Maisonneuve
& J.P. Montagne) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:88. 1883. On stems
of Scabiosa urceolata Desf., Cucurbitaceae.
Algeria.

obesula P.A. Saccardo in V. Mouton, Bulletin.
Société R. de Botanique de Belgique,
Bruxelles 26:176. 1887. On stems of Acer
pseudoplatanus L., Angelica sp., Spiraea

March 1991

ulmaria L. (=Filipendula ulmaria Maxim.),

Aceraceae, Umbelliferae, Rosaceae. Belgium.

obesula P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:318. 1881. On stems of most
herbaceous? plants. France.

obiones (P.L. Crouan & H.M. Crouan) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:24. 1883. =Pleospora obiones P.L. Crouan
& H.M. Crouan, Florule Du Finistere
Contenant Les Descriptions De 360 Espéces
Nouvelles De Sporogames, De Nombreuses
Observations et une synonymie des plantes
Cellulaires et Vasculaires Qui Croissent
Spontanément Dans ce Département, p. 22.
1867. =Passeriniella obiones (P.L. Crouan &
H.M. Crouan) K.D. Hyde & R. Mouzouras,
Transactions of the British Mycological
Society, London 91:183. 1988. On dead
stems of Obione sp. (=Atriplex), Chenopodi-
aceae. France.

obiones P.A. Saccardo var. evolutior W.B. Grove,
Journal of Botany, British and Foreign,
London 71:281—282. 1933. On dead stems of
Obione portulacoides (L.) Mog., Chenopodi-
aceae. Great Britain.

obtusispora C.L. Spegazzini, Anales de la Sociedad
Cientifica Argentina, Buenos Aires 12:179.
1881. =Paraphaeosphaeria obtusispora (C.L.
Spegazzini) O. Eriksson, Arkiv for Botanik,
Uppsala, Stockholm, Series 2, 6:406. 1967.
On decaying leaves of Yucca gloriosa L.,
Agavaceae. Argentina.

obtusispora C.L. Spegazzini forma agaves J.J.
Barthelet, Annales des Epiphyties, Paris, New
Series 8:118. 1942. On leaves of Agave ferox
C. Koch, Agave striata Zucc., Agavaceae.
France.

occidentalis J.B. Ellis & B.M. Everhart, Erythea. A
Journal of Botany, Berkeley 2:20. 1894.
=Phaeosphaeria occidentalis (J.B. Ellis &
B.M. Everhart) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1572. 1989. On
dead culms of Panicum crus-galli L.,
Gramineae. U.S.A.

occulta J, Lind, Danish Fungi as Represented in the
Herbarium of E. Rostrup, Nordisk Forlag,
Copenhagen, p. 218. 1913. =Phaeosphaeria
occulta (J. Lind) A. Leuchtmann, Sydowia.
Annales Mycologici, Horn, N.O. 37:128.
1984. On leaves of Carex hirta L., Cyper-
aceae. Denmark.

ocellata G. Niessl v. Mayendorf in J. Kunze, Fungi
Selecti Exsiccati, Century 4, No. 334. Anno
1880. Nom. nud. Art. 32.1. =Metasphaeria
ocellata (G. Niessl v. Mayendorf) ex P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:157. 1883. =Trichometasphaeria ocellata
(G. Niessl v. Mayendorf ex P.A. Saccardo) L.
Holm, Symbolae Botanicae Upsalienses

Nomenclator of Leptosphaeria 257

14(3):143. 1957. On dry stems of Hypericum
perforatum L., Guttiferae. Switzerland.

ocimicola S.R. Naphade, Sydowia. Annales
Mycologici, Horn, N.O. 24(1-6):242. (1970)
1971. On dead stems of Ocimum sanctum L.,
Labiatae. India.

octophragmia G.B. Traverso & R. Gonzalez Fragoso
in G.B. Traverso, Bollettino Della Societa
Botanica Italiana, Firenze 1915:23. 1915. On
dry stems of Aloysia citriodora Ort. ex Pers.,
Verbenaceae. Spain.

octophragmia G.B. Traverso & R. Gonzalez Fragoso
var. major W.B. Grove, Journal of Botany,
British and Foreign, London 71:282. 1933.
On very old thick dead stem of Lippia
(Aloysia) citriodora Kunth, Verbenaceae.
Great Britain.

octoseptata L.E. Wehmeyer, Lloydia: a quarterly
journal of biological science, Manasha
9:239-240. 1946. =Nodulosphaeria octosep-
tata (L.E. Wehmeyer) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):91.
1957. On stems of Senecio crassulus A.
Gray, Compositae. U.S.A. (Wyoming).

odora (M.C. Cooke & H.W. Harkness) A.N. Berlese
& P. Voglino, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
Additamenta Ad Volumina I-IV, p. 137.
1886. =Sphaeria (Leptosphaeria) odora M.C.
Cooke & W.H. Harkness, Grevillea, London
13:19. 1885. On branches of Umbellularia
sp., Lauraceae. U.S.A. (California).

ogilviensis (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:235. 1863.
=Sphaeria (Caulicolae) ogilviensis M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 9:379. 1852. On dead stems of Senecio
jacobaea L., Compositae. Canada, Germany,
Great Britain, Italy, Switzerland, U.S.A.

ogilviensis M.J. Berkeley & C.E. Broome forma
achilleae H. Rehm, Hedwigia, Dresden
39:(193). 1900. Nom. nud. Art. 32.1.
Ascomyceten, Fascicle 27, No. 1336. Anno
1900. Nom. nud. Art. 32.1. On stems of
Achillea millefolium auct., Compositae.
Germany.

ogilviensis (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris forma /epidii F. Fautrey in C.
Roumeguere, Fungi Selecti Gallici Exsiccati,
Century 62, No. 6138. Anno 1892; Revue
Mycologique, Toulouse 14:172. 1892. On
capsules of Lepidium campestre (L.) R. Br.,
Cruciferae. France.

ogilviensis (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris forma megalospora F.
Fautrey in C. Roumeguere, Fungi Selecti
Gallici Exsiccati, Century 61, No. 6035. Anno
1892; Revue Mycologique, Toulouse 14:107.

258

1892. On stems of Solidago virgaurea L.,
Compositae. France.

ogilviensis (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris forma myrrhis-odorata F.
Fautrey in C. Roumeguere, Fungi Selecti
Gallici Exsiccati, Century 56, No. 5540. Anno
1891; Revue Mycologique, Toulouse 13:8.
1891. On stems of Myrrhis odorata (L.)
Scop., Umbelliferae. France.

ogilviensis (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris var. pleurospermi O.A. Ade,
Allgemeine botanische Zeitschrift f. Systema-
tik, Floristik, Pflanzengeographie, etc.,
Karlsruhe 30/31:22(134). 1926. On dead
stems of Pleurospermum austriacum (L.)
Hoffm., Umbelliferae. Germany.

ogilviensis (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris var. senecionis-cordati C.B.
Massalongo, Malpighia. Rassegna Mensuale
di Botanica. Messina, Genova 8:194. 1894;
Nuova Contribuzione alla Micologia
Veronese Genova (Tipografia di Angelo
Ciminago), p. 37. 1894. On stems of Senecio
cordatus Koch, Compositae. Italy.

olericola (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:21. 1883. =Sphaeria olericola M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:150. 1876. On stems of
Brassica sp., Cruciferae. U.S.A.

oligospora (E.A. Vainio) P.A. Saccardo & D.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
17:730. 1905. =Xenosphaeria oligospora
E.A. Vainio, Meddelanden af Societas pro
Fauna et Flora Fennica, Helsingfors 10:203.
1883. On thalli of Solorina crocea (L.) Ach.,
Lichenes (Peltigeraceae). Finland.

oligotheca F. Petrak & H. Sydow, Annales Mycol-
ogici, Berlin 22:359. 1924. [Described from
the type material of Laestadia potentillae E.
Rostrup, Botanisk Tidsskrift, Kjobenhavn
25:300. 1903.] On leaves of Potentilla
maculata Pour. (=Potentilla aurea L.),
Rosaceae. Denmark, Iceland.

olivacea J.B. Ellis, Bulletin of the Torrey Botanical
Club (and Torreya), New York 10:53. 1883.
=Ophiobolus olivaceus (J.B. Ellis) A.N.
Berlese & P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
186. 1886. =Nodulosphaeria olivacea (J.B.
Ellis) L. Holm, Svensk Botanisk Tidskrift,
Stockholm 55:75. 1961. On dead herbaceous
stems of unknown host, unknown family.
U.S.A. (Utah).

olivaespora (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:85-86. 1883. =Sphaeria olivaespora M.J.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:148. 1876. On branches
of Cornus florida L., Cornaceae. U.S.A.

onagrae H. Rehm, Annales Mycologici. Berlin
11:398. 1913. On dead stems of Onagra
strigosa Rydb., Onagraceae. U.S.A.

onobrychidicola L. Hollés, Magyar Botanikai
Lapok, Budapest (Ungarische Botanische
Blatter) 28:50. (1929) 1930. On dried stems
of Onobrychis sativa Link, Leguminosae.
Hungary.

onobrychidis L. Hollos, Magyar Botanikai Lapok,
Budapest (Ungarische Botanische Blatter)
28:50. (1929) 1930. On dry stems of
Onobrychis sativa Link, Leguminosae.
Hungary.

ophioboloides P.A. Saccardo, Atti del Istituto
Veneto di Scienze, Lettere ed Arti, Venezia,
Series 6, 2:457. 1884. =Leptosphaeriopsis
ophioboloides (P.A. Saccardo) A.N. Berlese,
Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:89. 1892. =Ophiobolus
ophioboloides (P.A. Saccardo) L. Holm,
Svensk Botanisk Tidskrift, Stockholm 42:345.
1948. On stems of Tragopogon sp., Composi-
tae. Switzerland.

ophiopogonis (P.A. Saccardo) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:321.
1875. =Sphaerella ophiopogonis P.A.
Saccardo, Atti dell’ Accademia Scientifica
Veneto-Trentino-Istriana, Padova 2(2):
142-143. 1873. On fallen decaying leaves of
Ophiopogon japonicus Ker-Gawl., Liliaceae.
Italy.

ophiopogonis (P.A. Saccardo) P.A. Saccardo var.
graminum P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:68. 1883. On rotten leaves of
unknown host, Gramineae. France.

opizii T. Nitschke in J. Schroter, Kryptogamen-Flora
von Schlesien. Im Namen Der Schlesischen
Gesellschaft fiir vaterlandische Cultur
herausgegeben von Prof. Dr. Ferdinand Cohn,
Secretair der Botanischen Section, Breslau
3(2):358-359. 1894. On dead stems of
Solanum dulcamara L., Solanaceae. Poland.

opuntiae B.O. Dodge, Mycologia, Lancaster,
Pennsylvania 29:708. 1937. On stems of
Opuntia lindheimeri Engelm., Cactaceae.
U.S.A. (Texas).

orae-maris D.H. Linder, Farlowia. A Journal of
Cryptogamic Botany, Cambridge, Massachu-
setts 1:413. 1944. On driftwood of unknown
host. U.S.A. (California).

orchidearum P.A. Karsten, Hedwigia, Dresden
26:125. 1887. On dead stems of Gymnadenia
conopsea (L.) R. Br., Orchidaceae. Finland.

oreophila P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos

March 1991

Illustrans 1:120. 1878. On dead stems of
Tofieldia calyculata (L.) Wahlenb., Liliaceae.
Austria.

oreophiloides P.A. Saccardo & O. Penzig in P.A.
Saccardo, Michelia Commentarium Mycol-
ogicum Fungos in Primis Italicos Ilustrans
2:599-600. 1882. On dead stems of Laser-
pitium angustifolium Georgi, Umbelliferae.
France.

oreophiloides P.A. Saccardo subsp. scrophulariae
P.A. Karsten, Hedwigia, Dresden 22:178.
1883; Acta Societatis pro Fauna et Flora
Fennica, Helsingforsiae 2(6):51. (1884) 1885.
On dead stems of Scrophularia nodosa L.,
Scrophulariaceae. Finland.

ornithogali §.A. Gucevicz, Novosti Sistematiki
Nizshikh Rastenij, Novitates Systematicae
Plantarum Non Vascularium 4:209-210.
1967. On dead stems of Ornithogalum
pyrenaicum L., Liliaceae. U.S.S.R. (Crimea).

orthogramma (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:60. 1883. =Sphaeria orthogramma M.J.
Berkeley & M.A. Curtis, Grevillea, London
4:144-145. 1876 [as orthogrammi]. On stems
of Erianthus alopecuroides (L.) Ell., Gramin-
eae. U.S.A. (Pennsylvania, South Carolina).

orthrosanthi E. Miiller in E. Miiller and R.W.G.
Dennis, Kew Bulletin. Royal Botanic
Gardens, Kew 19:373—375, 384. 1965 [as
orthosanthi]. On leaves of Orthrosanthus
chimboracensis Baker (=Orthrosanthus),
Iridaceae. Venezuela.

oryzae A. Cattaneo, Atti dell’ Istituto Botanico della
Universita e Laboratorio Crittogamica di
Pavia, Milano 2 & 3:127. 1879. =Metas-
phaeria oryzae (A. Cattaneo) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:180.
1883. On culms and leaves of Oryza sativa
L., Gramineae. China, Italy.

oryzicola K. Hara, A Monograph of Rice Diseases,
p. 113. 1959 [as oryzaecola]. On living plant
of Oryza sativa L., Gramineae. Japan.

(Leptosphaerella) oryzina P.A. Saccardo, Atti dell’
Accademia Scientifica Veneto-Trentino-
Istriana, Padova 10:67—-68. 1919. On dead
glumes of Oryza sativa L., Gramineae.
Philippines.

(Clypeosphaeria) osculanda (C.G.T. Preuss) M.C.
Cooke, Grevillea, London 17:91. 1889.
=Sphaeria osculanda C.G.T. Preuss, Linnaea.
Institut Botanique de |’ Université de Geneve
26:713. 1853. =Clypeosphaeria osculanda
(C.G.T. Preuss) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:91. 1883. On
branches of Rubus sp., Rosaceae. Germany.

oubanguiensis A.M. Saccas, Etude de la Flore
Cryptogamique des Caféiers en Afrique

Nomenclator of Leptosphaeria

i)
wn
vo)

Centrale. Bulletin de Institut Frangais du Café
du Cacao et D’ Autres Plantes Stimulantes
(Bulletin IFCC No. 16), pp. 257-259. 1981.
Nom. inval. Art. 37.1. On dead stems of
Coffea robusta L. Linden (=Coffea canephora
Pierre ex Froehn.), Rubiaceae. Central
African Republic.

owaniae K. Kalchbrenner & M.C. Cooke, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:33. 1883 [as
Sphaeria owaniae K. Kalchbrenner & M.C.
Cooke, Grevillea, London 9:29. 1880; falsely
so cited in P.A. Saccardo]. On stems of
Artemisia sp., Compositae. South Africa.

oxyriae E. Rostrup, Meddelelser om Grgnland,
Kjobenhavn 3:559. 1888. On dry stems of
Oxyria digyna (L.) Hill, Polygonaceae.
Greenland.

oxyspora J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg
1(2):165—166. 1901. On dry stems of Rubus
idaeus L., Rosaceae. Luxembourg.

pachyasca G. Niessl v. Mayendorf, Hedwigia,
Dresden 21:111. 1882. Nom. illegit. Art. 64.1
[as pachyascus|. =Metasphaeria pachyasca
(G. Niessl v. Mayendorf) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:171.
1883. On dead leaves of Campanula zoysii
Wulfen, Campanulaceae. Yugoslavia.

pachyasca G. Niessl v. Mayendorf, Osterreichische
Botanische Zeitschrift, Wien 31:345—346.
1881 [as pachyascus]. =Metasphaeria
pachyasca (G. Niessl v. Mayendorf) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:171-172. 1883. =Sphaerulina pachyasca
(G. Niessl v. Mayendorf) A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:126. 1894. =Pseudosphaeria pachyasca (G.
Niessl v. Mayendorf) F. v. Héhnel,
Sitzungsberichte der Akademie der Wissen-
schaften in Wien, Mathematisch-naturwissen-
schaftliche Klasse, Abt. I, 116:635. 1907.
=Saccothecium pachyascum (G. Niessl v.
Mayendorf) W. Kirschstein, Annales Mycol-
ogici, Berlin 37:105. 1939. =Wettsteinina
pachyasca (G. Niessl v. Mayendorf) F. Petrak,
Sydowia. Annales Mycologici, Horn, N.O.
1:56. 1947. [Combination not actually made! ]
On dead or overwintered leaves, on leaves of
Androsace lactea L., Campanula zoysii
Wulfen, Primulaceae, Campanulaceae.
Yugoslavia,

pachycarpa P.A. Saccardo & E. Marchal, Revue
Mycologique, Toulouse 7:145. 1885. On
culm of grass of unknown host, Gramineae.
Unknown country.

pachytheca P.A, Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze, New Series 22:39. 1915.

260

Nom. illegit. Art. 64.1. =Trematosphaeria
pachycarpa (P.A. Saccardo & E. Marchal)
R.A. Shoemaker, Canadian Journal of Botany,
Ottawa 67:1595. 1989. On dead branches of
Amygdalus communis L. [=Prunus dulcis
(Mill.) D.A. Webb], Rosaceae. Yugoslavia.
pachytheca P.A. Hariot & P.A. Briard, Revue
Mycologique, Toulouse 12:178. 1890. On dry
culms and leaves of Nardus stricta L.,
Gramineae. France.
pacifica H. Rehm, Annales Mycologici, Berlin
9:364. 1911. On leaves of Yucca sp. [as
Yucca colusplei], Agavaceae. U.S.A.
paludosa J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg
1(2):157—158. 1901. On fallen leaves of
Carex vesicaria L., Cyperaceae. Luxem-
bourg.
palustris E. Miller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):207. 1950. =Massari-
osphaeria palustris (E. Miiller) A. Leucht-
mann, Sydowia. Annales Mycologici, Horn,
N.O. 37:171. 1984. On dead stems of Typha
latifolia L., Typhaceae. Switzerland.
palustris J. Schroter, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
15:191. 1901. [Falsely so cited in P.A.
Saccardo; for Leptospora palustris J. Schroter
in F.J. Cohn, Jahresbericht der Schlesischen
Gesellschaft fiir vaterlandische Kultur,
Breslau 61:179. 1884.] =Ophiobolus
palustris (J. Schroter) A.N. Berlese & P.
Voglino, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
Additamenta Ad Volumina I-IV, p. 190. 1886.
pampaniniana P.A. Saccardo, Annales Mycologici,
Berlin 11:566. 1913. On decayed leaves of
Stipa tenacissima L., Gramineae. Libya.
pampini (P.A. Saccardo) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 15:191. 1901.
=Sphaerella (Leptosphaeria) pampini P.A.
Saccardo, Revue Mycologique, Toulouse 4:1.
1882. =Metasphaeria pampini (P.A.
Saccardo) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:164. 1883 [as
pampinea]. On vines of Vitis sp., Vitaceae.
France.
pampini (F. v. Thiimen) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:31. 1883. =Sphae-
rella pampini F. v. Thiimen, Die Pilze des
Weinstockes, p. 27. 1878. On wilted, living
runners of Vitis vinifera L., Vitaceae. Italy.
pandani F. Tassi, Bollettino del R. Orto Botanico,
Siena 6:125. 1904. On dry leaves and sheaths
of Pandanus utilis Bory, Pandanaceae. Italy.
pandanicola K. Sawada, Descriptive Catalogue of
Taiwan (Formosan) Fungi, Part XI (Special
Publication of the National Taiwan University

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Taipaei No. 8), pp. 71-72. 1959. Nom. inval.
Art. 36.1. On leaves of Pandanus odoratis-
simus Nor. (=Pandanus laevis Lowr.),
Pandanaceae. Taiwan.

panici H. Sydow, Leaflet of Philippine Botany,
Manila 9:3121—3122. 1925. =Phaeosphaeria
panici (H. Sydow) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1535. 1989. On
dead leaves of Panicum palmaefolium Koen..,
Gramineae. Philippines.

paoluccii M. Curzi, Atti dell’ Istituto Botanico della
Universita e Laboratorio Crittogamica di
Pavia, Milano, Series 3, 3:204. 1927. In spots
on upper surface and margins of leaves of
Laurus nobilis L., Lauraceae. Italy.

papaveris E. Rostrup, Botanisk Tidsskrift, Kjoben-
havn 25:305-306. 1930. On stems of
Papaver radicatum Rottb., Papaveraceae.
Iceland.

papillata (H.F. Bonorden) P.A. Saccardo & G.B.
Traverso, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
19:1109. 1910. =Myriocarpium papillatum
H.F. Bonorden, Abhandlungen auf dem
Gebiete der Mykologie, Halle 1:154—155.
1864. On branchlets of unknown host,
unknown family. Germany.

papillosa M. de Sousa da Camara, Agronomia
Lusitana, Sacavém 13:127. 1951. On culms
of Oryzopsis miliacea (L.) Bentham &
Hooker ex Ascherson & Graebner, Gramin-
eae. Portugal.

papulosa (M.C. Durieu de Maisonneuve & J.P.
Montagne) P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:313. 1875.
=Sphaeria papulosa M.C. Durieu de Maison-
neuve & J.P. Montagne in M.C. Durieu de
Maisonneuve, Exploration Scientifique de
L’ Algérie Publiée Par Ordre du Gouverne-
ment. Series Naturelles. Botanique I, pp.
536-537. 1868. =Metasphaeria papulosa
(M.C. Durieu de Maisonneuve & J.P.
Montagne) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:168—169. 1883. On somewhat
rotten leaves of Citrus aurantium L., Hedera
helix L., Smilax mauritanica Poiret, Rutaceae,
Araliaceae, Liliaceae. Algeria.

papyri P.A. Saccardo & P. Sydow in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 16:514-.
1902. =Leptosphaeria papyricola F. Tassi.
Nec J.B. Ellis & B.M. Everhart. On dry
peduncles of Cyperus papyrus L., Cyperaceae.
China.

papyricola F. Tassi, Bolletino del R. Orto Botanico,
Siena 2:141. 1899. Nom. illegit. Art. 64.1.
=Leptosphaeria papyri P.A. Saccardo & P.
Sydow.

papyricola J.B. Ellis & B.M. Everhart, Proceedings
of the Academy of Natural Sciences of

:
)

March 1991

Philadelphia 1894:337. 1894. On old
pasteboard lying by roadside. U.S.A.

papyricola L. Vouaux in M. Bouly de Lesdain,
Recherches sur les Lichens des Environs de
Dunkerque, p. 73. 1912. Nom. illegit. Art.
64.1. On cardboard. France.

paraguariensis A. Maublanc, Bulletin de la Société
Mycologique de France, Paris 36:35. 1920.
On leaves of /lex paraguariensis A. St. Hil.,
Aquifoliaceae. Brazil.

parietariae P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):154. 1873. On rotten stems of Parietaria
officinalis L., Urticaceae. Italy.

parietariae P.A. Saccardo forma lamii P.A.
Saccardo, Fungi Selecti Gallici Exsiccati,
Century 72, No. 7138. Anno 1897; Revue
Mycologique, Toulouse 19:149-150. 1897.
On dry stems of Lamium album L., Labiatae.
France?

parmeliarum (W. Phillips & C.B. Plowright) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:83. 1883. =Sphaeria parmeliarum W.
Phillips & C.B. Plowright, Grevillea, London
4:124. 1876. On unknown substrate of
Parmelia saxatilis (L.) Arch., Lichenes
(Parmeliaceae). Great Britain.

parvula G. Niessl v. Mayendorf, Verhandlungen des
Naturforschenden Vereins in Briinn
10:173-174. (1871) 1872; Hedwigia, Dresden
12:119. 1873. =Phaeosphaeria parvula (G.
Niess! v. Mayendorf) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:109. 1984. On wilting leaves of /ris
pseudacorus L., Iridaceae. Austria, Czecho-
slovakia, Switzerland.

parvula G. Niessl v. Mayendorf var. iridis-germani-
cae C.B. Massalongo ex C.B. Massalongo,
Madona Verona. (Verona. Museo Civico)
2(1):38. 1908. On leaves of /ris germanica
L., Iridaceae. Italy.

parvula G. Niess| vy. Mayendorf var. iridis-germani-
cae C.B. Massalongo, Novitates Florae My-
cologicae Veronensis (Fungi Potissimum in
Valle Tremniacensi Detecti) Cum 10 Tabulis
Coloratis, Verona, p. 65. 1902. Nom. inval.
Art. 33.1. On leaves of /ris germanica L.,
Iridaceae. Italy.

passeriniana P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:785. 1891. =Leptosphaeria
asparagi G. Passerini. Non C.H. Peck. On
dry stems of Asparagus officinalis L.,
Liliaceae. Italy.

passerinii P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:37—38. 1877. On stems of
Scabiosa columbaria L., Dipsacaceae. Italy.

passerinii P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.

Nomenclator of Leptosphaeria

261

Saccardo 2:53. 1883. Nom. illegit. Art. 64.1.
=Leptosphaeria marginalis G. Passerini. On
wilted leaves of Achillea ptarmica L.,
Compositae. Italy.

patellaeformis G. Passerini, Atti dell’ Accademia
Nazionale dei Lincei, Rendiconti, Roma,
Series 4, 4(2):59. 1888. On wilted culms of
Zea mays L., Gramineae. Italy.

paucispora A.B. Cribb & J.W. Cribb, Papers from
the Department of Botany, University of
Queensland, Brisbane 4(3):41-44. 1960. On
wood of unknown host, unknown family.
Australia (Queensland).

pelagica E.B. Gareth Jones, Transactions of the
British Mycological Society, London 45:105.
1962. On dying culms of Spartina sp.,
Gramineae. Great Britain.

pelargonti H. Rehm, Hedwigia, Dresden 44:5. 1904
{as pelagerinii]. On leaves of cultivated
Pelargonium sp., Geraniaceae. Brazil.

pellita (F. Currey) P.A. Saccardo, Michelia Com-
mentarium Mycologicum Fungos in Primis
Italicos Illustrans 2:63. 1880 [as Lepto-
sphaeria pellita G.L. Rabenhorst & K.
Kalchbrenner]. =Sphaeria pellita F. Currey,
Transactions of the Linnean Society of
London 22:331. 1859. Nec Sphaeria pellita
E.M. Fries:E.M. Fries. Nec Pleospora pellita
(E.M. Fries:E.M. Fries) E.L.R. Tulasne. On
various stems of unknown host, unknown
family. Unknown country.

peltigerarum F. Armold, Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wien
28:271. 1878. =Ophiobolus peltigerarum (F.
Arnold) A.N. Berlese & P. Voglino, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo Additamenta Ad
Volumina I-IV, p. 189. 1886. On thallus of
Peltigera aphthosa (L.) Willd. forma
complicata Th. Fries, Lichenes (Peltiger-
aceae). Italy.

peltigerea (G.K. Merrill) ex L. Vouaux, Bulletin de
la Société Mycologique de France, Paris
29:119-120. 1913. =Trypethelium (Mela-
notheca) peltigereum G.K. Merrill, Lichenes
Exsiccati, Series I, Fascicle 4, No. 85. Anno
1910. Nom. inval. Art. 32.1. On thallus of
Peltigera sp., Lichenes (Peltigeraceae).
Jamaica.

penicillus (E.M. Fries:E.M. Fries) P.A. Saccardo,
Atti dell’ Accademia Scientifica Veneto-
Trentino-Istriana, Padova 2(2):258. 1873.
=Sphaeria penicillus Schmidt, unpublished.
=Sphaeria penicillus E.M. Fries:E.M. Fries,
Systema Mycologicum Sistens Fungorum
2:508. 1823. =Rhaphidospora penicillus
(E.M. Fries:E.M. Fries) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:233. 1863.
=Pyrenophora penicillus (E.M. Fries:E.M.

262

Fries) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:282. 1883 [as (Schmidt)].
=Pleospora penicillus (E.M. Fries:E.M. Fries)
L. Fuckel, Symbolae Mycologicae Dritter
Nachtrag, pp. 23-24. 1873. On stem of
Peucedanum cervaria (L.) Lapeyr., Umbellif-
erae. Austria, Germany.

penicillus P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:316. 1875. Non (E.M.
Fries:E.M. Fries) P.A. Saccardo. 1873. Nom.
illegit. Art. 64.1. On stems of Peucedanum
cervaria (L.) Lapeyr., Umbelliferae. Italy.

penniseti C.G. Hansford, Proceedings of the Linnean
Society of London 153:24. 1941. On dead
culms of Pennisetum purpureum Schum.,
Gramineae. Uganda.

penniseticola F.C. Deighton, Sydowia. Annales
Mycologici, Horn, N.O. 6(1-4):311. 1952.
On living leaves of Pennisetum purpureum
Schum., Gramineae. Sierra Leone.

perforans (M.R. Roberge & J. Desmaziéres) B.
Auerswald in M. Marcucci, Unio Itineraria
Cryptogama Exsiccata No. XXXIII. Anno
1866. =Sphaeria perforans M.R. Roberge &
J. Desmaziéres in J. Desmaziéres, Plantes
Cryptogames Du Nord De La France, Edition
1, Fascicle 26, No. 1288. Anno 1843; Annales
des Sciences Naturelles, Paris, Botanique,
Series 2, No. 19, p. 357. 1843. =Sphaerella
perforans (M.R. Roberge & J. Desmaziéres)
V. Cesati & G. de Notaris, Commentario della
Societa Crittogamologica Italiana, Milan
1:238. 1863. On leaves of Ammophila
arenaria (L.) Link, Gramineae. France, Italy.

periclymeni C.A. Oudemans, Nederlandsch
kruidkundig Archief. Verslagen en Med-
edeelingen der Nederlandsche Botanische
Vereeniging, Leiden, Series 2, 5:168. 1888.
On branches of Lonicera periclymenum L.,
Caprifoliaceae. Netherlands.

periclymeni C.A. Oudemans var. tatarica A.A.
Potebnia, Annales Mycologici, Berlin 5:14.
1907. On living twigs of Lonicera tatarica
L., Caprifoliaceae. U.S.S.R.

perplexa P.A. Saccardo & C.E. Fairman in P.A.
Saccardo, Journal of Mycology, Columbus,
Ohio 12:49. 1906. On dead stems of Solidago
sp., Compositae. U.S.A.

perpusilla (J. Desmazieres) P.A. Karsten, Mycologia
Fennica Pars 2, Pyrenomycetes, p. 99. 1873.
=Sphaeria (folliicola) perpusilla J. Des-
mazieres, Annales des Science Naturelles,
Paris, Botanique, Series 3, 6:80. 1846.
=Laestadia perpusilla (J. Desmaziéres) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
1:423. 1882. On stems of Phragmites
communis Trin., Gramineae. Belgium,
France, Great Britain, Italy.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

perpusilla (J. Desmaziéres) P.A. Karsten var. typhae
P.A. Karsten, Mycologia Fennica Pars 2,
Pyrenomycetes, p. 99. 1873. =Sphaeria
perpusilla J. Desmaziéres var. typhae B.
Auerswald in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi
Mycologici Continuatio Edita Nova, Series
Secunda, Century 9, No. 831. Anno 1865.
Nom. nud. Art. 32.1. =Leptosphaeria typhae
(P.A. Karsten) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:321.
1875. [P.A. Saccardo, Fungi Italici autogra-
phice delineati (additis nonnullis extra-italicis
asterisco notatis), Patavii, Table 491. 1879:
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Diggesit P.A. Saccardo
2:64. 1883.] On rotting culms of Typha
latifolia L., Typhaceae. Finland.

personata G. Niessl v. Mayendorf in G.L. Raben-
horst, Fungi Europaei Exsiccati, Klotzschii
Herbarii vivi Mycologici Continuatio Edita
Nova, Series 2, Century 20, No. 1933. Anno
1875; Botanische Jahrbiicher fiir Systematik,
Pflanzengeschichte und Pflanzengeographie,
Leipzig 3:262. (1875) 1877. =Scleropleella
personata (G. Niessl v. Mayendorf) F. v.
Hohnel, Annales Mycologici, Berlin 16:158.
1918. =Mycotodea personata (G. Niessl v.
Mayendorf) W. Kirschstein, in O.C. Schmidt,
Kryptogamenflora der Mark Brandenburg und
Angrenzender Gebiete herausgegeben von
dem Botanishen Verein der Provinz Bran-
denburg 7(3):433. 1938. =Leptosphaerulina
personata (G. Niessl v. Mayendorf) M.E.
Barr, Contributions from the Univerity of
Michigan Herbarium, Ann Arbor 9:542. 1972.
On dry stems of Aira cespitosa L., Gramineae.
Austria.

peruviana C.L. Spegazzini, Anales de la Sociedad
Cientifica Argentina, Buenos Aires 12:179.
1881; Fungi Argentini Pugillus 4, No. 168.
1881. =Leptosphaeria promontorii P.A.
Saccardo. Nec M.C. Cooke.

petiolaris J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg
1(3):216-217. 1903. On petioles of Juglans
regia L., Juglandaceae. Luxembourg.

petiolicola P.A. Saccardo, Atti dell” Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):153. 1873. On petioles of Robinia
pseudacacia L., Leguminosae. Italy.

petkovicensis F. Bubak & N. Ranojevic in N.
Ranojevic, Annales Mycologici, Berlin 8:361.
1910. =Metasphaeria petkovicensis (F.
Bubak & N. Ranojevic) W. Kirschstein,
Annales Mycologici, Berlin 37:134. 1939.
=Phaeosphaeria petkovicensis (F. Bubak &
N. Ranojevic) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1508. 1989. On

March 199]

dry culms of Juncus effusus L., Juncaceae.
Czechoslovakia, Germany, Great Britain,
Switzerland, Yugoslavia.

petkovicensis F. Bubak & N. Ranojevi¢ var. elymi
L.E. Wehmeyer, Mycologia, Lancaster,
Pennsylvania 55:322. 1963. =Phaeosphaeria
elymi (L.E. Wehmeyer) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1503.
1989. On stems of Blysmus compressus (L.)
Panzer ex Link (=Scirpus), Cyperaceae.
India.

petrakii P.A. Saccardo in F. Petrak, Annales
Mycologici, Berlin 12:287. 1914. On
diseased culms of Luzula nemorosa (Pollich)
E.H.F. Meyer, Juncaceae. Czechoslovakia.

petri J. Chevaugeon, Encyclopedie Mycologique,
Paris 28:24. 1956. On leaves of Manihot
utilissima Pohl, Euphorbiaceae. Ivory Coast.

phacae E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 5(3-6):49-S0. 1951. On dead
culms of Phaca alpina L. (=Astragalus),
Leguminosae. Switzerland.

phaeospora E. Miiller, Sydowia. Annales Mycol-
ogici, Horn, N.O. 4(1—-6):208. 1950. =Mas-
sariosphaeria phaeospora (E. Miiller) P.G.
Crivelli, Ueber die Heterogene
Ascomycetengattung Pleospora Rabh.;
Vorschlag fiir Eine Aufteilung. Abhandlung
zur Erlangung des Titels eines Doktors de
Naturwissenchaften der Eidgendssischen
Technischen Hochschule, Ziirich, No.
7318:141. 1983. =Trematosphaeria
phaeospora (E. Miller) L. Holm, Symbolae
Botanicae Upsalienses, Uppsala 14(3):165.
1957. =Chaetomastia phaeospora (E. Miiller)
M.E. Barr, Mycotaxon. An International
Journal Designed to Expedite Publication of
Research on Taxonomy & Nomenclature of
Fungi & Lichens, Ithaca, New York 34:512.
1989. On dead stems of Artemisia campestris
L., Compositae. Switzerland.

phaeosticta B. Auerswald in W. Gonnermann and
G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Sammtlicher Pilze Europa’s,
Hefte 5 und 6, Synopsis Pyrenomycetum
Europaeorum, Table 11, Figure 154. 1869.
Nom. nud. Art. 32.1. [Sphaeria phaeosticta
M.J. Berkeley is a facultative synonym, fide
P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:374. 1878.] =Anthostomella
phaeosticta (M.J. Berkeley) P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 1:374.
1878. Europe.

phaseoli F. Fautrey & C. Roumeguére in C.
Roumeguére, Revue Mycologique, Toulouse
14:6. 1892; Fungi Selecti Gallici Exsiccati,
Century 60, No. 5951. Anno 1892. On dry
stems of Phaseolus vulgaris L., Leguminosae.
France.

Nomenclator of Leptosphaeria

263

phaseolorum J.B. Ellis & B.M. Everhart, American
Naturalist, Lancaster, Pennsylvania
31:341—342. 1897. On old bean vines (stems)
of Phaseolus vulgaris L., Leguminosae.
U.S.A.

phiala (M.C. Durieu de Maisonneuve & J.P.
Montagne) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:19. 1883. =Sphaeria phiala M.C.
Durieu de Maisonneuve & J.P. Montagne in
M.C. Durieu de Maisonneuve, Exploration
Scientifique de L’ Algérie Publi¢e Par Ordre
du Gouvernement. Series Naturelles.
Botanique 1:519. 1868. On cortex of
branches of Genista ferox Poir., Leguminosae.
Algeria.

phlogis C.A. Oudemans, Verslagen van de Gewone
Vergadering der Wis-en Natuurkundige
Afdeeling. K. Academie van Wetenschappen
Te Amsterdam 9(2):141. 1900. On leaves of
Phlox decussata Lyon, Polemoniaceae.
Netherlands.

phoenicis §.T. Tilak, Mycopathologia et Mycologia
Applicata, Den Haag 28:83—84. 1966. On
dried petioles of Phoenix dactylifera L.,
Palmae. India (Aurangabad).

phoradendri L. Bonar, Proceedings of the California
Academy of Sciences, San Francisco, Series
4, 22:198-199. 1939. On living leaves of
Phoradendron townsendii Trel., Viscaceae.
Chile (Socorro Island).

phormicola M.C. Cooke & H.W. Harkness,
Grevillea, London 14:10. 1885. On unknown
substrate of Phormium sp., Agavaceae.
U.S.A.

phormii W.B. Grove, Kew Bulletin. Royal Botanic
Gardens, Kew 1921:150. 1921. On dead
leaves of Phormium sp., Agavaceae. Great
Britain (Scotland).

phragmiticola (P.L. Crouan & H.M. Crouan) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:87. 1883. =Pleospora phragmitecola P.L.
Crouan & H.M. Crouan, Florule Du Finistere
Contenant Les Descriptions De 360 Espeéces
Nouvelles De Sporogames, De Nombreuses
Observations et une synonymie de plantes
Cellulaires et Vasculaires Qui Croissent
Spontanément Dans ce Département, p. 23.
1867. On culms of Phragmites communis
Trin., Gramineae. France.

phyllachoricola F. Petrak, Sydowia. Annales
Mycologici, Horn, N.O. 2(1-6):337-338.
1948. On stroma of Phyllachora disseminata
Syd., Fungi (Phyllachoraceae). Ecuador.

phyllachorivora F. Petrak, Annales Mycologici,
Berlin 20:302—303. 1922. Parasitic on stroma
of Phyllachora graminis (Pers.:Fr.) Fuckel,
Fungi (Phyllachoraceae). Czechoslovakia.

(Metasphaeria) phyllostachydis K. Hara, Botanical
Magazine, Tokyo 27(317):250. 1913 [as

264

phllostachydis|. On unknown substrate of
Phyllostachys bambusoides Sieb. & Zucc. var.
bambusoides [as Phyllostachys reticulata},
Gramineae. Japan.

physalidis J.B. Ellis & B.M. Everhart, Bulletin of the
Torrey Botanical Club (and Torreya), New
York 11:42. 1884. On dirty white round spots
on leaves of Physalis pubescens L., Sola-
naceae. U.S.A.

physostegiae C.E. Fairman, Proceedings of the
Rochester Academy of Science, Rochester,
New York 4:216. 1906. On dead stems of
Physostegia virginiana (L.) Benth., Labiatae.
U.S.A.

phyteumatis (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:37. 1883. =Ple-
ospora (Nodulosphaeria) phyteumatis L.
Fuckel, Fungi Rhenani Exsiccati A Leopoldo
Fuckel Collecti, Fascicle XX V(X), No. 2439.
Anno 1872; Symbolae Mycologicae Zweiter
Nachtrag, p. 25. 1873. On dry stems of
Phyteuma spicatum L., Campanulaceae.
Switzerland.

phyteumatis (L. Fuckel) H.G. Winter forma knautiae
J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(2):164.
1901. On unknown substrate of Knautia
arvensis (L.) Coulter, Dipsacaceae. Luxem-
bourg.

phytolaccae F. Cavara, Revue Mycologique,
Toulouse 11:181. 1889. On old stems of
Phytolacca decandra L., Phytolaccaceae.
Italy.

picastra (E.M. Fries:E.M. Fries) F. v. Hohnel,
Mitteilungen des Botanischen Instituts der
Technischen Hochschule, Wien 4:44. 1927.
=Trematosphaeria morthieri L. Fuckel,
Symbolae Mycologicae Nachtrag 1:306. 1871.
=Trematosphaeria picastra (E.M. Fries:E.M.
Fries) L. Fuckel, Symbolae Mycologicae, p.
162. 1870. =Sphaeria picastra E.M.
Fries:E.M. Fries, Kungliga Svenska
Vetenskapsakademiens Handlingar, Uppsala
and Stockholm 38:260—261. 1817; E.M. Fries,
Systema Mycologicum Sistens Fungorum
2:463. 1823. On resin-hardened wood of
Pinus sylvestris L., Pinaceae. Sweden.

picridis F. Fautrey & J.B. Lambotte, Revue
Mycologique, Toulouse 16:75—76. 1894. On
unknown substrate of Picris hieracioides L.,
Compositae. France.

pilulariae A. Ade, Kryptogramische Forschungen,
Minchen 2:26. 1929. On dead leaves of
Pilularia sp., Marsileaceae. Germany
(Bavaria).

pimpinellae R. Lowen & A. Sivanesan, Mycotaxon.
An International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens, Ithaca,
New York 35:205—209. 1989. On standing

Illinois Natural History Survey Bulletin

Vol. 34 Ant. 3

dead stems of Pimpinella anisum L., Umbel-
liferae. Israel.

pini (D. Cruchet) E. Miller, Sydowia. Annales
Mycologici, Horn, N.O. 4(1-6):277. 1950.
=Metasphaeria pini D. Cruchet, Bulletin de la
Société Vaudoise des Sciences Naturelles,
Lausanne 55:167. 1923. On cones of Pinus
sylvestris L., Pinaceae. France, Switzerland.

pinnarum G. Passerini, Revue Mycologique,
Toulouse 2:34. 1880. =Metasphaeria
pinnarum (G. Passerini) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:179.
1883. On soft pinnae of Phoenix dactylifera
L., Cyperaceae. Italy.

pinnarum G. Passerini var. rachidis G. Passerini,
Revue Mycologique, Toulouse 2:34. 1880 [as
pinnosum). =Metasphaeria rachidis (G.
Passerini) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:180. 1883. On pinnae of Phoenix
dactylifera L., Palmae. Italy.

piperis P.C. Hennings in H.G.A. Engler, Die
Pflanzenwelt Ost-Afrikas und der Nach-
bargébiete, Theil C, p. 33. 1895. On leaves of
Piper capense L., Piperaceae. East Africa.

plagia (M.C. Cooke & G.E. Massee) L. Holm,
Svensk Botanisk Tidskrift, Stockholm 62:234.
1968. =Cucurbitaria (Melanomma) plagia
M.C. Cooke & G.E. Massee, Grevillea,
London 17:8. 1888. =Gibberidea plagia
(M.C. Cooke & G.E. Massee) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 9:820.
1891. On living twigs of Cassinia aculeata R.
Br., Compositae. Australia.

planiuscula (H. Riess) V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:236. 1863. =Sphaeria
planiuscula H. Riess, Hedwigia, Dresden 1,
Table IV, Figure 7. 1854. Nom. nud. Art.
32.1; in G.L. Rabenhorst, Klotzschii Herbar-
ium Vivum Mycologicum Sistens Fungorum
Per Totam Germaniam Cresentium Collec-
tionem Perfectam, Dresden, Edition 1,
Century 19, No. 1829. 1854. On dead stems
of Solidago virgaurea L., Compositae.
Finland, Germany, Switzerland.

planiuscula (H. Riess) V. Cesati & G. de Notaris
forma cruciferarum H. Rehm, Hedwigia,
Dresden 22:55—56. 1883. On stems of
Cruciferae. Unknown country.

planiuscula V. Cesati & G. de Notaris forma
succisae J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(2):161. 1901. On unknown substrate of
Succisa pratensis Moench., Dipsacaceae.
Luxembourg.

plantaginicola (N.T. Patouillard) F. Petrak, Sydowia.
Annales Mycologici, Horn, N.O. 4(1-6):496.

March 1991

1950. =Sphaerella? plantaginicola N.T.
Patouillard in N.T. Patouillard and N.G. de
Lagerheim, Bulletin de la Société Mycolo-
gique de France, Paris 9:153. 1893. =Myco-
sphaerella plantaginicola (N.T. Patouillard)
F.L. Stevens, Illinois Biological Monographs,
University of Illinois at Urbana-Champaign
11:199-200. 1927. On leaves of Plantago sp.,
Plantaginaceae. Ecuador.

platanicola (E.C. Howe) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2 (Addenda Ad
Volumen Secundum), p. LVII. 1883.
=Sphaeria platanicola E.C. Howe, Bulletin of
the Torrey Botanical Club (and Torreya), New
York 5:43. 1874. On branchlets of Platanus
sp., Platanaceae. U.S.A. (New York).

platanicola F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 7(1-4):122. 1953. Nom. illegit.
Art. 64.1. On dry stems of Platanus occiden-
talis L., Platanaceae. U.S.A.

platycarpa P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:342—343. 1878. On branches of
Cornus sanguinea L., Cornaceae. Italy.

platychorae E. Miiller, Sydowia. Annales Mycol-
ogici, Horn, N.O. 7(1-4):275-276. 1953. On
stroma of Platychora ulmi (Schleich.:Fr.)
Petrak, Fungi (Venturiaceae). Switzerland.

platypus (L.D. v. Schweinitz) F. Petrak & H. Sydow,
Annales Mycologici, Berlin 21:349. 1923.
=Sphaeria platypus L.D. v. Schweinitz,
Synopsis Fungorum in America Boreali
Media Degentium. Secundum Observationes

=Macrobasis platypus (L.D. v. Schweinitz) K.
Starbick, Bihang till K. Svenska Vetenskaps-
akademiens Handlingar, Stockholm 19(Afd.
III, No. 2):97-98. 1894. On stems of
Anemone virginiana L., Ranunculaceae.
U.S.A. (Pennsylvania).

plectrospora J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(2):162—163. 1901. On dry stems of Galium
mollugo L., Rubiaceae. Luxembourg.

plemeliana G. Niess| v. Mayendorf, Osterreichische
Botanische Zeitschrift, Wien 31:346. 1881;
Hedwigia, Dresden 21:111—112. 1882. On
leaves of Campanula zoysii Wulfen, Cam-
panulaceae. Yugoslavia.

pleosporoides B. Auerswald in G.L. Rabenhorst,
Fungi Europaei Exsiccati, Klotzschii Herbarii
vivi Mycologici Continuatio Edita Nova,
Series 2, Century 13, No. 1253. Anno 1869.
On runners of Clematis vitalba L., Ranuncu-
laceae. France.

pleurospora G. Niess| v. Mayendorf in H. Rehm,
Ascomyceten, Fascicle 19, No. 937. Anno
1888; Hedwigia, Dresden 27:172. 1888.
=Phaeosphaeria pleurospora (G. Niessl v.

Nomenclator of Leptosphaeria 265

Mayendorf) A. Leuchtmann, Sydowia.
Annales Mycologici, Horn, N.O. 37:162.
1984. =Sulcispora pleurospora (G. Niessl v.
Mayendorf) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1594. 1989. On
dry culms of Aira cespitosa L., Gramineae.
Austria, Switzerland, U.S.S.R. (Latvia).

plocamae F. Petrak, Beiblatt zu den Botanischen
Jahrbiichern 62(No. 142), Heft 3:110-111.
1928. On decorticated wood of Plocama
pendula W. Aiton, Rubiaceae. Canary Islands
(Tenerife Island).

plumbaginis N.T. Patouillard, Revue Mycologique,
Toulouse 8:181. 1886. On dead stems of
Plumbago sp., Plumbaginaceae. China.

plurisepta (L.R. Tehon & E.Y. Daniels) J.A. v. Arx
& E. Miiller, Studies in Mycology, Baarn,
Netherlands 9:76. 1975. =Exilispora
plurisepta L.R. Tehon & E.Y. Daniels,
Mycologia, Lancaster, Pennsylvania 19:1 13.
1927. On stems of Erigeron sp., Compositae.
U.S.A.

poae G. Niessl v. Mayendorf in H. Rehm ex G.
Niessl v. Mayendorf, Ascomyceten, Century
13, No. 643. Anno 1881. Nom. nud. Art.
32.1; Hedwigia, Dresden 21:83. 1882.
=Metasphaeria poae (G. Niessl v. Mayendorf
ex G. Niessl v. Mayendorf) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:175.
1883. On dry culms of Poa nemoralis L.,
Gramineae. Czechoslovakia.

poae G. Niessl v. Mayendorf var. agrostidis F.A.
Hazslinszky, Matematikai és Természettudo-
manyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 25(2):131. 1892. On
unknown substrate of Agrostis vulgaris With.,
Gramineae. Hungary.

polaris P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:83. 1883. [Published as Sphaeria No. 8 by
T. Fries, Hedwigia, Dresden 20:59. 1881.] On
thalli of Rhizocarpon geographicum (L.) DC.,
Lichenes (Rhizocarpaceae). Greenland.

polini S.A. Gucevicz, Trudy Gosudarstvennogo
Nikitskogo Botanicheskogo Sada, Yalta
29:189-190. 1959. On dried branches of
Buddleja davidii Franch., Loganiaceae.
U.S.S.R. (Crimea).

politis F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 10(1-6):308-309. (1956) 1957.
On dry stems of Asperula sp., Rubiaceae.
Greece.

polygonati E. Miller & M. TomaSevic, Phytopa-
thologische Zeitschrift, Berlin 29:289. 1957.
On dry overwintered stems of Polygonatum
sp., Liliaceae. Switzerland.

polytrichina A. Racovitza, Memoires du Museum
Nationale d’ Histoire Naturelle, Paris, Series
B, Botanique 10:150. 1959. On dead plant of

266

Polytrichum juniperinum Willd. ex Hedw.
var. alpinum Schimp., Musci (Polytrichaceae).
France.

pomiformis (C.H. Persoon:E.M. Fries) V. Cesati &
G. de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:235. 1863.
=Sphaeria pomiformis C.H. Persoon:E.M.
Fries, Synopsis Methodica Fungorum, p. 65.
1801; E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:455. 1823. On dry stems
of unknown host, unknown family. Europe.

(Leptosphaerella) pomona P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 8:176.
1876. On upper surface of leaves of Pyrus
malus L. (=Malus domestica Borkh.),
Rosaceae. Italy.

pomona P.A. Saccardo forma transilvanica A. Negru
et I. Ditzu, Notulae Systematicae e Sectione
Cryptogamica Instituti Botanici Nomine V.L.
Komarov Academicae Scientiarum U.R.S.S.,
Petropolis 16:152—153. 1963. On fruits of
Crataegus monogyna Jacq., Rosaceae.
Romania.

pontiformis (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:78. 1883. =Ple-
ospora pontiformis L. Fuckel, Symbolae
Mycologicae, p. 139. 1870. =Phaeosphaeria
pontiformis (L. Fuckel) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:134. 1984. On culms and rhizomes of
Cynodon sp., Secale sp., Triticum sp.,
Gramineae. Czechoslovakia, Germany, Great
Britain, Hungary.

porellae M.J. Thirumalachar, Transactions of the
British Mycological Society, London 31:11.
1948. On thallus of Porella sp., Porellaceae.
India (Mysore State).

portoricensis L.A. Alvarez Garcia, Journal of
Agriculture of the University of Puerto Rico,
Rio Pedras 47(1):56. 1963. On leaves of
Asparagus officinalis L., Liliaceae. Puerto
Rico.

poterti G. Passerini, Atti dell’ Accademia Nazionale
dei Lincei. Memoire, Rome, Series 4, 6:459.
(1889) 1890. On wilted stems of Porerium
sanguisorba L., Rosaceae. Italy.

praeclara P.A. Karsten, Hedwigia, Dresden 23:2.
1884; Acta Societatis pro Fauna et Flora
Fennica, Helsingforsiae 2(6):54. 1885. On
dead stems of Asparagus officinalis L.,
Liliaceae. Finland.

praeclara P.A. Karsten forma typhiseda (P.A.
Saccardo & A.N. Berlese) A.N. Berlese,
Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:75. 1892. =Leptosphaeria
typhiseda P.A. Saccardo & A.N. Berlese.

praetermissa (P.A. Karsten) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Digessit P.A. Saccardo 2:26. 1883.
=Sphaeria praetermissa P.A. Karsten, Fungi
Fenniae Exsiccati, Century 9, No. 852. Anno
1869. On dried canes of Rubus idaeus L.,
Rubus odoratus L., Rosaceae. Finland.

pratensis P.A. Saccardo & P.A. Briard, Revue
Mycologique, Toulouse 7:209. 1885. On
dried stems of Medicago sativa L., Legumino-
sae. France.

preandina (C.L. Spegazzini) L. Holm, Svensk
Botanisk Tidskrift, Stockholm 62:235. 1968.
=Gibberidea praeandina C.L. Spegazzini,
Anales del Museo Nacional de Historia
Natural de Buenos Aires 23:59-60. 1912. On
dead branches of Eupatorium saucechicoense
Hieron., Compositae. Argentina (Mendoza).

primulaecola (H.G. Winter) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:49. 1883. =Sphae-
rella primulaecola H.G. Winter, Hedwigia,
Dresden 19:166. 1880. =Sphaerulina
primulicola (H.G. Winter) A.N. Berlese,
Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:123. 1894. =Scleropleella
primulicola (H.G. Winter) F. v. Hohnel,
Annales Mycologici, Berlin 18:76. 1920.
=Mycodotea primulicola (H.G. Winter) W.
Kirschstein in O.C. Schmidt, Krypto-
gamenflora der Mark Brandenburg und
Angrenzender Gebiete herausgegeben von
dem Botanishen Verein der Provinz Bran-
denburg, Leipzig 7(3):432. 1938. On leaves
and petioles of Primula latifolia Lapeyr.,
Primulaceae. Switzerland.

primulana A. Allescher, Bericht der Bayerischen
Botanischen Gesellschaft zur Erforschung der
Heimischen Flora, Miinchen 5:14. 1897. On
dry stems of Primula elatior (L.) Hill,
Primulaceae. Germany.

priuscheggiana F. Petrak, Annales Mycologici.,
Berlin 16:225. 1919. On rotting leaves of
Tilia sp., Tiliaceae. France.

proliferae? J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg 1(4):50.
1905. On dry stems of Dianthus prolifer L..
Caryophyllaceae. Luxembourg.

promontorii P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:22-23. 1883. =Leptosphaeria
peruviana C.L. Spegazzini, Anales de la
Sociedad Cientifica Argentina, Buenos Aires
12:179. 1881; Fungi Argentina Pugillus 4, No.
168. 1881. On fallen decaying stems of
Salicornia peruviana H.B.K., Chenopodi-
aceae. Argentina.

protearum H, Sydow & P. Sydow, Annales
Mycologici, Berlin 10:441—442. 1912. On
leaves of Protea melaleuca R. Br., Proteaceae.
South Africa.

March 1991

proteispora C.L. Spegazzini, Anales des Museo
Nacional de Buenos Aires 6:282—283. 1898;
Fungi Argentini Novi vel Critici, No. 629.
Anno 1899. On decaying culms of Paspalum
giganteum Baldw. ex Vasey, Gramineae.
Argentina.

pruni N.N. Woronichin, Vestnik Tiflisskogo
Botanicheskogo Sada, Tiflis (Moniteur du
Jardin Botanique de Tiflis) 28:17-18. 1913.
On living leaves of Prunus domestica L.,
Rosaceae. U.S.S.R. (Caucasus).

pruni N.N. Woronichin forma plurivora N.N.
Woronichin, Trudy Akademiia Nauk SSSR
Botanicheskii Muzei 21:125—126. 1927. On
leaves of Chamaerops sp., Palmae. U.S.S.R.
(Caucasus).

pseudo-diaporthe C.A. Oudemans, Nederlandsch
kruidkundig Archief. Verslagen en Med-
edeelingen der Nederlandsche Botanische
Vereeniging, Leiden, Series 2, 6:34. 1892. On
dry stems of Juncus effusus L., Juncaceae.
Netherlands.

pseudohleria C.E. Fairman, Proceedings of the
Rochester Academy of Science, Rochester,
New York 6:127. 1922. On stems of Typha
latifolia L., Typhaceae. U.S.A.

psilospora B. Auerswald, Hedwigia, Dresden 7:184.
1868. On dry stems of Phyteuma scheuchzeri
All., Campanulaceae. Germany.

ptarmicae P.A. Karsten, Fungi Fenniae Exsiccati,
Century 9, No. 888. Anno 1869. On stems of
Achillea ptarmica L., Compositae. Finland.

pterocelastri E.M. Doidge, Bothalia, National
Herbarium, Pretoria 4:859-860. 1948. On
leaves of Pterocelastrus tricuspidatus Walp.,
Celastraceae. South Africa.

puccinioides C.L. Spegazzini in P.A. Saccardo,
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 1:459.
1879. =Metasphaeria puccinioides (C.L.
Spegazzini) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:182. 1883. On fallen leaves of
Carex paludosa Good., Cyperaceae. Italy.

puiggarii C.L. Spegazzini, Fungi Puiggariani
1:143-144. No. 281. 1889. Boletin de la
Academia Nacional de Ciencias en Cordoba
11:521—522. 1889. On dead leaves of
Andropogon saccharoides Swartz, Gramin-
eae. Brazil.

pulchra (H.G. Winter) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:53-54. 1883.
=Sphaerella pulchra H.G. Winter, Hedwigia,
Dresden 11:145-146. 1872. =Mycodotea
pulchra (H.G. Winter) W. Kirschstein,
Annales Mycologici, Berlin 34:201. 1936;
Kryptogamenflora der Mark Brandenburg und
Angrenzender Gebiete herausgegeben von
dem Botanischen Verein der Provinz
Brandenburg, Leipzig 7(3):433. 1938.

Nomenclator of Leptosphaeria

267

=Leptosphaerulina pulchra (H.G. Winter)
M.E. Barr, Contributions de |’Institut
Botanique de L’ Université de Montréal 73:7.
1959. On dry stems and leaves of Potentilla
caulescens L., Rosaceae. Austria.

punctiformis G. Passerini, Atti dell’ Reale Ac-
cademia Nazionale dei Lincei. Rendiconti,
Rome, Series 4, 7(2):45. 1891. On wilted
culms of Zea mays L., Gramineae. Italy.

punctiformis V. Mouton, Bulletin. Société R. de
Botanique de Belgique, Bruxelles 39:44-45.
1900. Nom. illegit. Art. 64.1. =Lepto-
sphaeria moutoniana P.A. Saccardo & P.
Sydow. On culm of grass of unknown host,
Gramineae. Belgium.

punctillum H. Rehm, Annales Mycologici, Berlin
10:356. 1912. =Phaeosphaeria punctillum
(H. Rehm) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):125.
1957. On leaves of Typha latifolia L.,
Typhaceae. Canada.

punctoidea P.A. Karsten, Hedwigia, Dresden 23:2.
1884; Acta Societatis pro Fauna et Flora
Fennica, Helsingforsiae 2(6):53. 1884. On
dead stems of Asparagus officinalis L.,
Liliaceae. Finland.

puniciae A.K. Kar & M.K. Maity, Canadian Journal
of Botany, Ottawa 48:1302. 1970. On
branches of Punica granatum L., Punicaceae.
India (West Bengal).

punjabensis L.E. Wehmeyer, Mycologia, Lancaster,
Pennsylvania 55:322. 1963. On stems of
Berberis petiolicola Wall., Berberidaceae.
India, Pakistan.

purpurea H. Rehm, Ascomycetes Lojkani Lecti in
Hungaria, Transylvania et Galicia, Berlin, p.
54. 1882. =Melanomma purpurea (H. Rehm)
L. Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):64. 1957. On dry stems of
Artemisia vulgaris L., Compositae. Hungary.

purpurearum B. Auerswald in W. Gonnermann and
G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Sammtlicher Pilze Europa’s,
Hefte 5 und 6, Synopsis Pyrenomycetum
Europaeorum, Table 12, Figure 168. 1869.
Nom. nud. Art. 32.1. =Didymella purpure-
arum (B, Auerswald) ex P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 1:549.
1882. On leaves of Salix purpurea L.,
Salicaceae. Germany.

pusilla (C.L. Spegazzini) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 22:999-1000. 1928.
=Leptosphaerella pusilla C.L. Spegazzini,
Boletin de la Academia Nacional de Ciencias
en Cordoba 23:558—559. 1918. On living
leaves of Coffea sp., Rubiaceae. Costa Rica.

puteana J.B. Ellis & W.A. Kellerman in J.B. Ellis
and B.M. Everhart, The North American
Pyrenomycetes. A Contribution to Mycologic

Botany, p. 368. 1892. On inside and outside
of an oaken well bucket, Quercus sp.,
Fagaceae. U.S.A.

puttemansii A. Maublanc, Bulletin de la Société
Mycologique de France, Paris 21:88—89.
1905. On leaves of Eriobotrya japonica
(Thunb.) Lindley, Rosaceae. Brazil.

pycnostigma (W. Nylander) P.A. Saccardo & D.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
17:731. 1905. =Verrucaria pycnostigma W.
Nylander, Flora, Jena und Regensburg 52:297.
1869; in A.F. Malbranche, Bulletin de la
Société des Amis des Sciences Naturelles de
Rouen 6:307. 1870. =Phaeospora pycnos-
tigma (W. Nylander) F. Arnold, Flora, Jena
und Regensburg 57:151. 1874. =Sphaeria
pycnostigma (W. Nylander) H. Olivier,
Bulletin de Géographie Botanique, LeMans
17:169. 1907. On thallus of Baeomyces rufus
(Huds.) Rebent., Lichenes (Baeomycetaceae).
France.

pycnostigma (W. Nylander) P.A. Saccardo & D.
Saccardo var. morbosa (F. Arnold) K.
Keissler in Dr. L. Rabenhorst’s Kryptogaman-
Flora von Deutschland, Oesterreich und der
Schweiz, Second edition, 9(1 Abt., Teil
2):371. 1938. =Sagedia morbosa F. Arnold,
Verhandlungen der Zoologisch-Botanischen
Gesellschaft in Wien 21:1133. 1871. =Porina
morbosa (F. Arnold) G. Lettau, Hedwigia,
Dresden 52:105. 1912. On thalli of unknown
host, unknown family [originally described as
a lichen]. Italy.

pyrenopezizoides P.A. Saccardo & C.L. Spegazzini
in P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:394. 1878. =Heptameria
(Leptosphaeria) pyrenopezizoides (P.A.
Saccardo & C.L. Spegazzini) M.C. Cooke,
Grevillea, London 18:30. 1889. On decaying
twigs of Clematis vitalba L., Ranunculaceae.
Italy.

quadriseptata J.W.H. Trail, Transactions and
Proceedings of the Botanical Society of
Edinburgh 17:491. 1889. On old stems of
Lactuca muralis (L.) Gaertner, Rumex acetosa
L., Thalictrum minus L., Valeriana officinalis
L., Compositae, Polygonaceae, Ranuncu-
laceae, Valerianaceae. Norway.

quamoclidii C.E. Fairman, Mycologia, Lancaster,
Pennsylvania 10:246. 1918. On old stems of
Quamoclidion multiflorum Torr. ex A. Gray,
Nyctaginaceae. U.S.A. (New Mexico).

quinta L.N. Vasil’eva, Pirenomitsety i Lokulo-
askomitsety Severa Dal’nego Vostoka
(Leningrad:“*Nauka”), p. 120. 1987. =Ple-
ospora vagans G. Niessl v. Mayendorf, Ver-
handlungen des Naturforschenden Vereins in
Briinn 14:174—175. (1875) 1876. On Aira

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

cespitosa L., Calamagrostis sylvatica Host,
Elymus arenarius L., Gramineae. Germany.

rajasthanensis K.S. Panwar & S. Kaur, Kavaka,
Madras 5:41-43. 1977. On dead wood of
Lantana camara L., Verbenaceae. India (Mt.
Abu, Rajasthan).

ramalinae (M.R. Roberge) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:84. 1883 [as (Desm.)
Sacc.]. =Sphaeria (Subtecta) ramalinae MLR.
Roberge in J. Desmaziéres, Annales des
Sciences Naturelles, Paris, Botanique, Series
3, 11:354. 1849. On thallus of Ramalina sp..
Lichenes (Ramalinaceae). France.

ramsaugiensis H. Schmid-Heckel, Pilze in den
Berchtesgadener Alpen. Nationalpark
Berchtesgaden Forschungsberichte No. 15
(Regensberg: Nationalpark Bercht-
esgaden):48. 1988. Non vidi. On stem of
Saxifraga paniculata Miller, Saxifragaceae.
Germany.

ramulicola (C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:26-27. 1883.
=Sphaeria ramulicola C.H. Peck, Report.
New York State Museum of Natural History,
Albany 25:104. 1873. =Neodeightonia
ramulicola (C.H. Peck) M.E. Bart in M.E.
Barr, C.T. Rogerson, S.J. Smith, and J.H.
Haines, Bulletin of the New York State
Museum, Albany 459:39. 1986. =Do-
thidotthia ramulicola (C.H. Peck) M.E. Barr,
Mycotaxon. An International Journal
Designed to Expedite Publication of Research
on Taxonomy & Nomenclature of Fungi &
Lichens, Ithaca, New York 34:524. 1989. On
dead twigs of U/mus sp., Ulmaceae. U.S.A.

ranunculi E. Rostrup, Meddelelser om Gronland,
Kjobenhavn 3:558. 1888. On stems of
Ranunculus affinis R. Br., Ranunculaceae.
Greenland.

ranunculi-polyanthemi C. Sandu-Ville & M. Mititiuc
in C. Sandu-Ville, V. Jacob, M. Hatmanu, and
M. Mititiuc, Lucrarile Stintifice, Institutul
Agronomic ‘Professor Ion Ionescu de la
Brad’, Iasi, Bucharest 1967:173—174. 1967.
On dead stems of Ranunculus polyanthemos
L., Ranunculaceae. Romania.

ranunculoides A. Noelli, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze, New Series 19:403. 1912.
On stems of Bupleurum ranunculoides L.,
Umbelliferae. Italy.

raphani D. Hawksworth & A. Sivanesan, Transac-
tions of the British Mycological Society,
London 64:104. 1975. On dead culms of
Raphanus maritimus Sm., Cruciferae. Great
Britain.

raphidophora F.A. Hazslinszky, Matematikai és
Természettudomanyi Kézlemenyek Von-

March 1991

atkozolag a Hazai Viszonyokra, Budapest
25(2):147. 1892. On scapes of Anemone
narcissiflora L., Ranunculaceae. Hungary.

recessa G. Passerini, Atti dell’ Accademia Nazionale
dei Lincei. Memoire, Rome, Series 4, 6:459.
(1889) 1890. =Phaeosphaeria recessa (G.
Passerini) R.A. Shoemaker, Canadian Journal
of Botany, Ottawa 67:1555. 1989. On wilted
stems of Arundo donax L., Gramineae. Italy.

recutita (L. Fuckel) J. Lind, Danish Fungi as
Represented in the Herbarium of E. Rostrup,
Nordisk Forlag, Copenhagen, p. 219. 1913.
=Sphaeria recutita L. Fuckel, Fungi Rhenani
Exsiccati A Leopoldo Fuckel Collecti,
Fascicle 25(10), No. 2434. Anno 1872 [as
(Fr.) Fekel.]. =Sphaeria recutita L. Fuckel,
Symbolae Mycologicae 2:21—22. 1873 [as
Sphaeria recutita (Fr.) Fuckel]. Type based
on Fungi Rhenani Exsiccati A Leopoldo
Fuckel Collecti, Fascicle 25, No. 2434. Anno
1872. Nom. nud. Art. 32.1. Non Sphaeria
recutita E.M. Fries:E.M. Fries, Systema
Mycologicum Sistens Fungorum 2:524. 1823.
=Metasphaeria recutita (L. Fuckel) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:176. 1883. On wilted leaves of Dactylis
glomerata L., Glyceria aquatica (L.)
Wahlenb., Gramineae. Denmark, Germany.

recutita (E.M. Fries:E.M. Fries) E. Miiller [as (L.
Fuckel)], Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):269. 1950. =Sphaeria
recutita E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:524. 1823. =Metas-
Phaeria recutita (E.M. Fries:E.M. Fries) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:176. 1883. On preceding year’s stubble of
Hordeum distichon L., Triticum vulgare L.,
Gramineae. Austria, Germany, Switzerland.

rehmiana W. Voss, Mitteilungen des Musealvereines
fiir Krain, Carniola, Laibach 4(2):19. 1891.
On dead leaves of Drypis spinosa L.,
Caryophyllaceae. Yugoslavia.

rehmii V. Mouton, Bulletin. Société R. de Botanique
de Belgique, Bruxelles 39:44. 1900. On
leaves of grass of unknown host of Gramin-
eae, Gramineae. Italy.

reidiana H. Sydow, Annales Mycologici, Berlin
22:297. 1924. On dead culms of Radicula sp.
(=Rorippa sp.), Cruciferae. New Zealand.

resedae G. Passerini, Atti dell’ Reale Accademia
Nazionale dei Lincei. Rendiconti, Rome,
Series 4, 4(2):58. 1888. On wilted stems of
Reseda lutea L., Resedaceae. Italy.

restionis C.G. Hansford, Proceedings of the Linnean
Society of New South Wales, Sydney 79:121.
1954. On stems of Restio australis R. Br.,
Restionaceae. Australia.

revocans P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo

Nomenclator of Leptosphaeria

269

des A.N. Berlese. Nec F. Fautrey, 1891. On
dead branches of Buxus sempervirens L.,
Buxaceae. Belgium.

rhizomatum G. Passerini, Atti dell’ Accademia
Nazionale dei Lincei, Rendiconti, Roma,
Series 4, 4(2):59. 1888. On rhizomes and
runners of Agrostis vulgaris With., Cynodon
dactylon (L.) Pers., Gramineae. Italy.

rhodiolicola F. Petrak, Annales Mycologici, Berlin
34:444-445. 1936. On stems of Rhodiola
rosea L., Crassulaceae. Finland.

(Melanomma) rhododendri (H. Rehm) M.C. Cooke,
Grevillea, London 17:91. 1889. =Mela-
nomma rhododendri H. Rehm, Ascomyceten,
Fascicle 4, No. 186. Anno 1873. [=Cucurbi-
taria rhododendri G. Niessl v. Mayendorf,
Verhandlungen des Naturforschenden Vereins
in Briinn 10:200. (1871) 1872, is listed as a
synonym in H. Rehm.}] On dry branches of
Rhododendron ferrugineum L., Ericaceae.
Italy.

rhododendri P.C. Hennings, Verhandlungen des
Botanischen Vereins Der Provinz Bran-
denburg, Berlin 40:157. 1898. On leaves of
Rhododendron sp., Ericaceae. Himalayas.

rhodophaea G. Bizzozero, Atti del Istituto Veneto di
Scienze, Lettere ed Arti, Venezia, Series 6,
3:304-305. 1885. =Montagnula rhodophaea
(G. Bizzozero) A. Leuchtmann, Sydowia.
Annales Mycologici, Horn, N.O. 37:176.
1984. On dead culms of Arundo donax L.,
Gramineae. Italy.

rhoina J.B. Ellis & B.M. Everhart, Bulletin of the
Torrey Botanical Club (and Torreya), New
York 24:131. 1897. On decorticated wood of
Rhus glabra L., Anacardiaceae. U.S.A.

rhopalispora A.N. Berlese, cones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:81.
1892. On stems of Crithmum maritimum L.,
Inula crithmoides L., Umbelliferae, Composi-
tae. France.

rhopographoides H. Rehm, Hedwigia, Dresden
41:(204). 1902. =Rhopographus caulincola
C.A. Oudemans, Nederlandsch kruidkundig
Archief. Verslagen en Mededeelingen der
Nederlandsche Botanische Vereeniging,
Leiden, Series 3, 2(3):712. 1902. [Contribu-
tions a la Flore Mycologique des Pays-Bas
XVIII.] Nec L.D. v. Schweinitz. On stem of
unknown host of Umbelliferae, Umbelliferae.
Netherlands.

ribis P.A. Karsten, Revue Mycologique, Toulouse
7:106. 1885. On dry branches of Ribes
alpinum L., Saxifragaceae. Finland.

richoni P.A. Saccardo & P. Sydow, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 14:571. 1899.
=Leptosphaeria stellariae C.E. Richon. Nec
Leptosphaeria stellaria E. Rostrup. On old

stems of Stellaria media (L.) Vill., Caryophyl-
laceae. France.

rimalis G. Niessl v. Mayendorf in J. Kunze, Fungi
Selecti Exsiccati, Century 3, No. 337. 1880.
Nom. nud. Art. 36.1. =Leptosphaeria
agminalis P.A. Saccardo & P. Morthier. Nec
V. Cesati & G. de Notaris. On dry runners of
Clematis vitalba L., Ranunculaceae. Switzer-
land.

rimicola (G.H. Otth) P.A. Saccardo, Hedwigia,
Dresden 35:XXIX. 1896; Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 11:XXIX. 1896. =Cladosphaeria
rimicola G.H. Otth, Mitteilungen der
Naturforschenden Gesellschaft in Bern
1870:106. 1871. Nom. inval. Art. 43.1. [The
genus Cladosphaeria was validated in 1894. ]
On fallen branches of Prunus avium L.,
Rosaceae. Switzerland.

riofrioi R. Gonzalez Fragoso, Memorias de la R.
Sociedad Espanola de Historia Natural,
Madrid 11:92. 1919. On dry stems and
branchlets of Coronilla emerus L., Legumino-
sae. Spain.

riparia P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:39. 1877. On culms of Juncus
effusus L., Juncaceae. Spain.

rivalis J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(2):157.
1901. [Description effectively published:
Vorstudien zu einer Pilz-Flora des Grossher-
zogthums, Luxemburg 1:256. 1899.] On dry
stems, leaves, and leaf sheaths of Carex
paludosa Good., Cyperaceae. Luxembourg.

rivana (G. de Notaris) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:83. 1883.
=Sphaeria rivana G. de Notaris, Commentario
della Societa Crittogamologica Italiana, Milan
2:485. 1867. On thallus of Peltigera canina
(L.) Willd., Lichenes (Peltigeraceae). Italy.

rivana (G. de Notaris) P.A. Saccardo forma
solorinae H. Rehm, Osterreichische Botanis-
che Zeitschrift, Wien 54:84-85. 1904. On
thallus of Solorina crocea (L.) Ach., Lichenes
(Peltigeraceae). Italy.

rivularis E.C. Bommer, M.H. Rousseau & P.A.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:787. 1891. On stems of Alisma
plantago L., Alismataceae. Belgium.

robusta (P. Strasser) E. Miiller, Sydowia. Annales
Mycologici, Horn, N.O. 4(1-6):298. 1950.
=Leptosphaeria derasa (M.J. Berkeley & C.E.
Broome) B. Auerswald forma robusta P.
Strasser, Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien 57:315.
1907. =Nodulosphaeria robusta (P. Strasser)
L. Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):90. 1957. =Ophiobolus

Illinois Natural History Survey Bulletin

Vol. 34 An. 3

robustus (P. Strasser) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 54:2389.
1976. On dry stems of Senecio nemorensis
L., Senecio rupestris Waldst. & Kit.,
Compositae. Switzerland.

roseotinctum J.B. Ellis & B.M. Everhart, The North
American Pyrenomycetes. A Contribution to
Mycologic Botany, p. 237. 1892. Nom. nud.
Art. 32.1. [Apparently an error for Lophio-
stoma roseotinctum J.B. Ellis & B.M.
Everhart.]

rostrata M.L. Farr & H.T. Horner, Nova Hedwigia.
Zeitschrift fiir Kryptogramenkunde, Wein-
heim 15:250—251. 1968. On leaves of
Selaginella arenicola subsp.
acanthonota (Underwood) R. Tryon,
Selaginella rupestris (L.) Spring, Selaginel-
laceae. U.S.A. (Virginia).

rostrupii P.A. Saccardo & D. Saccardo in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
17:721. 1905. =Leptosphaeria dryadis E.
Rostrup. Nec E. Rostrup, 1903. On upper
surface of dried leaves of Dryas octopetala L.,
Rosaceae. Norway.

rostrupii J. Lind, Annales Mycologici, Berlin 13:17.
1915. Nom. illegit. Art. 64.1. On stems of
Daucus carota L., Umbelliferae. Denmark,
Germany.

rothomagensis P.A. Saccardo, Michelia Commentar-
ium Mycologicum Fungos in Primis Italicos
Illustrans 2:63. 1880. On stems of Senecio
erucifolius L., Compositae. France.

rothomagensis P.A. Saccardo var. artemisiae N.
Patouillard, Exploration Scientifique de la
Tunisie, p. 13. 1892. On dead stems of
Artemisia herba-alba Asso, Compositae.
Tunisia.

roumegueri P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:62. 1880. =Metasphaeria
roumegueri P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:181. 1883. =Massariosphaeria
roumegueri (P.A. Saccardo) A Leuchtmann,
Sydowia. Annales Mycologici, Hom, N.O.
37:173. 1984. On decaying cortex of
Phormium sp., Agavaceae. France.

rousseliana (J. Desmaziéres) V. Cesati & G. de
Notaris, Commentario della Societa Crittoga-
mologica Italiana, Milan 1:236. 1863.
=Sphaeria (Foliicola) rousseliana J. Des-
mazieres, Annales des Sciences Naturelles,
Paris, Botanique, Series 3, 11:355. 1849.
=Phaeosphaeria rousseliana (J. Desmaziéres)
L. Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):114. 1957. On sheath of
Phleum boehmeri Wibel., Gramineae.
Canada, Czechoslovakia, Germany, Nether-
lands, Switzerland.

March 1991

rubelloides (C.B. Plowright) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:77. 1883.
=Sphaeria rubelloides C.B. Plowright in M.C.
Cooke, Grevillea, London 5:120. 1877. On
decaying culms of Triticum repens L.,
Gramineae. Great Britain.

rubellula (J. Desmaziéres) F. v. Héhnel, Osterreic-
hische Botanische Zeitschrift, Wien 63:377.

1913. =Sphaeria modesta J. Desmaziéres var.

rubellula J. Desmaziéres, Annales des
Sciences Naturelles, Paris, Botanique, Series
3, 16:312. 1851. On stems of Crepis
vesicaria L. subsp. haenseleri (Boiss. ex DC.)
P.D. Sell [as Barkhousia taraxacifolia
(Thuill.) DC.], Compositae. France.
rubicunda H. Rehm ex H. Rehm in H.G. Winter,
Ascomyceten, Fascicle 2, No. 92. Anno 1871.
Nom. nud. Art. 32.1. Flora, Jena und
Regensburg 55:540. 1872. =Melanomma
rubicundum (H. Rehm ex H. Rehm) L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):65. 1957. On fallen stems of unknown

host of Umbelliferae, Umbelliferae. Hungary.

rubicunda H. Rehm, Ascomycetes Lojkani Lecti in
Hungaria, Transylvania et Galicia, Berlin, pp.
53-54. 1882. Nom. illegit. Art. 64.1. On
decayed stems of Conium maculatum L.,
Umbelliferae. Hungary.

rubrotincta J.B. Ellis & B.M. Everhart, Journal of

Mycology, Columbus, Ohio 1:150—151. 1885.

On dead twigs of Staphylea trifolia L.,
Staphyleaceae. U.S.A.

rudbeckiae (L.A. Kirchner) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:17. 1883. =Ple-
ospora rudbeckia L.A. Kirchner in F. v.
Thiimen, Fungi Austriaci Exsiccati, Century
9, No. 858. 1874. On dried stems of
Rudbeckia amplexicaulis Vahl., Compositae.
Czechoslovakia.

rugosa J. Dearness & G.R. Bisby in G.R. Bisby, A.
Buller, and J. Dearness, The Fungi of
Manitoba, p. 74. 1929. On dead stems of
Cornus stolonifera Michx., Cornaceae.
Canada.

rulingiae F. Tassi, Bolletino del R. Orto Botanico,
Siena 3:95—96. 1900. On branchlets of
Rulingia prostrata Maiden & Betche,
Sterculiaceae. Australia.

rumicicola §. Ahmad, Monographs, Biological
Society of Pakistan, Lahore 5:13. 1969.
=Leptosphaeria rumicis A.C. Batista, W.A.
Cavalcanti & S.H. Iqbal. Nec Leptosphaeria
rumicis F, Fautrey. Nec Leptosphaeria
rumicis H. Wegelin. On stems of Rumex

nepalensis Sprengel, Polygonaceae. Pakistan.

rumicis A.C. Batista, W.A. Cavalcanti & S.H. Iqbal
in A.C. Batista, G.E.P. Peres, and S.H. Iqbal,
Atas do Instituto de Micologia, Recife

Nomenclator of Leptosphaeria 271

4:135-136. 1967. Nom. illegit. Art. 64.1. On
stems of Rumex nepalensis Sprengel,
Polygonaceae. Pakistan.

rumicis F. Fautrey in C. Roumeguére, Revue
Mycologique, Toulouse 13:168. 1891; Fungi
Selecti Gallici Exsiccati, Century 59, No.
5848. Anno 1891. On old stems of Rumex
patientia L., Polygonaceae. France.

rumicis H. Wegelin, Mitteilungen der Thurgauischen
Naturforschenden Gesellschaft, Frauenfeld
12:177. 1896. Nom. illegit. Art. 64.1.
=Leptosphaeria wegeliniana P.A. Saccardo &
P. Sydow. On dried stems of Rumex sp.,
Polygonaceae. Switzerland.

rusci (E.M. Fries) G. Niessl v. Mayendorf, Instituto.
Revista Scientifica e Litteraria, Coimbra
31:90. 1883. =Sphaeria rusci K.F.W.
Wallroth, Flora Cryptogamica Germaniae
2:776. 1833. =Leptosphaeria rusci (E.M.
Fries) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:74. 1883 [as (K.F.W. Wallroth)].

rusci (E.M. Fries) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:74. 1883. =Sphaeria atro-virens
E.M. Fries:E.M. Fries var. rusci E.M.
Fries:E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:501. 1823. =Sphaeria
rusci (E.M. Fries) K.F.W. Wallroth, Flora
Cryptogamica Germaniae 2:776. 1833.
=Sphaerella rusci (E.M. Fries) V. Cesati & G.
de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:237. 1863.
=Paraphaeosphaeria rusci (E.M. Fries) O.
Eriksson, Arkiv for Botanik, Uppsala,
Stockholm, Series 2, 6:406. 1967 [as
(Wallroth) O. Erikss.]. On cladodium and
stems of Ruscus aculeatus L., Ruscus
androgynus L., Ruscus hypoglossum L.,
Ruscaceae. Austria, Belgium, France, Great
Britain, Italy, Portugal.

rusci (E.M. Fries:E.M. Fries) P.A. Saccardo forma
caulina D. Saccardo, Mycotheca Italica,
Sistens Fungos Venetos Exsiccatos, Century
15, No. 1484. 1905. [Description in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:74. 1883 sub. Leptosphaeria rustici.| On
culms of Ruscus aculeatus L., Ruscaceae.
Italy.

rusci (E.M. Fries) P.A. Saccardo forma fourcroyae
P.A. Saccardo, Florula Mycologica Lusitanica
Sistens Contributionem Decimam ad Eamdem
Floram nec non Conspectum Fungorum
Omnium in Lusitania Hucusque Observa-
torum, Coimbra (Imprensa da Universidade),
p. 11. 1893. On dead leaves of Furcraea
bedinghausii C. Koch, Agavaceae. Portugal.

rusci (E.M. Fries) P.A. Saccardo var. hypophylli
R.C. Maire, Bulletin de la Société Botanique

de France, Paris 56:CCLXXII-CCLXXIIIL.
(1909) 1911. On dead cladodes of Ruscus
hypophyllum L., Ruscaceae. France.

rusci (E.M. Fries) G. Niessl v. Mayendorf var. rusci-
hypoglossi C. Massalongo, Malpighia.
Rassegna Mensuale di Botanica. Messina,
Genova 8:194. 1894; Nuovo Contribuzione
alla Micologia Veronese Genova (Tipografia
di Angelo Ciminago), p. 38. 1894. Nom. nud.
Art. 32.1. On leaves of Ruscus hypoglossum
L., Ruscaceae. Italy.

ruscicola P. Hariot & P.A. Karsten, Revue Mycolo-
gique, Toulouse 12:128—129. 1890. On
rotting stems of Ruscus sp., Ruscaceae.
France.

ruscicola P.A. Hariot & P.A. Karsten forma
cladodiicola R. Gonzalez Fragoso, Trabajos
del Museo Nacional de Ciencias Naturales,
Madrid, Series Botanica 10:90. 1916. On
dead cladodes of Ruscus aculeatus L.,
Ruscaceae. Spain.

russellii R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2711. 1984. On stems of
Asclepias sp.?, Asclepiadaceae. Canada.

rustica P.A. Karsten, Fungi Fenniae Exsiccati,
Century 10, No. 964. Anno 1870. =Metas-
phaeria rustica (P.A. Karsten) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:157.
1883. On stems of Spiraea ulmaria L.
(=Filipendula ulmaria Maxim.), Rosaceae.
Finland.

ruthenica F. Petrak, Annales Mycologici, Berlin
19:78-79. 1921. On dry stems of Sambucus
ebulus L., Caprifoliaceae. France.

sabalicola (J.B. Ellis & G.W. Martin) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo Addi-
tamenta Ad Volumina I-IV, 2:LVII-LVIII.
1882. =Sphaeria sabalicola J.B. Ellis & G.W.
Martin, American Midland Naturalist, Notre
Dame, Indiana 16:810. 1882. =Heptameria
sabalicola (J.B. Ellis & G.W. Martin) M.C.
Cooke, Grevillea, London 18:32. 1889. On
rachis of Sabal serrulata Roem & Schult.,
Palmae. U.S.A.

(Clypeosphaeria) sabaligera (M.J. Berkeley & M.A.
Curtis) M.C. Cooke, Grevillea, London 17:91.
1889. =Sphaeria sabaligera M.J. Berkeley &
M.A. Curtis in M.J. Berkeley, Grevillea,
London 4:147. 1876. =Clypeosphaeria sa-
baligera (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Diggesit P.A. Saccardo
2:91. 1883. On unknown substrate of Sahbal
sp., Palmae. U.S.A.

sabauda C.L. Spegazzini, Revue Mycologique,
Toulouse 4:79. 1882. On dead, decaying
leaves of Silene acaulis (L.) Jacg., Caryophyl-
laceae. France.

Illinois Natural History Survey Bulletin

sabauda C.L. Spagazzini forma arvaticae R.
Gonzalez Fragoso, Trabajos del Museo
Nacional de Ciencias Naturales, Madrid,
Series Botanica 4:17. 1914. On dead or rotting
leaves of Silene arvatica Lag., Caryophy!-
laceae. Spain.

sabuletorum (M.J. Berkeley & C.E. Broome) J.
Lind, Danish Fungi as Represented in the Her-
barium of E. Rostrup, Nordisk Forlag, Copen-
hagen, p. 219. 1913. =Metasphaeria sabule-
torum (M.J. Berkeley & C.E. Broome) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:180. 1883. =Sphaeria (Foliicolae) sabule-
torum M.J. Berkeley & C.E. Broome, Annals
and Magazine of Natural History, London, Se-
ries 2, 9:382. 1852. On dead leaves of
Festuca arundinacea Schreber, Gramineae.
Denmark, Great Britain.

sabuletorum (M.J. Berkeley & C.E. Broome) F. v.
Hohnel, Hedwigia, Dresden 60:141. 1918.
=Sphaeria (Foliicolae) sabuletorum M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 9:382. 1852. On dead leaves of Ammophila
arundinacea Host, Gramineae. Great Britain.

saccardiana J.H. Fabre, Annales des Sciences
Naturelles, Paris. Botanique, Series 6, 9:89.
1878. =Requienella saccardiana (J.H. Fabre)
J.H. Fabre, Annales des Sciences Naturelles,
Paris, Botanique, Series 6, 15:56. 1883. On
dried stems of Crataegus oxyacangha L.,
Rosaceae. France.

sacchari J. Van Breda De Haan, Mededelingen Van
Het Proefstation Voor Suikerriet in West-
Java, ‘Kagok’. Soerabaia 1892:25—28. 1892.
=Leptosphaerella sacchari (J. Van Breda De

Haan) S.C. Teng, Sinensia. Special Bulletin of

the Metropolitan Museum of Natural History,
Nanking 9:224. 1938. =Phaeosphaeria
sacchari (J. Van Breda De Haan) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1535. 1989. On sheaths and leaves
of Saccharum officinarum L., Gramineae.
Java.

sacchari C.L. Spegazzini, Revista. Universidad
Nacional de La Plata 2(18):232. 1896:
Hongos De La Cana De Azucar No. 9. Anno
1896. Nom. illegit. Art. 64.1. =Lepto-
sphaeria spegazzinii P.A. Saccardo & P.
Sydow. On wilted sheaths and leaves of
Saccharum officinarum L., Gramineae.
Argentina.

saccharicola P.C. Hennings, Hedwigia, Dresden
39:79. 1900. On leaves of Saccharum
officinarum L., Gramineae. Brazil.

sacculus G, Passerini & V. Beltrani, Atti dell’
Accademia Nazionale dei Lincei. Transunti,
Rome, Series 3, 7:36. 1883. =Metasphaeria
sacculus (G. Passerini & V. Beltrani) P.A.

Vol. 34 Art. 3

March 1991

Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:162. 1882. On dry stem of Euphorbia
dendroides L., Euphorbiaceae. Italy (Sicily).

saginata H. Sydow, Annales Mycologici, Berlin
37:377-378. 1939. =Mixtura saginata (H.
Sydow) O. Eriksson & J. Yue, Mycotaxon.
An International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens, Ithaca,
New York 38:203. 1990. On living or wilted
leaves of Chusquea serrulata Pilger, Gramin-
eae. Ecuador.

salebricola E.C. Bommer, M.H. Rousseau & P.A.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:783. 1891. =Phaeosphaeria
salebricola (E.C. Bommer, M.H. Rousseau &
P.A. Saccardo) A. Leuchtmann, Sydowia.
Annales Mycologici, Horn, N.O. 37:156.
1984. On drying leaves of Srellaria graminea
L., Caryophyllaceae. Belgium, Germany.

salebrosa P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:599. 1882. Nec C.G.T. Preuss.
=Leptosphaeria sapeyensis P.A. Saccardo.
On stems of Gentiana lutea L., Gentianaceae.
France.

salebrosa (C.G.T. Preuss) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:20. 1883. Nom.
illegit. Art. 64.1. =Phaeoderris salebrosa
(C.G.T. Preuss) F. v. H6hnel, Sitzungsber-
ichte der Akademie der Wissenschaften in
Wien, Mathematisch-naturwissenschaftliche
Klasse, Abt. I, 120:462. 1911. =Sphaeria
salebrosa C.G.T. Preuss, Linnaea. Institut
botanique de l'Université de Genéve 26:714.
1853. On trunks of Brassica crispa Rafin.,
Cruciferae. Germany.

salicaria G. Passerini, Revue Mycologique,
Toulouse 2:35. 1880. On dry stems of
Lythrum salicaria L., Lythraceae. Italy.

salicinearum (G. Passerini) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:50. 1883.
=Sphaerella salicinearum G. Passerini. Nom.
nud. Art. 32.1 (herbarium name). =Sphae-
rella salicinearum G. Passerini forma populi-
nigrae G.L. Rabenhorst, Fungi Europaei
Exsiccati, Klotzschii Herbarii vivi Mycologici
Continuatio III, Century 25, No. 2444. Anno
1876; Hedwigia, Dresden 17:172. 1878.
=Sphaerella salicinearum G. Passerini forma
salicis-albae G.L. Rabenhorst, Fungi
Europaei III, Century 25, No. 2445. Anno
1876. On leaves of Populus nigra L., Salix
alba L., Salicaceae. Italy.

salsolae L. Hollés, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest 4:335.

Nomenclator of Lepfosphaeria

273

1906. On dry stems of Salsola kali L.,
Chenopodiaceae. Hungary.

salviae G. Passerini, Atti dell’ Reale Accademia
Nazionale dei Lincei. Rendiconti, Rome,
Series 4, 4(2):59. 1888. On dry branches of
Salvia officinalis L., Labiatae. Italy.

salviae G. Passerini forma minor C. Sandu-Ville,
Academia Republicii Populare Romine.
Filiala lasi. Studii si Cercetari Stiintifice.
Biologie si Stiinte Agricole 13:257. 1962. On
unknown substrate of Lavandula vera DC.,
Labiatae. Romania.

salvinii A. Cattaneo, Archivo del Laboratorio di
Botanica Crittogamica Universita di Pavia 2/
3:126. 1879. =Magnaporthe salvinii (A.
Cattaneo) R.A. Krause & R.K. Webster,
Mycologia, Lancaster, Pennsylvania 64:110.
1972. On leaf sheaths of Oryza sativa L.,
Gramineae. Italy.

sambuci F. Fautrey in C. Roumeguere, Revue
Mycologique, Toulouse 14:7. 1892; Fungi
Selecti Gallici Exsiccati, Century 60, No.
5960. Anno 1892. Nom. illegit. Art. 64.1. On
dry branches of Sambucus nigra L., Caprifoli-
aceae. France.

sambuci P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):152. 1873. On branches below the
epidermis of Sambucus nigra L., Caprifoli-
aceae. Italy.

sambucina J.B. Ellis & B.M. Everhart, Proceedings
of the Academy of Natural Sciences of
Philadelphia 1894:336—337. 1894. =Chaeto-
mastia sambucina (J.B. Ellis & B.M.
Everhart) M.E. Barr, Mycotaxon. An
International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens, Ithaca,
New York 34:513. 1989. On dead plant of
Sambucus melanocarpa A. Gray, Caprifoli-
aceae. U.S.A.

sanguisorbae P.A. Karsten in A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:78. 1892. On branches of Sanguisorba
officinalis L., Rosaceae. Germany.

sanguisorbae P.A. Karsten, Enumeratio Fungorum et
Myxomycetum in Lapponia Orientali Aestate
1861 Lectorum, p. 214. 1886; Notiser
Sallskapets Pro Fauna et Flora Fordhandlingar
8:214. 1866 [as Sphaeria (Leptosphaeria)
sanguisorbae|. On dead stems of San-
guisorba polygama F. Nyl., Rosaceae.
U.S.S.R.

sapeyensis P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:38. 1883. Nom. illegit. Art. 63.1.
=Leptosphaeria salebrosa P.A. Saccardo;
Michelia Commentarium Mycologicum
Fungos in Primis Italicos Illustrans 2:599.
1882. Nec C.G.T. Preuss. On stems of
Gentiana lutea L., Gentianaceae. France.

274 Illinois Natural History Survey Bulletin

saprophila H. Rehm, Hedwigia, Dresden 21:121.
1882. =Metasphaeria saprophila (H. Rehm)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:172. 1883. On decaying berries of Junipe-
rus nana Willd., Cupressaceae. Austria.

sarmenticia P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:63—64. 1880. On runners of
Solanum dulcamara L., Solanaceae. France.

sarothamni J.B. Lambotte & F. Fautrey in C.
Roumegueére, Revue Mycologique, Toulouse
15:115. 1893; in C. Roumeguere, Fungi
Selecti Gallici Exsiccati, Century 64, No.
6356. Anno 1893. On branches of Sarotham-
nus scoparius (L.) Wimmer ex Koch,
Leguminosae. France.

sarraziniana P.A. Saccardo & C. Roumeguere,
Revue Mycologique, Toulouse 7:159. 1885.
On nodes of unknown host, Caryophyllaceae.
France.

sasacola K. Hara, ByOchu-gai Zasschi (Journal of
Plant Protection), Tokyo 5:885. 1918. On
unknown substrate of Sasa sp., Gramineae.
Japan.

sasae K. Hara, Byéchu-gai Zasschi (Journal of Plant
Protection), Tokyo 5:884—885. 1918. On
dead leaves of Sasa sp., Gramineae. Japan.

saxonica F. vy. Hohnel, Annales Mycologici, Berlin
16:84-86. 1918. =Phaeosphaeria saxonica
(F. v. Hohnel) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1564. 1989. On
dead culms of Scirpus maritimus L., Cyper-
aceae. Germany.

scabiens (V. Cesati) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:69. 1883.
=Sphaeria scabiens V. Cesati, Atti dell’
Accademia delle Scienze Fisiche e Matemati-
che, Napoli 8(2):20. 1879. On culms of
unknown host, unknown family. Malaysia.

scabrispora S.C. Teng, Sinensia. Special Bulletin of
the Metropolitan Museum of Natural History,
Nanking 4:378. 1934. =Massariosphaeria
scabrispora (S.C. Teng) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1589.
1989. On culms of Bambusa sp., Gramineae.
China (Kiangsu).

scapophila (C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:43-44. 1883.
=Sphaeria (Caulicolae) scapophila C.H.
Peck, Report. New York State Museum of
Natural History, Albany 30:66—67. 1878. On
dead scapes of Sarracenia purpurea L.,
Sarraceniaceae. U.S.A.

schneideriana (J. Rick) J. Rick in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 22:231.
1913. =Trematosphaeria schneideriana J.

Vol. 34 Ar. 3

Rick, Broteria, Lisboa 5:47. 1906. On
unknown substrate of Bambusa sp., Gramin-
eae. Brazil.

schoenocauli H. Sydow, Annales Mycologici, Berlin
28:84-85. 1930. On dead decaying stems of
Schoenocaulon officinale A. Gray, Liliaceae.
Portugal.

scirpina H.G. Winter, Hedwigia, Dresden 11:146.
1872. =Metasphaeria scirpina (H.G. Winter)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:182. 1882. =Trichometasphaeria scirpina
(H.G. Winter) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):142. 1957.
=Massariosphaeria scirpina (H.G. Winter) A.
Leuchtmann, Sydowia. Annales Mycologici,
Horn, N.O. 37:174. 1984. On dry culms of
Scirpus lacustris L., Cyperaceae. Austria
(Tyrol?), Italy.

scitula H. Sydow, Annales Mycologici, Berlin
36:32 1-332. 1938. =Melanomma scitulum
(H. Sydow) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):66. 1957. On
rotten stems of Galium aparine L., Rubiaceae.
Germany.

sclavonica (S. Schulzer v. Miiggenburg & P.A.
Saccardo) F.A. Hazslinszky, Matematikai és
Természettudomanyi K6zlemenyek Von-
atkozolag a Hazai Viszonyokra, Budapest
25(2):150. 1892. =Metasphaeria sclavonca S.
Schulzer v. Miiggenburg & P.A. Saccardo,
Revue Mycologique, Toulouse 6:69—70. 1884.
On decorticated branches of Vitis vinifera L.
subsp. sylvestris (C.C. Gmelin) Hegi,
Vitaceae. Yugoslavia.

scolecosporarum M. de Sousa da Camara, Revista
Agronomica, Lisbon 20(1):23. 1932. On
leaves of Quercus suber L., Fagaceae.
Portugal.

scotophila P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:317. 1875. On stems of
Aconitum sp., Cynanchum sp., Vincetoxicum
sp., Ranunculaceae, Asclepiadaceae. Italy.

scotophila P.A. Saccardo forma aconti P.A.
Saccardo [falsely so cited in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:157.
1883. As published in Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:317. 1875: Fungi
Veneti Novi vel Critici, Series 2, p. 317.
1875.] =Leptosphaeria scotophila P.A.
Saccardo.

scrophulariae (J. Desmaziéres) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:57.
1883. =Sphaeria scrophulariae J. Des-
maziéres, Plantes Cryptogames Du Nord De
La France, Edition 1, Fascicle 15, No. 718.

March 1991

Anno 1834. Non vidi; Annales des Sciences
Naturelles, Paris, Botanique, Series 2,
6:245-246. 1836. On capsules of Scrophu-
laria aquatica L., Scrophulariaceae. France.

scutati S.A. Gucevicz, Novosti Sistematiki Nizshikh
Rastenij, Novitates Systematicae Plantarum
Non Vascularium 6:129-130. 1969 [as
scutata|. On dried stems of Rumex scutatus
L., Polygonaceae. U.S.S.R. (Ukraine).

secalina A. Jaczewski, Opredielitel’ Gribov’ 2,
Fungi Imperfecti (Petrograd), pp. 606—607.
1917. On unknown substrate of Secale
cereale L., Gramineae. U.S.S.R.

secalina Lachevskii in N.N. Lavrov, Trudy
Tomskogo Gosudarstvennogo Universiteta.
Tomsk. Series Biologicheskii 104:88. 1948.
Non vidi. Nom. illegit. Art. 64.1. On dead
culms of Secale cereale L., Gramineae.
U.S.S.R.

secalis F. Haberlandt in J. Schroter, Just’s Botanis-
che Jahresberichte, Berlin 6(2):319. 1878. On
unknown substrate of Secale cereale L.,
Gramineae. Unknown country.

semelina A. Caballero, Anales del Jardin Botanico
de Madrid 1:200. 1941. On living cladode of
Semele androgyna (L.) Kunth, Liliaceae.
Spain.

seminuda (C.H. Persoon:E.M. Fries) V. Cesati & G.
de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:235. 1863.

=Sphaeria seminuda C.H. Persoon:E.M. Fries,

Synopsis Methodica Fungorum, pp. 70-71.
1801; E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:464. 1823. =Tremato-
sphaeria seminuda (C.H. Persoon:E.M. Fries)
L. Fuckel, Symbolae Mycologicae Beitrage
zur Kenntnis Der Rheinischen Pilze, p. 162.
1870. On dry wood of Fagus sp., Fagaceae.
Austria, Germany.

senecionis (L. Fuckel) H.G. Winter, Dr. L.
Rabenhorst’s Kryptogamen-Flora von
Deutschland, Oesterreich und der Schweiz
1(2):469. 1867. =Pleospora senecionis L.
Fuckel, Fungi Rhenani Exsiccati A Leopoldo
Fuckel Collecti Supplement, Fascicle 6, No.
2032. Anno 1867. =Metasphaeria senecionis
(L. Fuckel) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:158. 1883. On dry stems of
Senecio nemorensis L., Compositae. Austria.

senegalensis G. Segretin, J. Baylet, H. Darasse & R.
Camain, Comptes Rendus hebdomadaire des
Sciences de |’ Académie des Sciences, Paris,
248:3732. 1959. On Homo sapiens, Hom-
inidae. Senegal.

sepalorum (J.S. Vleugel) J. Lind, Svensk Botanisk
Tidskrift, Stockholm 22:66. 1928. =Metas-
phaeria sepalorum J.S. Vieugel, Svensk
Botanisk Tidskrift, Stockholm 2:369-370.
1908. =Bricookea sepalorum (J.S. Vieugel)

Nomenclator of Leptosphaeria

M.E. Barr, Mycotaxon. An International
Journal Designed to Expedite Publication of
Research on Taxonomy & Nomenclature of
Fungi & Lichens, Ithaca, New York 15:346—
347. 1982. On dead sepals of Juncus
filiformis L., Juncus trifidus L., Juncaceae.
Norway (Spitsbergen).

sepincola (E.M. Fries:E.M. Fries) H.G. Winter, Dr.
L. Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):473. 1885. =Sphaeria
sepincola E.M. Fries, Observationes Mycol-
ogicae. Kjobenhavn 1:181. 1815; Systema
Mycologicum Sistens Fungorum 2:498. 1823
[as saepincola]. =Saccothecium sepincola
(E.M. Fries:E.M. Fries) E.M. Fries, Summa
Vegetabilium Scandinaviae, Seu Enumeratio,
Systematica et Critica, Plantarum tum
Cotyledonearum, Quam Nemearum Inter
Mare Occidentale Et Album, Inter Eidoram et
Nordkop, Hactenus Lectorum, una Cum
Singulae Distributione Geographica, p. 398.
1849. =Pringsheimia sepincola (E.M.
Fries:E.M. Fries) F. v. HOhnel, Annales
Mycologici, Berlin 18:97. 1920. =Metas-
phaeria sepincola (E.M. Fries:E.M. Fries)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:164. 1883 [as Fr? Fuckel]. =Sphaerulina
sepincola (E.M. Fries:E.M. Fries) K.
Starbick, Botaniska Notiser, Lund 1890:117.
1890; Botanisches Zentralblatt, Jena &
Dresden 46:261. 1891. =Pleosphaerulina
sepincola (E.M. Fries:E.M. Fries) H. Rehm in
F. y. Héhnel, Annales Mycologici, Berlin
18:96. 1920. =Sclerodothis sepincola (E.M.
Fries:E.M. Fries) F. Petrak, Annales Mycol-
ogici, Berlin 19:41. 1921. On dry stems of
Rosa sp., Rubus sp., Rosaceae. Sweden.

sepincola (M.J. Berkeley & C.E. Broome) F.A.
Hazslinszky, Matematikai és Természettu-
domanyi Kézlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 25(2):151. 1892.
Nom. illegit. Art. 64.1. =Sphaeria sepincola
M.J. Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 9:326. 1852. =Metasphaeria sepincola
(M.J. Berkeley & C.E. Broome) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:164. 1883. [Pro parte fide F.A.
Hazslinszky.] On herbaceous stems of Rumex
sp., Polygonaceae. Great Britain.

septemcellulata E. Miiller, Sydowia. Annales
Mycologici, Horn, N.O. 4(1-6):300. 1950.
=Nodulosphaeria septemcellulata (E. Miiller)
L. Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):86. 1957. On dry, dead stems
of Buphthalmum salicifolium L. (=Anthemis),
Compositae. Switzerland.

276

septovariata A.M. Saccas, Agronomie Tropicale,
Nogent-sur-Marne 9:271. 1954. On aerial
roots of Sorghum vulgare Pers., Gramineae.
French Equatorial Africa.

sequana A.L. Guyot, Revue de Mycologie, Paris
11:68-69. 1946. =Phaeosphaeria sequana
(A.L. Guyot) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1573. 1989. On
dry seeds of Phleum boehmeri Wibel.,
Gramineae. France.

serbica J. Schroter, Hedwigia, Dresden 29:59-60.
1890. On stems of Camphorosma monspeli-
aca L., Chenopodiaceae. Yugoslavia.

seriata G, Passerini, Rendiconti della Sedute della R.
Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:91. 1887. On wilted leaves
of Zea mays L., Gramineae. Italy.

setosa G. Niessl v. Mayendorf, Verhandlungen des
Naturforschenden Vereins in Briinn 10:178.
(1871) 1872. On dried stems of Angelica sp.,
Pastinaca sp., Umbelliferae. Czechoslovakia
(near Gratz and Briinn).

setosa (A. Leuchtmann) R.A. Shoemaker, Canadian
Journal of Botany, Ottawa 67:1580. 1989.
Nom. illegit. Art. 64.1. =Phaeosphaeria
setosa A. Leuchtmann, Sydowia. Annales
Mycologici, Horn, N.O. 37:159-160. 1984.
On Yucca elephantipes Hort. ex Regel,
Agavaceae. Mexico.

(Hendersonia?) setulosa P.A. Saccardo & C.
Roumeguere in C. Roumegueére and P.A.
Saccardo, Revue Mycologique, Toulouse
3:44. 1881; Reliquiae Mycologicae Liber-
tianae, Series Altera Reviserunt C.
Roumeguére & P.A. Saccardo, Toulouse, No.
123. 1881. =Pocosphaeria setulosa (P.A.
Saccardo & C. Roumeguére) A.N. Berlese,
Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae
Adcommodatae 1:89. 1892. On decayed
culms of Secale sp., Gramineae. Germany.

shahvarica F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 3(1-6):279. 1949. On dry stems
of Cousinia multiloba DC., Compositae. Iran.

Shastensis R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2712. 1984. On stems of
Lupinus albicaulis Dougl. ex Hook. var
shastensis C.P. Sm., Leguminosae. U.S.A.

sibirica F. v. Thiimen, I. Moskovskoe Obshchestvo
Ispytatelei Prirody, Bulletin 56(2):124. 1882.
On dead stems of Crepis sibirica L., Com-
positae. U.S.S.R.

sibtorpii J. Politis, Akademia Athénon, Pragmateiai,
Athens 1935:16. 1935. On dry stems of
Euphorbia sibthorpii Boiss., Euphorbiaceae.
Greece.

sicula P.A. Saccardo & V. Beltrani, Michelia
Commentarium Mycologicum Fungos in

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Primis Italicos Illustrans 1:397. 1878. On
decayed trunks of Opuntia ficus-indica (L.)
Miller, Cactaceae. Italy.

sieversiae (F.E. Clements) F. Petrak, Sydowia.
Annales Mycologici, Horn, N.O. 6(1-4):6.
1952. =Massaria sieversiae F.E. Clements,
Cryptogamae Formationum Coloradensium,
Century III, No. 234. Anno 1906. Nom. nud.
Art. 36.1. [Combination is not valid because
the basionym is an invalidly published
species.] On standing, dead stems of
Sieversia turbinata (Rydb.) Greene, Rosaceae.
U.S.A.

silenes-acaulis G. de Notaris, Commentario della
Societa Crittogamologica Italiana, Milan,
2:485-486. 1867. =Scleropleella silenes-
acaulis (G. de Notaris) F. v. Hohnel, Annales
Mycologici, Berlin 18:76. 1920.
=Phaeosphaeria silenes-acaulis (G. de
Notaris) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):128. 1957. On
withering leaves of Silene acaulis (L.) Jacq.,
Caryophyllaceae. Italy.

sileris G. Bresadola, Studi Trentini Di Scienze
Naturall, Trento, Series 2, 7:66-67. 1926. On
stems of Laserpitium siler L., Umbelliferae.
Italy.

silvestris J. Feltgen, Vorstudien zu einer Pilz-Flora
des Grossherzogthums, Luxemburg 1(2):162.
1901. On dry stems of Silene inflata Sm.,
Caryophyllaceae. Luxembourg.

simillima H. Rehm, Leaflet of Philippine Botany,
Manila 6:2203—2204. 1914. On dead
branches of Capparis horrida L., Cappar-
aceae. Philippines.

simmonsii P.A. Saccardo, Nuovo Giomale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze, New Series 27:77. 1920. On
dead stems of Heracleum lanatum Michx.,
Compositae (Umbelliferae?). U.S.A.

sinapis L. Hollos, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:34. 1926. On dry
stems of Sinapis arvensis L., Cruciferae.
Hungary.

slovacica R. Picbauer, Sbornik Vysoke Skoly
Zemédélské a Lesnické Faculty v. Bee
(Bulletin de L’Ecolo Superieure
D’Agronomie Bro, RCS, Faculte de
Silviculture) 1927(D7):7—8. 1927. On stems
of Ballota nigra L., Labiatae. Czecho-
slovakia.

smarodsti G. Moesz, Magyar Botanikai Lapok,
Budapest 31:37. 1932. On wilted leaves of
Convallaria majalis L., Liliaceae. U.S.S.R.
(Latvia).

smilacis T, Petch, Ceylon Journal of Science,
Colombo, Section A (Botany) 6:225. 1917.
Nom. illegit. Art. 64.1. On branches of
Smilax aspera L., Smilacaceae. Sri Lanka.

March 1991

smilacis C.E. Chardon, Farlowia. A Journal of
Cryptogamic Botany, Cambridge, Massachu-
setts 2:466-467. 1946. Nom. illegit. Art.
64.1. On dry branches of (on leaves of?)
Smilax sp., Smilacaceae. Dominican
Republic.

smilacis (J.L. Castagne) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:69. 1883.
=Sphaeria smilacis J.L. Castagne, Catalogue
Des Plantes qui Croissent Naturellement aux
Environs de Marseille, p. 169. 1845. On dry
branches of Smilax sp., Smilacaceae. France.

socia P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:533. 1882. =Metasphaeria socia
(P.A. Saccardo) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:165. 1883. On
living vines of Vitis vinifera L., Vitaceae.
Italy.

socialis A.N. Berlese, cones Fungorum Omnium
Hucusque Cognitorum ad usum Sylloges
Saccardianae Adcommodatae 1:63. 1890. On
stems of Asparagus officinalis L., Liliaceae.
Finland.

sodomaea (G. de Notaris) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:20. 1883.
=Sphaeria sodomaea G. de Notaris, Commen-
tario della Societa Crittogamologica Italiana,
Milan 2:485. 1867. On dried stems of
Solanum sodomeum L., Solanaceae. Italy.

solani L.G. Romell in A.N. Berlese, [cones Fun-
gorum Omnium Hucusque Cognitorum ad
usum Sylloges Saccardianae Adcommodatae
1:63. 1890. =Leptosphaeria solanicola P.A.
Saccardo. Non Leptosphaeria solani J.B.
Ellis & B.M. Everhart. 1893. =Melanomma
solani (L. Romell ex A.N. Berlese) L. Holm,
Symbolae Botanicae Upsalienses, Uppsala
14(3):56. 1957. On stems of Solanum
dulcamara L., Solanaceae. Sweden.

solani J.B. Ellis & B.M. Everhart, Proceedings of the
Academy of Natural Sciences of Philadelphia
1893:136. 1893. Nom. illegit. Art. 64.1. On
living leaves of Solanum dulcamara L.,
Solanaceae. Canada.

solanicola P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 24:99]. 1928. =Leptosphaeria
solani J.B. Ellis & B.M. Everhart. Non
Leptosphaeria solani L.G. Romell in A.N.
Berlese. On stems of Solanum dulcamara L.,
Solanaceae. Sweden.

solheimii F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 18(1-6):369. 1964. On dry culms
of Juncus drummondii E. Mey., Juncaceae.
U.S.A.

solidaginis (E.M. Fries:E.M. Fries) V. Cesati & G.
de Notaris, Commentario della Societa

Nomenclator of Leptosphaeria 277

Crittogamologica Italiana, Milan 1:236. 1863.
=Sphaeria solidaginis E.M. Fries:E.M. Fries,
Elenchus Fungorum Sistens Commentarium
in Systema Mycologicum 2:106. 1828. On
stems of Solidago sp., Compositae. France.

sophorae E.N. Koschkelova & I.P. Frolov,
Mikoflora Podgornoi Raviniy Kopetdaga 1
Tsentral’nykh Karakumoy [Mycoflora of the
Mountainous Plains of Kopet Dagh and
Central Kara Kum], p. 76. 1973. Nom. inval.
Art. 34.1, 36.1. On unknown substrate of
Sophora jabonica L., Leguminosae. U.S.S.R.

sorbi A. Jaczewski, Annales Mycologici, Berlin
1:30. 1903. On leaves of Sorbus aucuparia
L., Rosaceae. U.S.S.R.

sorgho-arundinacei M. Luc, Revue de Mycologie
22, Paris (Supplement Colonial No. 1)
18:15-18. 1953. =Phaeosphaeria sorgho-
arundinacei (M. Luc) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1535.
1989. On leaves of Sorghum arundinaceum
Roem & Schult., Gramineae. Canada, Ivory
Coast, U.S.A.

sorghophila (C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:60. 1883.
=Sphaeria (Caulicolae) sorghophila C.H.
Peck, Report. New York State Museum of
Natural History, Albany 31:51. 1879;
Hedwigia, Dresden 20:157. 1881. On brush
of an old broom. U.S.A.

sowerbyi (L. Fuckel) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:78. 1883. =Lepto-
sphaeria maculans (M.J. Sowerby) P.A.
Karsten, Mycologia Fennica 2:99. 1873. Non
Leptosphaeria maculans (J. Desmaziéres) V.
Cesati & G. de Notaris, Commentario della
Societa Crittogamologica Italiana, Milan
1:235. 1863. =Sphaeria maculans M.J.
Sowerby, Coloured Figures of English Fungi
or Mushrooms 3:Table 394, Figure 9. 1803; in
M.J. Berkeley and C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 9:378. 1852. =Sphaerella maculans (M.J.
Sowerby) B. Auerswald in W. Gonnermann
and G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Saimmtlicher Pilze Europa’s,
Hefte 5S und 6, Synopsis Pyrenomycetum
Europaeorum, p. 18. 1869. =Pleospora
sowerbyi L. Fuckel, Symbolae Mycologicae,
Erster Nachtrag, p. 301. 1871. Non Sphaeria
maculanas J. Desmazieres, Annales des
Sciences Naturelles, Paris, Botanique, Series
3, 6:77-78. 1846. =Phaeosphaeria sowerbyi
(L. Fuckel) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):130. 1957. On
culms of Heleocharis palustris (L.) Roem. &
Schult. (=Eleocharis), Schoenoplectus
lacustris (L.) Palla, Scirpus lacustris L.,

278

Cyperaceae. Belgium, Canada, Poland,
Switzerland.

sparganii (F. Fautrey) A. Munk, Dansk Botanisk
Arkiv, Kj@benhavn 17(1):376. 1957. Nom.
illegit. Art. 33.2. =Metasphaeria sparganii F.
Fautrey, Revue Mycologique, Toulouse
13:76-77. 1891. [Basionym not completely
cited as to place of publication and date.] On
dead stems of Sparganium erectum L.,
Sparganium ramosum Hudson, Spargani-
aceae. Ghana.

sparsa (L. Fuckel) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:77. 1883. =Pleospora sparsa L.
Fuckel, Symbolae Mycologicae, p. 138. 1870.
=Phaeosphaeria sparsa (L. Fuckel) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1556. 1989. On culms and leaves
of Agrostis sp., Calamagrostis sp., Poa sp.,
Gramineae. Switzerland.

sparsa (L. Fuckel) P.A. Saccardo var. elynae A.
Ade, Allgemeine Botanische Zeitschrift f.
Systematik, Floristik, Pflanzengeographie,
etc., Karlsruhe 30/31:22(134). 1926. On
unknown substrate of Elyna spicata Schrader,
Cyperaceae. Austria.

sparsa (L. Fuckel) P.A. Saccardo var. meizospora J.
Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):207.
1903. On dry leaves of Carex acutiformis
Ehrh., Sparganium ramosum Hudson,
Cyperaceae, Sparganiaceae. Luxembourg.

spartii F. Tassi, Bolletino del R. Orto Botanico,
Siena 7:60—61. 1905. On culms and sheaths
of Lygeum spartum L., Gramineae. Italy.

spartinae J.B. Ellis & B.M. Everhart, Journal of
Mycology, Columbus, Ohio 1:43. 1885.
=Phaeosphaeria spartinae (J.B. Ellis & B.M.
Everhart) R.A. Shoemaker, Canadian Journal
of Botany, Ottawa 67:1573. 1989. On dead
culms of Spartina sp., Gramineae. U.S.A.

spatharum V. Cesati in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotzschii Herbarii vivi
Mycologici Continuatio Edita Nova, Series 2,
Century 26, No. 2530. Anno 1881. Nom. nud.
Art. 32.1. =Metasphaeria spartharum (V.
Cesati) ex P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:179. 1883. On spath of Chamaer-
ops excelsior Boj, Palmae. Italy.

spectabilis G. Niessl v. Mayendorf, Verhandlungen
des Naturforschenden Vereins in Brinn
10:179-180. (1871) 1872. On dried stems of
Laserpitium sp., Umbelliferae. Austria (near
Vienna).

spegazzini P.A. Saccardo & P. Sydow, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 14:570. 1899.
=Leptosphaeria sacchari C.L. Spegazzini.
Nec Leptosphaeria sacchari H.J. Van Breda.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

On wilting or drying leaves and sheaths of
Saccharum officinarum L., Gramineae.
Argentina.

spegazzini P.A. Saccardo & P. Sydow var. minor
C.L. Spegazzini, Anales del Museo Nacional
de Historia Natural de Buenos Aires 19(Series
3, 12):383. 1909; Mycetes Argentinenses,
Series 4, p. 383, No. 629. 1909. On culms of
Saccharum officinarum L., Gramineae.
Argentina.

sphyridiana (J.G. Lahm) H.G. Winter in Dr. L.
Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz
1(2):442-443. 1887. (As Leptosphaeria?).
=Xenosphaeria sphyridiana J.G. Lahm in F.
Arnold, Flora, Jena und Regensburg 57:150.
1874. On thallus of Sphyridium fungiforme
Flotow [=Baeomyces rufus (Huds.) Rebent.],
Lichenes (Baeomycetaceae). Austria,
Germany.

spiraeae P.A. Karsten, Hedwigia, Dresden 27:260.
1888. On dry fallen branches of Spiraea
sorbifolia L. (=Sorbaria sorbifolia A. Br.),
Rosaceae. Finland.

sporoboli J.B. Ellis & B.T. Galloway, Journal of
Mycology, Columbus, Ohio 5:66. 1889. On
dead culms of Sporobolus depauperatus Torr.
ex Hemsl. [=Muehlenbergia richardsonis
(Trin.) Rydb.], Gramineae. U.S.A.

squamata (M.C. Cooke & J.B. Ellis) J.B. Ellis,
Catalogue of Plants Found in New Jersey.
Geological Survey of New Jersey, Final
Report of the State Geologist, Trenton
2(1):525. 1889. =Sphaeria (Obtectae)
squamata M.C. Cooke & J.B. Ellis, Grevillea,
London 7:10. 1878. On bark of dead pine
limbs of Pinus sp., Pinaceae. U.S.A. (New
Jersey).

Staritzii P.C. Hennings in R. Staritz, Verhandlungen
des Botanischen Vereins Der Provinz
Brandenburg, Berlin 55:78. 1913. On dead
stems of Armeria vulgaris Willd., Plumbag-
inaceae. Germany.

Staritzii R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2713. 1984. Nom. illegit.
Art. 64.1. On stems of Achillea millefolium
auct., Compositae. Germany.

staticicola L.M. Unamuno Yrigoyen, Asociacion
Espanola Para el Progresso de las Ciencias,
Congreso de Coimbra 1923:40. 1923. On dry
leaves of Statice occidentalis Lloyd (=Arm-
eria), Plumbaginaceae. Spain.

steinti (G.W. Koerber) ex H.G. Winter, Dr. L.
Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):443. 1885. (As
Leptosphaeria?). =Leptorhaphis steinii G.W.
Koerber, Jahresbericht der Schlesischen
Gesellschaft fiir vaterlandische Kultur,
Breslau 46:91. 1869. Nom. nud. Art. 32.1.
=Ophiobolus steinii (G.W. Koerber) A.N.

March 1991

Berlese & P. Voglino, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo Additamenta Ad Volumina I-IV, p.
189. 1886. On thallus of Lecanora frustulosa
(Dicks.) Ach., Lichenes (Lecanoraceae).
Czechoslovakia, Germany.

steironematis J.B. Ellis & B.M. Everhart, Proceed-
ings of the Academy of Natural Sciences of
Philadelphia 1890:237. (1890) 1891. On dead
stems of Steironema ciliatum (L.) Rafin.
(=Lysimachia ciliata L.), Primulaceae.
Canada.

stellariae C.E. Richon, Catalogue Raisonné Des
Champignons Qui Croissent Dans le Départe-
ment de la Marne, p. 299. 1889. Nom. illegit.
Art. 64.1. =Leptosphaeria richoni P.A.
Saccardo & P. Sydow, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 14:571. 1899. On old stems of
Stellaria media (L.) Vill., Caryophyllaceae.
France.

stellariae E. Rostrup, Meddelelser om Gronland,
Kj@benhavn 3:557. 1888. =Phaeosphaeria
stellariae (E. Rostrup) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:157. 1984. On leaves of Stellaria
humifusa Rottb., Caryophyllaceae. Green-
land.

stellata N.T. Patouillard in N.T. Patouillard and N.G.
de Lagerheim, Bulletin de la Société Mycolo-
gique de France, Paris 11:225. 1895. On dead
stems of Chusquea sp., Gramineae.
Colombia.

stereicola J.B. Ellis, American Naturalist, Lancaster,
Pennsylvania 17:317. 1883. On hymenium of
Stereum bicolor (Pers.) Pers. [=Laxitextum
bicolor (Pers.), Lentz], Fungi (Stereaceae).
U.S.A.

stereocaulorum F. Arnold, Flora, Jena und Re-
gensburg 57:175. 1874. =Metasphaeria
stereocaulorum (F. Arnold) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:183.
1883. Parasitic on thallus of Stereocaulon
alpinum Laur., Lichenes (Stereocaulaceae).
Italy.

Sticta J.B. Ellis & B.M. Everhart, Journa! of
Mycology, Columbus, Ohio 1:43. 1885. On
dead culms of Spartina sp., Gramineae.
U.S.A.

Stictoides (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:86. 1883. =Sphaeria stictoides M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:149. 1876. On branches
of Liriodendron sp., Magnoliaceae. U.S.A.

Stictostoma (M.J. Berkeley & M.A. Curtis) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo

Nomenclator of Leptosphaeria

279

2:84. 1883. =Sphaeria stictostoma M.J.
Berkeley & M.A. Curtis in M.J. Berkeley,
Grevillea, London 4:151. 1876. On stems of
herbaceous plant of unknown host, unknown
family. U.S.A.

stipae-minor L. Lacoste, Revue de Mycologie 22,
Paris (Supplement Colonial No. 1) 22:12-13.
1957. On leaves of Stipa tenacissima L.,
Gramineae. Tunisia.

stipae L. Trabut, Etude sur L’Halfa (Stipa tenacis-
sima) Alger (Adolphe Jourdan...), p. 48. 1889.
[Leptosphaeria pampaniniana Sacc. may be a
synonym.] On decayed leaves of Stipa
tenacissima L., Gramineae. Algeria.

stipae A.\. Lobik, Materialy po Floristicheskim i
Faunisticheskim Obsledovaniyam Terskogo
Okruga [Data from Investigations on the Flora
and Fauna of the Ter Region], p. 24. 1928.
Nom. inval. Art. 36.1. On leaves of Stipa
capillata L., Gramineae. U.S.S.R.

straminis M.C. Cooke & H.W. Harkness, Grevillea,
London 14:10. 1885. =Pleospora harknessii
A.N. Berlese & P. Voglino, Sylloge Fun-
gorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo Additamenta Ad
Volumina I-IV, p. 174. 1886. Nec Pleospora
straminis P.A. Saccardo & C.L. Spegazzini.
On culms of straw, Gramineae. U.S.A.

stratiotis C.A. Oudemans, Beihefte zum Botanischen
Zentralblatt, Cassel 11:528. 1902. On leaves
of Stratiotes aloides L., Hydrocharitaceae.
Netherlands.

striata H.G. Winter, Hedwigia, Dresden 11:147.
1872. On dry stems of Veronica latifolia
auct., Scrophulariaceae. Germany.

striolata G., Passerini, Atti della R. Accademia dei
Lincei Memoriae, Rome, Series 4, No. 6, p.
459. 1890; Diagnosi Di Funghi Nuovi, Nota
IV, p. 5. 1890. =Mycotodea striolata (G.
Passerini) W. Kirschstein, Kryptogamenflora
der Mark Brandenburg und Angrenzender
Gebiete herausgegeban von dem Botanischen
Verein der Provinz Brandenburg), Leipzig
7(3):434. 1938. On dry stems of Scirpus
holoschoenus L., Cyperaceae. Italy.

striolata G. Passerini var. caricis-glaucae L.M.
Unamuno Yrigoyen, Anales del Jardin
Botanico de Madrid 2:43-44. 1942. On dry
leaves of Carex glauca Scop., Cyperaceae.
Spain.

stromatoidea F.A. Hazslinszky, Matematikai és
Természettudomanyi Ké6zlemenyek Von-
atkoz6lag a Hazai Viszonyokra, Budapest
25(2):135. 1892. On culms of Festuca sp.,
Gramineae. Hungary.

suaedae C.G. Hansford, Proceedings of the Linnean
Society of New South Wales, Sydney 82:215.
1957. On dead branches of Suaeda australis
Mogq., Chenopodiaceae. Australia.

subalpina F. Bubak, Botanikai K6zlemények,
Budapest 14:(59). 1915. =Phaeosphaeria

280

subalpina (F. Bubak) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1573.
1989. On crack (cleft) of sheaths of Phrag-
mites communis Trin., Gramineae. Yugosla-
Via.

subarticulata (F. Arnold) K. Keissler, Dr. L.
Rabenhorst’s Kryptogamen-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 9(1 Abt., Teil 2):373. 1937.
=Sagedia subarticulata F. Arnold, Verhand-
lungen der Zoologisch-Botanischen
Gesellschaft in Wien 21:1133—1134. 1871.
=Segestria subarticulata (F. Arnold) Dalla
Torre & L. Sarntheim, Flora der Gefiirsteten
Grafschaft Tirol, des Landes Vorarlberg und
des Fiirstenthumes Liechtenstein 4:568. 1902.
=Porina subarticulara (F. Arnold) G. Lettau,
Hedwigia, Dresden 52:105. 1912. On thallus
of unknown host, unknown family [originally
described as a lichen]. Italy.

subcaespitosa M.C. Cooke & H.W. Harkness,
Grevillea, London 14:10. 1885. On stems of
Geranium sp., Geraniaceae. U.S.A.

subcompressa J.H. Miller & M.G. Burton, Mycolo-
gia, Lancaster, Pennsylvania 34:4. 1942. On
dead culms of Andropogon glomeratus
(Walt.) B.S.P., Gramineae. U.S.A.

subconica (G.W. Clinton & C.H. Peck) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:15. 1883. =Sphaeria (Caulicolae) subconia
G.W. Clinton & C.H. Peck, Report. New
York State Museum of Natural History,
Albany 29:65. 1878. On dead stems of herbs,
unknown host, unknown family. U.S.A.

subcutanea (M.C. Cooke & J.B. Ellis) J.B. Ellis in
N.L. Britton, Catalogue of Plants Found in
New Jersey. Geological Survey of New
Jersey, Final Report of the State Geologist,
Trenton 2(1):525. 1889. =Sphaeria (sub-
tectae) subcutanea M.C. Cooke & J.B. Ellis,
Grevillea, London 7:41. 1878. =Metas-
Phaeria subcutanea (M.C. Cooke & J.B. Ellis)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:167. 1883. =Conisphaeria subcutanea
(M.C. Cooke & J.B. Ellis) M.C. Cooke,
Grevillea, London 16:87. 1888. =Endophlaea
subcutanea (M.C. Cooke & J.B. Ellis) M.C.
Cooke, Grevillea, London. 17:89. 1889. On
decorticated limbs of Pyrus communis L., Ro-
saceae. U.S.A. (New Jersey).

subiculifera? C.L. Spegazzini, Anales del Museo
Nacional de Historia Natural de Buenos Aires
6:283. 1898; Fungi Argentini Novi vel Critici,
No. 630. Anno 1899. On decaying leaves of
Gynerium argenteum Nees, Gramineae.
Argentina.

sublanosa (M.C. Cooke) J.B. Ellis in N.L. Britton,
Catalogue of Plants Found in New Jersey.
Geological Survey of New Jersey, Final

Illinois Natural History Survey Bulletin

Vol. 34 Ar. 3

Report of the State Geologist, Trenton
2(1):525. 1889. =Sphaeria (Obtectae)
sublanosa M.C. Cooke in M.C. Cooke and
J.B. Ellis, Grevillea, London 7:41. 1878. On
decorticated parts of Andromeda sp.,
Ericaceae. U.S.A. (New Jersey).

submaculans L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):38. 1957. On dry
stems of Berteroa incana (L.) DC., Erysimum
cheiranthoides L., Sisymbrium loeselii L.,
Cruciferae. Germany.

submodesta E. Miiller, Sydowia. Annales Mycol-
ogici, Horn, N.O. 5(3-6):53—54. 1951. On
dead stems of Tofieldia calyculata (L.)
Wahlenb., Liliaceae. Switzerland.

subriparia V. Mouton, Bulletin. Société R. de
Botanique de Belgique, Bruxelles 39:44.
1900. On stalks of Juncus glaucus Sibth.,
Juncaceae. Belgium.

subsimilis (S. Schulzer v. Miiggenburg & P.A.
Saccardo) F.A. Hazslinszky, Matematikai és
Természettudomanyi K6zlemenyek Von-
atkoz6lag a Hazai Viszonyokra, Budapest
25(2):151. 1892. =Metasphaeria subsimilis S.
Schulzer v. Miiggenburg & P.A. Saccardo,
Revue Mycologique, Toulouse 6:70. 1884.
On branches of Carpinus sp., Betulaceae.
Yugoslavia.

substerilis C.H. Peck, Bulletin of the New York
State Museum, Albany 105:24—25. 1906. On
living leaves of Mentha piperita L., Labiatae.
U.S.A.

subsuperficialis P.A. Saccardo & P. Sydow in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
16:516. 1902. =Montagnula subsuperficialis
(P.A. Saccardo & P. Sydow) R.A. Shoemaker,
Canadian Journal of Botany, Ottawa 67:1591.
1989. =Leptosphaeria melanommoides C.L.
Spegazzini. On fallen rotten culms of
Panicum grumosum Nees, Gramineae.
Argentina.

subtecta H.G. Winter, Hedwigia, Dresden 11:146.
1872. On dry leaf of Erica carnea L..,
Ericaceae. Italy.

suffulta (C.G. Nees v. Esenbeck:E.M. Fries) G
Niessl v. Mayendort in G.L. Rabenhorst,
Fungi Europaei Exsiccati, Klotzschii Herbarii
vivi Mycologici Continuatio Edita Nova,
Series 2, Century 16, No. 1549. Anno 1872.
=Sphaeria suffulta C.G. Nees v. Esenbeck:
E.M. Fries, Das System der Pilze und
Schwamme, p. 316. 1816; E.M. Fries,
Systema Mycologicum Sistens Fungorum
2:508-509. 1823. On dry stems of Melampy-
rum pratense L., Scrophulariaceae. Austria,
Italy.

superficialis F.A. Hazslinszky, Matematikai és
Természettudomanyi Kézlemenyek Von-
atkozolag a Hazai Viszonyokra, Budapest
25(2):146. 1892. On unknown substrate of
Potentilla argentea L., Rosaceae. Hungary.

March 1991]

surculorum W. Kirschstein, Annales Mycologici,
Berlin 33:212—213. 1935. On hard, living,
annual shoots of Lonicera tatarica L.,
Caprifoliaceae. Germany.

swertiae M.J. Thirumalachar, Bulletin of the Torrey
Botanical Club (and Torreya), New York
75:176. 1948. Nom. nud. Arts. 32.1 and 34.1.
On leaves of Swertia sp., Gentianaceae.
India.

sydowiana H. Rehm in P. Sydow, Mycotheca
Marchica, Century 15, No. 1456. Anno 1887.
Nom. nud. Art. 32.1. On unknown substrate
of Artemisia annua L., Compositae. Ger-
many.

sydowiana H. Rehm in P. Sydow, Mycotheca
Marchica, Century 15, No. 1484. Anno 1887.
Nom. nud. Art. 32.1. On unknown substrate
of Artemisia campestris L., Compositae.
Germany.

sydowiana H. Rehm in P. Sydow, Mycotheca
Marchica, Century 9, No. 849. Anno 1885.
Nom. nud. Art. 32.1. On unknown substrate
of Artemisia campestris L., Compositae.
Germany.

sylvatica G. Passerini in G.L. Rabenhorst, Hedwigia,
Dresden 16:118. 1877; Fungi Europaei
Exsiccati, Klotzschii Herbarii vivi Mycologici
Continuatio Edita Nova, Series 2, Century 23,
No. 2235. Anno 1877. =Phaeosphaeria
silvatica (G. Passerini) A. Hedjaroude,
Sydowia. Annales Mycologici, Horn, N.O.
22(1-4):91. (1968) 1969. On leaves of
Brachypodium sylvaticum (Hudson) Beauv.,
Gramineae. Italy, Switzerland, Turkey.

taichungensis J. Yen, Bulletin de la Société
Mycologique de France, Paris 89:67—-69.
1973. On leaves of Musa sp., Musaceae.
Taiwan (Puli).

taiwanensis W.Y. Yen & C.C. Chi, Journal of Sugar
Cane Research, Taiwan 6:213. 1952.
=Didymella taiwanensis (W.Y. Yen & C.C.
Chi) R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 67:1576. 1989. On decaying
leaves of Saccharum officinarum L., Gramin-
eae. Taiwan.

tamaricis L. Hollos, Botanikai K6zlemények,
Budapest 25:130. 1928. Nom. illegit. Art.
64.1. =Leptosphaeria hollosii G. Moesz. On
dry branches of Tamarix gallica L., Tamari-
caceae. Hungary.

tamaricis (R.K. Greville) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:26. 1883. =Crypto-
sphaeria tamaricis R.K. Greville, Scottish
Cryptogamic Flora 1, Fascicle 9, Table 45.
1823. [Text unpaginated; as tamariscinis.|
=Sphaeria tamaricis (R.K. Greville) F.
Currey, Transactions of the Linnean Society
of London 22:324. 1859 [as tamariscinis]. On
dead branches of Tamarix germanica L.,
Tamaricaceae. Great Britain.

Nomenclator of Leptosphaeria

281

(Metasphaeria) taminensis H. Wegelin, Mitteilungen
der Thurgauischen Naturforschenden
Gesellschaft, Frauenfeld 12:173. 1896.
=Trichometasphaeria taminensis (H.
Wegelin) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):142. 1957. On
dried culms of Festuca sp., Gramineae.
Switzerland.

tanaceti (L. Fuckel) L. Holm, Symbolae Botanicae
Upsalienses, Uppsala 14(3):49. 1957. Nom.
illegit. Art. 64.1. [A later homonym of
Leptosphaeria tanaceti A. Jaczewski.]
=Sphaerulina tanaceti L. Fuckel, Jahrbuch
des Nassauischen Vereins fiir Naturkunde,
Wiesbaden 15:77—78. 1860 [as Sphaerolina].
=Ohiobolus tanaceti (L. Fuckel) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:348. 1883. On wilted leaves of Achillea sp.,
Anthemis tinctoria L., Chrysanthemum
leucanthemum L., Chrysanthemum vulgare
(L.) Bernh., Compositae. Austria, Germany.

tanaceti A. Jaczewski, Bulletin de la Société
Mycologique de France, Paris 9:217. 1893.
On stems of Tanacetum vulgare L., Composi-
tae. U.S.S.R.

tartarina? (W. Nylander) W. Zopf, Hedwigia,
Dresden 35:341. 1896. =Verrucaria tartarina
W. Nylander, Flora, Jena und Regensburg
57:318. 1874. On thallus of Ochrolechia
tartarea (L.) Massal, Lichenes (Pertusari-
aceae). Unknown country.

taurica N.A. Naumov & T.L. Dobrozrakova,
Materialy po Mikologii i Fitopatologii Rossi,
Petrograd 8(2):134. 1929. On dry stems of
Cephalaria uralensis (Murray) Roemer &
Scultes, Dipsacaceae. U.S.S.R.

taxicola (C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:85. 1883.
=Sphaeria taxicola C.H. Peck, Report. New
York State Museum of Natural History,
Albany 24:99. 1872. =Metasphaeria taxicola
(C.H. Peck) C.H. Peck, Report. New York
State Museum of Natural History, Albany
39:58. 1886. [Combination not formerly
proposed.] =Sphaerulina taxicola (C.H.
Peck) A.N. Berlese, [cones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:125.
1894. =Saccothecium taxicolum (C.H. Peck)
W. Kirschstein, Kryptogamenflora der Mark
Brandenburg und Angrenzender Gebiete
herausgegeben von dem Botanischen Verein
der Provinz Brandenburg, Leipzig 7(3):427.
1938. =Dothiora taxicola (C.H. Peck) M.E.
Barr, Contributions from the University of
Michigan Herbarium, Ann Arbor 9:575. 1972.
On dead leaves of Taxus canadensis Marsh.,
Taxaceae. U.S.A.

282 Illinois Natural History Survey Bulletin

tenera J.B. Ellis, Bulletin of the Torrey Botanical
Club (and Torreya), New York 8:124—-125.
1881 [as Leptosphaeria tenera (J.B. Ellis)
P.A. Saccardo in Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:39. 1883]. On dead herbaceous stems of
unknown host, unknown family. U.S.A.

tenuis E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):221. 1950. On dead stems
of Thalictrum aquilegifolium L., Ranuncu-
laceae. Switzerland.

tephrosiae (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:27.
1883. =Sphaeria (Obtectae) tephrosiae M.C.
Cooke & J.B. Ellis, Grevillea, London 7:10.
1878. On stems of Tephrosia virginiana (L.)
Pers., Leguminosae. U.S.A.

ternata F. Hazslinszky, Matematikai és Természettu-
domanyi K6zlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 25(2):150. 1892. On
branches of Pyrus malus L. (=Malus domes-
tica Borkh.), Rosaceae. Hungary.

tetonensis L.E. Wehmeyer, Lloydia: a quarterly
journal of biological science, Manasha 9:234.
1946. Nom. illegit. Art. 64.1. =L. jacksonen-
sis R.A. Shoemaker.

tetonensis (J.B. Ellis & B.M. Everhart) H. Rehm,
Annales Mycologici, Berlin 9:289. 1911.
=Melanomma tetonensis J.B. Ellis & B.M.
Everhart, Proceedings of the Academy of
Natural Sciences of Philadelphia 1890:240.
1890. =Melanomma occidentale (J.B. Ellis)
P.A. Saccardo var. tetonensis J.B. Ellis &
B.M. Everhart, The North American Pyreno-
mycetes. A Contribution to Mycologic
Botany, p. 183. 1892. =Syncarpella tetonen-
sis (J.B. Ellis & B.M. Everhart) M.E. Barr &
J.R. Boise, Memoirs of the New York
Botanical Garden, Bronx 49:303. 1989. On
stems of Artemisia frigida Willd., Composi-
tae. U.S.A. (California, Colorado, Montana,
Utah). ‘

tetraspora A.M. Saccas, Etude de la Flore Cryptoga-
mique des Caféiers en Afrique Centrale.
Bulletin de Institut Frangais du Café du Cacao
et d’Autres Plantes Stimulantes (Bulletin
IFCC No. 16), pp. 260-262. 1981. Nom.
inval. Art. 37.1. On dead branches of Coffea
robusta L. Linden (=Coffea canephora Pierre
ex Froehn.), Rubiaceae. Central African
Republic.

teucrii (P.L. Crouan & H.M. Crouan) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:85.
1883. =Sphaeria teucrii P.L. Crouan & H.M.
Crouan, Florule Du Finistére Contenant Les
Descriptions De 360 Espéces Nouvelles De
Sporogames, De Nombreuses Observations et

Vol. 34 Art. 3

une synonymie des plantes Cellulaires et
Vasculaires Qui Croissent Spontanément
Dans ce Département, p. 28. 1867. On dead
stems of Teucrium scorodonia L., Labiatae.
France.

thalictri H.G. Winter, Hedwigia, Dresden 11:147.
1872. =Metasphaeria thalictri (H.G. Winter)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:156. 1883. =Scleropleella thalictri (H.G.
Winter) F. v. Héhnel, Annales Mycologici,
Berlin 18:76. 1920. =Buergenerula thalictri
(H.G. Winter) E. Miiller, Sydowia. Annales
Mycologici, Horn, N.O 4(1-6):307. 1950. On
dry stems of Thalictrum aquilegifolium L.,
Ranunculaceae. Italy.

thalictricola L. Hollos, Annales Historico-Naturales
Musei Nationalis Hungarici, Budapest
7:51-52. 1909. On dry stems of Thalictrum
collinum Wallr., Ranunculaceae. Hungary.

thalictrina L. Hollés, Matematikai és Természettu-
domanyi Kézlemenyek Vonatkozolag a Hazai
Viszonyokra, Budapest 35:34. 1926. On dry
stems of Thalictrum aquilegifolium L.,
Ranunculaceae. Hungary.

theobromicola R. Ciferri & R. Gonzalez Fragoso,
Boletin de la R. Sociedad Espanola de
Historia Natural, Madrid 26:474-475. 1926:
Publicaciones Estaci6n Agronomica de Moca,
Santa Domingo. Series B, Botanica 8:23.
1927. Nom. nud. Art. 34.1b. On living leaves
of Theobroma cacao L., Sterculiaceae.
Dominican Republic.

therophila B. Auerswald in W. Gonnermann and
G.L. Rabenhorst, Mycologia Europaea,
Abbildungen Sammtlicher Pilze Europa’s,
Hefte 5 und 6, Synopsis Pyrenomycetum
Europaeorum, Table 12, Figure 158. 1869.
Nom. nud. Art. 32.1. =Sphaeria (Caulicola)
therophila J. Desmaziéres, Annales des
Sciences Naturelles, Paris, Botanique, Series
3, 16:310. 1851. =Phomatospora therophila
(J. Desmaziéres) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 1:433. 1882. On dry
culms of Juncus articulatus L., Juncaceae.
France.

thielensii (G.D. Westendorp) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:46. 1883. =Ple-
ospora thielensti (G.D. Westendorp) J.B.
Lambotte, Flore Mycologique de la Belgique,
Comprenant la description des especes
trouvées jusqu’a ce Jour. 2:274—275. 1880.
=Sphaeria thielensii G.D. Westendorp,
Bulletin. Société R. de Botanique de Bel-
gique, Bruxelles 5:37. 1866. On dead stems
of Tanacetum vulgare L., Compositae.
Belgium.

March 1991

thomasiana P.A. Saccardo & C. Roumeguére, Revue
Mycologique, Toulouse 5:236. 1883. On
dead runners of Rubus sp., Rosaceae. France.

thorae O. Jaap, Annales Mycologici, Berlin 6:210.
1908. On living leaves of Ranunculus thora
L., Ranunculaceae. Austria.

thuemeniana G. Niessl v. Mayendorf in F. v.
Thiimen, Instituto. Revista Scientifica e
Litteraria, Coimbra 28:29. 1881; Contribu-
tiones ad Floram Mycologicam Lusitanicam,
Series 3, p. 29. 1881. =Heptameria thuemeni-
ana (G. Niessl v. Mayendorf) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:89.
1883. On somewhat rotten stems of Antir-
rhinum majus L., Scrophulariaceae. Portugal.

thujaecola K. Hara, Reports of the Shizuokaken
Prefecture Agricultural Society 34:(49)-(S0).
1930. [In Japanese.] On leaves of Thuja
orientalis L., Cupressaceae. Japan.

thurgoviensis E. Miiller, Sydowia. Annales Mycol-
ogici, Horn, N.O. 4(1-6):280. 1950. =Mas-
sariosphaeria thurgoviensis (E. Miiller) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1589. 1989. On dead grass culms
of unknown host of Gramineae, Gramineae.
Switzerland.

thyphicola G. Passerini & V. Beltrani, Atti dell’
Accademia Nazionale dei Lincei. Transunti,
Rome, Series 3, 7:36. (1882) 1883. =Lepto-
sphaeria licatensis P.A. Saccardo. Non P.A.
Karsten. On dry leaves of Typha latifolia L.,
Typhaceae. Italy (Sicily).

tigrisoides K. Hara, The Diseases of Trees (Zikken-

‘ Zyumoku-Byogaihen), p. 331. 1927. On
unknown substrate of Bambusa sp., Gramin-
eae. Japan.

tint J.B. Ellis & B.M. Everhart, Journal of Mycol-
ogy, Columbus, Ohio 4:64. 1888. On leaves
of Viburnum tinus L., Caprifoliaceae. U.S.A.

tiroliensis W. Kirschstein, Annales Mycologici,
Berlin 33:213. 1935. On dry branches of
Sambucus racemosa L., Caprifoliaceae.
Austria.

tofieldiae E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 5(3-6):53. 1951. =Phaeosphaeria
tofieldiae (E. Miiller) A. Leuchtmann,
Sydowia. Annales Mycologici, Horn, N.O.
37:110. 1984. On leaves of Tofieldia
calyculata (L.) Wahlenb., Liliaceae. Switzer-
land.

tolgorensis F. Petrak, Sydowia. Annales Mycologici,
Horn, N.O. 3(1-6):281. 1949. On dry stems
of Euphorbia sp., Euphorbiaceae. Iran.

tompkinsii A.S. El-Ani, Mycologia, Lancaster,
Pennsylvania 58:409. 1966. On Homo
sapiens, Hominidae. Mauritania.

tonduzi C.L. Spegazzini, Boletin de la Academia
Nacional de Ciencias en Cérdoba 23:562.

Nomenclator of Leptosphaeria 283

1919; Reliquiae Mycologicae Tropicae et
Fungi Costaricenses Nonnulli, pp. 200-201.
1919. On living leaves of Coffea sp.,
Rubiaceae. Costa Rica.

torbolensis W. Kirschstein, Annales Mycologici,
Berlin 37:108. 1939. On dry stems of
Scrophularia canina L., Scrophulariaceae.
Italy.

tornatospora T. Petch, Ceylon Journal of Science,
Colombo, Section A (Botany) 9:319. 1925.
On dead twigs of Camellia theifera Griff.,
Theaceae. Sri Lanka.

torrendii G.B. Traverso & C. Spessa, Boletim da
Sociedade Broteriana, Coimbra 25:171. 1910
[as torrendi]. On dry branches of Ricinus
communts L., Euphorbiaceae. Portugal.

torulispora (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:42. 1883.
=Sphaeria (Caulicolae) torulaespora M.C.
Cooke, Journal of the Linnean Society
(Botany) London 17:144. 1878. On herba-
ceous stems of unknown host, unknown
family. U.S.A.

translucens H.G. Winter, Boletim da Sociedade
Broteriana, Coimbra 2:42. (1883) 1884. On
dry leaves of Furcraea sp., Labiatae.
Portugal.

treatiana P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 10:923. 1892 [in a footnote]. Nom.
illegit. Art. 63.1.

trematostoma J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(3):214. 1903. On dry barked branches of
Syringa vulgaris L., Oleaceae. Luxembourg.

trevoae C.L. Spegazzini, Fungi Chilenses, p. 79.
1910; Revista de La Facultad de Agronomia
Y Veterinaria, Universidad Nacional de La
Plata, Series 2, 6:79. 1910. On dead branches
of Trevoa trinervia Gill. & Hook., Rham-
naceae. Chile.

trichopterygis C.G. Hansford, Proceedings of the
Linnean Society of London 153:25. 1941. On
dead culms of Trichopteryx sp. [as T.
afroflammida\, Gramineae. Uganda.

trichostoma G. Passerini in F. v. Thiimen, Myco-
theca Universalis, Century 15, No. 1455.
Anno 1879; Flora, Jena und Regensburg
63:324. 1880. =Metasphaeria trichostoma
(G. Passerini) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:158. 1883. On
stems and branches of Chondrilla juncea L.,
Compositae. Italy.

trifolit J. Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):210.
1903. On dry stems of Trifolium medium L.,
Leguminosae. Luxembourg.

Illinois Natural History Survey Bulletin

trifolii-alpestris T. Dominick, Acta Societatis

Botanicorum Poloniae, Warszawa
11:239-240. 1934. On living leaves of
Trifolium alpestre L., Leguminosae. Poland.

triglochinicola (F. Currey) P.A. Saccardo, Sylloge

Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:69. 1883.
=Sphaeria triglochinicola F. Currey,
Transactions of the Linnean Society of
London 24:158—159. (1863) 1864.
=Phaeosphaeria triglochinicola (F. Currey)
A. Leuchtmann, Sydowia. Annales Mycol-
ogici, Horn, N.O. 37:111. 1984. On carpels
and stems of Triglochin palustris L., Juncag-
inaceae. Great Britain, Switzerland.

triglochinis J. Schroter, Kryptogamen-Flora Von

Schlesien. Im Namen Der Schlesischen
Gesellschaft fiir vaterlandische Cultur
herausgegeben von Prof. Dr. Ferdinand Cohn,
Secretair der Botanischen Section, Breslau
3(2):366. 1894. On dead stems of Triglochin
palustris L., Juncaginaceae. Poland.

trimera P.A. Saccardo, Nuovo Giornale Botanico

Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:319. 1875. On dead stems
of Andropogon ischaemum L., Gramineae.
Italy.

trimerioides H. Rehm, Annales Mycologici, Berlin

13:2. 1915. On stems of Liatris scariosa (L.)
Willd., Compositae. U.S.A.

tritici (S. Garovaglio) G. Passerini in G.L. Raben-

horst, Fungi Europaei Exsiccati, Klotzschii
Herbarii vivi Mycologici Continuatio Edita
Nova, Series 2, Century 24, No. 2333. Anno
1876. =Pleospora tritici S. Garovaglio,
Rendiconti dell’ Istituto Lombardo di Scienze
e Lettere, Milano, Series 2, 6:61 1—612. 1873.
=Leptosphaeria eustoma (E.M. Fries:E.M.
Fries) P.A. Saccardo forma tritici (S.
Garovaglio) A.N. Berlese, Icones Fungorum
Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 1:56.
1892 [as (Garov.) Pass.]. =Phaeosphaeria
tritici (S. Garovaglio) A. Hedjaroude,
Sydowia. Annales Mycologici, Horn, N.O.
22:74. 1968. On leaves of Triticum vulgare
Vill., Gramineae. Italy, Kenya.

tritici (S. Garovaglio) G. Passerini var. papyricola

J.B. Ellis & B.M. Everhart, Report of the
Michigan Academy of Science, Lansing 7:66.
1905 [as papicola]. Nom. nud. Art. 32.1. On
old paper. U.S.A. (Michigan).

tritorulosa (M.J. Berkeley & C.E. Broome) V. Cesati

& G. de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:236. 1863.
=Sphaeria (Caulicolae) tritorulosa M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London, Series
2, 13:466. 1854. =Metasphaeria tritorulosa

Vol. 34 Ar. 3

M.J. Berkeley & C.E. Broome, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 15:190. 1901 [as
Metasphaeria bitorulosa}. On dead stems of
Epilobium hirsutum L., Onagraceae. Great
Britain.

(Astrosphaeria) trochus (O.A.J. Penzig & P.A.

Saccardo) F. v. Hohnel, Sitzungsberichte der
Akademie der Wissenschaften in Wien,
Mathematisch-naturwissenschaftliche Klasse
118(Abt. 1):328. 1909. =Melanomma trochus
O.A.J. Penzig & P.A. Saccardo, Malpighia.
Rassegna Mensuale di Botanica. Messina,
Genova 11:401. 1897; Icones Fungorum
Bambusicolorum Japonicorum, p. 16. 1904.
=Trematosphaeria trochus (O.A.J. Penzig &
P.A. Saccardo) S.C. Teng, Sinensia. Special
Bulletin of the Metropolitan Museum of
Natural History, Nanking 9:257. 1938.
=Asterotheca trochus (O.A.J. Penzig & P.A.
Saccardo) I. Hino, Bulletin Miyazaki College
of Agriculture and Forestry 10:57. 1938.
=Asterella trochus (O.A.J. Penzig & P.A.
Saccardo) K. Hara, Nippon Baikin-Gaku, p.
187. 1936. =Astrosphaeriella trochus (O.A.J.
Penzig & P.A. Saccardo) D.L. Hawksworth,
Journal of the Linnean Society (Botany)
London 82:46. 1981. On rachis of palm,
Palmae. China, Java.

trollii (P.A. Karsten) E. Miiller, Sydowia. Annales

Mycologici, Horn, N.O. 4(1-6):249. 1950.
=Metasphaeria trolii P.A. Karsten, Hedwigia,
Dresden 23:86. 1884. On dry stems of
Cardamine sylvatica Link, Trollius europaeus
L., Cruciferae, Ranunculaceae. Switzerland.

tucumanensis C.L. Spegazzini, Revista. Universidad

Nacional de La Plata 2(18):232—233. 1896;
Hongos De La Cana De Azitcar No. 10. Anno
1896. On culms of Saccharum officinarum
L., Gramineae. Argentina.

tumefaciens (J.B. Ellis & H.W. Harkness) F. Petrak,

Annales Mycologici, Berlin 32:360—361.
1934. =Sphaeria (Montagnella) tumefaciens
J.B. Ellis & H.W. Harkness, Journal of
Mycology, Columbus, Ohio 2:41. 1886.
=Montagnella tumefaciens (J.B. Ellis & H.W.
Harkness) A.N. Berlese & B. Voglino in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
Additamenta Ad Volumina I-IV, p. 244.
1886. =Phaeoderris tumefaciens (J.B. Ellis &
H.W. Harkness) F. v. Hohnel, Sitzungsber-
ichte der Akademie der Wissenschaften in
Wien, Mathematisch-naturwissenschaftliche
Klasse, Abt. I, 120:462. 1911. =Syncarpella
tumefaciens (J.B. Ellis & H.W. Harkness) F.
Theissen & H. Sydow, Annales Mycologici,
Berlin 13:631. 1915. On dead limbs of
Artemisia californica Less., Artemisia sp..

March 1991

Compositae. U.S.A. (California), U.S.S.R.
(Turkistan).

tungurahuensis F. Petrak, Sydowia. Annales
Mycologici, Horn, N.O. 2(1-6):338. 1948.
On stroma of Phyllachora tungurahuensis
Petrak, Fungi (Phyllachoraceae). Philippines.

tupae C.L. Spegazzini, Fungi Chilenses, pp. 79-80.
1910; Revista de La Facultad de Agronomia
Y Veterinaria, Universidad Nacional de La
Plata, Series 2, 6:79-80. 1910. On year-old
shoots of Lobelia tupa L., Campanulaceae.
Chile.

typhae (P.A. Karsten) P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:321.
1875. =Leptosphaeria perpusilla (J. Des-
maziéres) P.A. Karsten var. typhae P.A.
Karsten, Mycologia Fennica Pars 2, Pyreno-
mycetes, p. 99. 1873; Bidrag till Kannedom
om Finlands Natur och Folk. Utgifna af
Finska Vetenskaps-Societeten, Helsingfors
23:99. 1873. =Sphaeria perpusilla J.

Desmazieres var. typhae B. Auerswald in G.L.

Rabenhorst, Fungi Europaei Exsiccati,
Klotzschii Herbarii vivi Mycologici Continu-
atio Edita Nova, Series Secunda, Century 9,
No. 831. Anno 1865. Nom. nud. Art. 32.1.
=Phaeosphaeria typhae (P.A. Karsten) R.A.
Shoemaker, Canadian Journal of Botany,
Ottawa 67:1536. 1989. On decaying culms of
Typha latifolia L., Typhaceae. Finland,
Germany.

typharum (J. Desmaziéres) P.A. Karsten, Mycologia
Fennica Pars 2, Pyrenomycetes, p. 100. 1873;
Bidrag till Kannedom om Finlands Natur och
Folk. Utgifna af Finska Vetenskaps-Soci-
eteten, Helsingfors 23:100. 1873. =Sphaeria
scirpicola A.P. de Candolle:E.M. Fries var.
typharum J. Desmazieéres, Plantes Crypto-
games Du Nord De La France, Edition 2,
Fascicle 36, No. 1778. Anno 1849.
=Sphaeria typharum (J. Desmaziéres) G.L.
Rabenhorst, Klotzschii Herbarium Vivum
Mycologicum Sistens Fungorum Per Totam
Germaniam Cresentium Collectionem
Perfectam, Dresden, Editio Novo, Century 8,
No. 731. Anno 1858. =Pleospora typharum
(J. Desmaziéres) L. Fuckel, Fungi Rhenani
Exsiccati A Leopoldo Fuckel Collecti,
Fascicle 9, No. 858. Anno 1864; Symbolae
Mycologicae, p. 137. 1870. =Phaeosphaeria
typharum (J. Desmaziéres) L. Holm, Symbo-

lae Botanicae Upsalienses, Uppsala 14(3):126.

1957. On decaying leaves of Typha latifolia
L., Typhaceae. Belgium, Canada, Finland,

France, Germany, Italy, New Zealand, U.S.A.,

U.S.S.R.

typharum (L. Fuckel) P.A. Saccardo, Atti dell’
Accademia Scientifica Veneto-Trentino-
Istriana, Padova 2(2):154. 1873. =Pleospora

Nomenclator of Leptosphaeria

typharum (J. Desmazieres) L. Fuckel, Fungi
Rhenani Exsiccati A Leopoldo Fuckel
Collecti, Fascicle 9, No. 858. Anno 1864.
{P.A. Saccardo distinctly attributed the
basionym to L. Fuckel.]

typharum (G.L. Rabenhorst) B. Auerswald in J.
Kunze, Fungi Selecti Exsiccati, Series 11, No.
256. Anno 1877 [as Tauschverein for possible
earlier place of publication]. =Sphaeria
typharum G.L. Rabenhorst, Fungi Europaei
Exsiccati, Klotzschii Herbarii vivi Mycologici
Continuatio, Edition 2, Century 8, No. 731.
Anno 1858. =Pleospora typharum (G.L.
Rabenhorst) L. Fuckel, Fungi Rhenani
Exsiccati A Leopoldo Fuckel Collecti,
Fascicle 9, No. 858. Anno 1864; Symbolae
Mycologicae, p. 137. 1870. =Leptosphaeria
kunzeana A.N. Berlese. On dry leaves and
standing stems of Typha angustifolia L.,
Typha latifolia L., Typhaceae. Germany.

typharum (J. Desmaziéres) P.A. Karsten forma acori
W.B. Grove, Journal of Botany, British and
Foreign, London 68:97. 1930. On unknown
substrate of Acorus calamus L., Araceae.
Great Britain.

typharum (J. Desmaziéres) P.A. Karsten subsp.
papyrogena P.A. Saccardo, Annales Mycol-
ogici, Berlin 6:558. 1908. On putrid paper.
France.

typharum (J. Desmazieres) P.A. Karsten subsp.
phragmatina P.A. Karsten, Hedwigia,
Dresden 23:5. 1884; Acta Societatis pro
Fauna et Flora Fennica, Helsingforsiae
2(6):54. 1885. On decaying leaves of
Phragmites communis Trin., Gramineae.
Finland.

typhicola G. Passerini & V. Beltrani, Atti dell’
Accademia Nazionale dei Lincei. Transunti,
Rome, Series 3, 7:36. 1883. =Leptosphaeria
licatensis P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:70. 1883. Nec Leptosphaeria
typhicola P.A. Karsten, fide P.A. Saccardo
1883. =Leptosphaeria typhicola P.A.
Karsten. On dry leaves of Typha /atifolia L.,
Typhaceae. Italy.

typhicola P.A. Karsten, vide P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:70. 1883 sub.
Leptosphaeria licatensis P.A. Saccardo.
=Leptosphaeria maculans (M.J. Sowerby)
P.A. Karsten var. typhicola P.A. Karsten,
Mycologia Fennica Pars 2, Pyrenomycetes, p.
100. 1873. =Massariosphaeria typhicola
(P.A. Karsten) A. Leuchtmann, Sydowia.
Annales Mycologici, Horn, N.O. 37:168.
1984. =Phaeosphaeria typhicola (P.A.
Karsten) A. Hedjaroude, Sydowia. Annales
Mycologici, Horn, N.O. 22:86. 1968.
=Chaetomastia typhicola (P.A. Karsten) M.E.

286 Illinois Natural History Survey Bulletin

Barr, Mycotaxon. An International Journal
Designed to Expedite Publication of Research
on Taxonomy & Nomenclature of Fungi &
Lichens, Ithaca, New York 34:514. 1989.
Canada, Czechoslovakia, Switzerland.

typhiseda F. Petrak, Hedwigia, Dresden 65:220-221.
1925. Nom. illegit. Art. 64.1. On dry flower
stalks of Typha angustifolia L., Typhaceae.
Poland.

typhiseda P.A. Saccardo & A.N. Berlese, Revue
Mycologique, Toulouse 8:33. 1886. =Lepto-
sphaeria praeclara P.A. Karsten forma
typhiseda (P.A. Saccardo & A.N. Berlese)
A.N. Berlese. On dead leaves of Typha
angustifolia L., Typhaceae. Algeria.

typhiseda P.A. Saccardo & A.N. Berlese forma
sodoloci F. Fautrey in C. Roumeguére, Fungi
Selecti Gallici Exsiccati, Century 54, No.
5357. Anno 1890; Revue Mycologique,

angustifolia L., Typhaceae. France.

uliginosa (W. Phillips & C.B. Plowright) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:47. 1883. =Sphaerella uliginosa W.
Phillips & C.B. Plowright, Grevillea, London
10:74. 1881. On leaves of Stellaria uliginosa
Murray, Caryophyllaceae. Great Britain.

ulmicola C. Massa, Annales Mycologici, Berlin
10:287. 1912. On leaves of Ulmus campestris
auct., Ulmaceae. Italy.

umbilicariae (L. Lindsay) P.A. Saccardo & D.
Saccardo in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 17:731. 1905. =Microthelia
umbilicariae W.L. Lindsay, Proceedings of
the Royal Society of Edinburgh 6:535. 1869.
Nom. nud Art. 32.1c; Transactions of the
Royal Society of Edinburgh 25(2):538-539.
1869. Parasitic on sterile thallus of Umbili-
carta pustulata (L.) Hoffm., Lichenes
(Umbilicariaceae). Great Britain.

umbrosa G. Niessl v. Mayendorf in G.L. Rabenhorst,

Fungi Europaei Exsiccati, Klotzschii Herbarii
vivi Mycologici Continuatio, Edition 3 (Edita
Nova), Series 2, Century 20, No. 1934. Anno
1875; Just’s Botanisch Jahresberichte, Berlin
3:262. 1887. =Massaria umbrosa (G. Niessl
v. Mayendorf) H. Rehm in P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 9:761.
1891. On unknown substrate of Actaea
spicata L., Astrantia major L., Gentiana
asclepiadea L., Solanum dulcamara L.,
Spiraea aruncus L. (=Aruncus dioicus Fern.),
Ranunculaceae, Umbelliferae, Gentianaceae,
Solanaceae, Rosaceae. Austria.

uncinata (G. Niessl vy. Mayendorf) ex P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 9:798.
1881. =Heptameria uncinata G. Niessl v.

Vol. 34 Art. 3

Mayendorf in H. Rehm, Ascomyceten,
Fascicle 15, No. 735. Anno 1883. Nom. nud.
Art. 32.1b [as (Niessl) Rehm]. On stems of
Artemisia vulgaris L., Compositae. Czecho-
slovakia.

usneae N.W. Woronichin, Trudy Akademiia Nauk
SSSR Botanicheskii Muzei 21:126. 1927. On
thallus of Usnea florida (L.) Web., Lichenes
(Usneaceae). U.S.S.R.

utahensis J.B. Ellis & B.M. Everhart, The North
American Pyrenomycetes. A Contribution to
Mycologic Botany, p. 361. 1892. On dead
stems of unknown host of Umbelliferae,
Umbelliferae. U.S.A.

vagabunda P.A. Saccardo, Nuovo Giomale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:318. 1875. =Sphaeria
fuscella P.A. Saccardo, Atti dell’ Accademia
Scientifica Veneto-Trentino-Istriana, Padova
2(2):145. 1873 [as Sphaeria fuscella M.J.
Berkeley & C.E. Broome. Non Leptosphaeria
fuscella (M.J. Berkeley & C.E. Broome) V.
Cesati & G. de Notaris]. On branches of
Abies excelsa (Lam.) Poiret, Acer campestre
L., Alnus glutinosa (L.) Gaertner, Ampelopsis
heterophylla (Thunb.) Sieb. & Zucc.,
Clematis vitalba L., Cornus sanguinea L.,
Corylus avellana L., Dulcamara sp..
Hypericum calycinum L., Kerria japonica (L.)
DC., Quercus pedunculata Ehrh., Salix
purpurea L., Pinaceae, Aceraceae, Betu-
laceae, Vitaceae, Ranunculaceae, Comaceae,
Solanaceae, Guttiferae, Rosaceae, Fagaceae,
Salicaceae. Italy.

vagabunda P.A. Saccardo forma abietis F. Fautrey in
C. Roumeguére, Fungi Selecti Gallici
Exsiccati, Century 59, No. 5846. Anno 1891;
Revue Mycologique, Toulouse 13:167. 1891.
On stems of Abies excelsa (Lam.) Poiret,
Pinaceae. France.

vagabunda P.A. Saccardo forma citri-limonii P.A.
Saccardo, Fungi Italici autographice delineati
(additis nonnullis extra-italicis asterisco
notatis), Patavii, No. 520. 1879. On stems of
Citrus limonum Risso, Rutaceae. Italy.

vagabunda P.A. Saccardo forma daphnes? M. de
Sousa da Camara, Agronomia Lusitana,
Sacavém 11:172. 1949. On branches of
Daphne gnidium L., Thymelaeaceae.
Portugal.

vagabunda P.A. Saccardo forma lonicerae F.
Fautrey in C. Roumeguére, Fungi Selecti
Gallici Exsiccati, Century 61, No. 6040. Anno
1892; Revue Mycologique, Toulouse 14:108.
1892. On young shoots of Periclymenwm sp.
(=Lonicera sp.), Caprifoliaceae. France.

vagabunda P.A. Saccardo forma salicis-capreae F.
Fautrey in C. Roumeguére, Fungi Selecti
Gallici Exsiccati, Century 65, No. 6440. Anno
1894; Revue Mycologique, Toulouse 16:8.

March 1991

1894. On stems of Salix caprea L., Sali-
caceae. France.
vagabunda P.A. Saccardo subsp. alvarensis K.
Starbiack, Bihang till K. Svenska Vetenskap-
sakademiens Handlingar, Stockholm, Series
15(2):13. 1889. On rhizomes on uncovered
soil of Plantago maritima L., Plantaginaceae.
Sweden.
vagabunda P.A. Saccardo var. caulium P.A.
Saccardo, Revue Mycologique, Toulouse
3:44. 1881; Reliquiae Mycologicae Liber-
tianae, Series Altera Reviserunt C.
Roumeguére & P.A. Saccardo, Toulouse, p. 6,
No. 119. 1881. On stems of Perasites sp.,
Compositae. France.
vagabunda P.A. Saccardo var. divergens P.A.
Saccardo, Fungi Italici autographice delineati
(additis nonnullis extra-italicis asterisco
notatis), Patavii, No. 518. 1879; Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:31. 1883. On
branches of Kerria sp., Rosaceae. Italy.
vagabunda P.A. Saccardo var. dulcamarae P.A.
Karsten, Meddelanden af Societas pro Fauna
et Flora Fennica, Helsingfors 16:34. 1888;
Symbolae ad Mycologiam Fennicam
XXVII:34. 1888. On dead stems of Solanum
dulcamara L., Solanaceae. Finland.
vagabunda P.A. Saccardo var. sarmenti P.A.
Saccardo, Fungi Italici autographice delineati
(additis nonnullis extra-italicis asterisco
notatis), Patavii, No. 518. 1879; Sylloge
Fungorum Omnium Hucusque Congitorum
Digessit P.A. Saccardo 2:31. 1883. On
runners of Rubus sp., Rosaceae. Italy.
vagans P.A. Karsten, Svenska Vetenskapsakademien
Stockholm Ofversigt af Forhandlingar,
Stockholm 29(2):101. 1872. On rotting leaves
of Dupontia fisheri R. Br., Gramineae.
Norway.
vagans P.A. Karsten forma scirpi P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:59.
1883. On leaves of Dupontia fisheri R. Br.,
Gramineae. France, Norway.
vaginae G. Passerini, Atti dell’ Reale Accademia
Nazionale dei Lincei. Rendiconti, Rome,
Series 4, 7(2):45. 1891. On wet sheaths of
Phragmites communis Trin., Gramineae.
Italy.
vahlii E. Rostrup, Meddelelser om Grgnland,
Kj@benhavn 3:557—558. 1888. On dry stems
of Melandrium triflorum J. Vahl. ex Liebm.,
Caryophyllaceae. Greenland.
valdiviensis C.L. Spegazzini, Fungi Chilenses, pp.
80-81. 1910; Revista de La Facultad de
Agronomia Y Veterinaria, Universidad
Nacional de La Plata, Series 2, 6:80-81. 1910.

3

Ys

Nomenclator of Leptosphaeria

287

On decaying shoots of Digitalis purpurea L.,
Scrophulariaceae. Chile.
valdobbiae T. Ferraris, Malpighia. Rassegna
Mensuale di Botanica. Messina, Genova
18:488-489. 1904. On leaves of Fagus
sylvatica L., Fagaceae. Italy.
valesiaca H. Wegelin, Mitteilungen der Thurgauis-
chen Naturforschenden Gesellschaft,
Frauenfeld 12:175. 1896. On preceding
year’s stems of Artemisia campestris L.,
Compositae. Switzerland.
vanhoeffeniana A. Allescher in A. Allescher and
P.C. Hennings, Biblotheca Botanica, Stuttgart
42(1):47. 1897. [Issued also in Botanische
Ergebnisse der von der Gesellschaft fiir
Erdkunde zu Berlin unter Leitung Dr. v.
Drygalski’s ausgesandten Grénlandexpedition
nach Dr. Vanhoffen’s Sammlungen Bearbe-
itet, A. Kryptogamen...Stuttgart, in 1897.]
Non vidi. On dead leaves of Melandrium
apetalum (L.) Fenzl, Caryophyllaceae.
Greenland.
variabilis L.M. Unamuno Yrigoyen, Anales del
Jardin Botanico de Madrid 1:29-31. 1941.
On culms of Juncus sp., Juncaceae. Spain.
variegata C.H. Peck, Bulletin of the New York State
Museum, Albany 67:31—32. 1903. On dead
stems of Phytolacca americana L., Phytolac-
caceae. U.S.A.
variiseptata G.L. Stout, Mycologia, Lancaster,
Pennsylvania 22:276—-277. 1930.
=Phaeosphaeria variiseptata (G.L. Stout)
R.A. Shoemaker, Canadian Journal of Botany,
Ottawa 67:1536. 1989. On leaves of Zea
mays L., Gramineae. U.S.A.
vectis (M.J. Berkeley & C.E. Broome) V. Cesati &
G. de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:236. 1863.
=Sphaeria (Caulicolae) vectis M.J. Berkeley
& C.E. Broome, Annals and Magazine of
Natural History, London, Series 2, 13:467.
1854. =Paraphaeosphaeria vectis (M.J.
Berkeley & C.E. Broome) A. Hedjaroude,
Sydowia. Annales Mycologici, Horn, N.O.
22(1-4):98. (1968) 1969. On dead leaves of
Iris foetidissima L., Iridaceae. Great Britain.
veratri F.S. Earle in E.L. Greene, Plantae Bakerianae
2:20-21. 1901. On dead weathered stems of
Veratrum sp., Liliaceae. U.S.A.
verwoerdiana §.J. DuPlessis, South African Journal
of Science, Cape Town 30:207—208. 1933.
On leaves of Aloe affinis A. Berger, Aloe
striatula Haw., Liliaceae. South Africa.
viciae E. Miiller, Sydowia. Annales Mycologici,
Horn, N.O. 4(1-6):268. 1950. On dead stems
of Vicia cracca L., Leguminosae. Switzer-
land.
vincae (E.M. Fries) P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in

288

Primis Italicos Ilustrans 1:37. 1877.
=Sphaeria vincae E.M. Fries in J.E. Duby,
Aug. Pyrami de Candolle Botanicon Gallicum
seu synopsis plantarum in flora gallica
descriptum. Pars Secunda, p. 709. 1830.
=Sphaerella vincae (E.M. Fries) B. Auerswald
in W. Gonnermann and G.L. Rabenhorst,
Mycologia Europaea, Abbildungen Sammtli-
cher Pilze Europa’s, Hefte 5 und 6, Synopsis
Pyrenomycetum Europaeorum, p. 10. 1869.
=Metasphaeria vincae (E.M. Fries) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:171. 1883. On dry leaves of Vinca minor
L., Apocynaceae. France, Germany, Italy.

vindobonensis F. Petrak, Annales Mycologici, Berlin
42:75. 1944. On dead stems of Cynoglossum
officinale L., Boraginaceae. Austria.

vinealis G. Passerini in F. v. Thiimen, Die Pilze des
Weinstockes, pp. 133-134. 1878. On dry
runners of Vitis vinifera L., Vitaceae. Italy.

vinosa C.L. Spegazzini, Anales de la Sociedad
Cientifica Argentina, Buenos Aires 10:142.
1880. On decaying leaves and stems of
Eryngium agavifolilum Griseb., Umbelliferae.
Argentina.

virginica (M.C. Cooke & J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:35.
1883. =Sphaeria (Caulicolae) virginica M.C.
Cooke & J.B. Ellis, Grevillea, London 8:16.
1879. On stems of Lepidium virginicum L.,
Cruciferae. U.S.A.

viridella (C.H. Peck) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:18. 1883.
=Sphaeria (Caulicolae) viridella C.H. Peck,
Report. New York State Museum of Natural
History, Albany 30:66. 1878. =Phaeo-
sphaeria viridella (C.H. Peck) A. Leucht-
mann, Sydowia. Annales Mycologici, Horn,
N.O. 37:108. 1984. On dead stems of Juncus
effusus L., Melilotus sp., Cyperaceae,
Leguminosae. U.S.A.

vitalbae G. Niessl v. Mayendorf in H. Rehm, As-
comyceten, Fascicle 19, No. 938. Anno 1888;
Hedwigia, Dresden 27:172. 1888. On
decorticated twigs of Clematis vitalba L.,
Ranunculaceae. Austria.

vitalbae (G. de Notaris ex V. Cesati & G. de Notaris)
H.G. Winter in J. Kunze, Fungi Selecti
Exsiccati, Century 3, No. 331. Anno 1880.
(=Fungi Helvetica, No. 31.) =Sphaeria
vitalbae G. de Notaris, Memorie della
Accademia delle Scienze di Torino, Series 2,
13:124. 1853. Nom. nud. Art. 32.1; Micro-
mycetes Italici Novi vel minus Cogniti Decas
8, p. 124. 1854; Commentario della Societa

Illinois Natural History Survey Bulletin

Vol. 34 Art. 3

Crittogamologia Italiana, Milan 1:221—222.
1863. On dry runners of Clematis vitalba L.,
Ranunculaceae. Switzerland.

vitalbae G. Niessl v. Mayendorf var. sarmenticola J.
Feltgen, Vorstudien zu einer Pilz-Flora des
Grossherzogthums, Luxemburg 1(3):221—222.
1903. On dry shoots of Clematis vitalba L.,
Ranunculaceae. Luxembourg.

vitensis L.M. Unamuno Yrigoyen, Boletin de la R.
Sociedad Espanola de Historia Natural,
Madrid 29:394-395. 1929. On above-ground
parts of reeds, bracts, and peduncles of
inflorescence of Juncus glaucus Sibth.,
Juncaceae. Spain.

viticola F. Fautrey & C. Roumeguere in C.
Roumegueére, Revue Mycologique, Toulouse
14:6. 1892; Fungi Selecti Gallici Exsiccati,
Century 60, No. 5950. Anno 1892. On vines
of Vitis vinifera L., Vitaceae. France.

vitigena P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:29. 1883. =Sphaerella vitis S. Schulzer v.
Miiggenburg, Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien 20:643.
1870. Non Leptosphaeria vitis (J.L. Cas-
tagne) R. Pirotta 1882, non Leptosphaeria
vitis S. Schuizer v. Miiggenburg, 1870.
=Sphaerella sarmentorum R. Pirotta, fide P.A.
Saccardo, 1883. On living branches of Vitis
vinifera L., Vitaceae. Austria.

vitis (J.L. Castagne) R. Pirotta, Archivo del Labora-
torio di Botanica Crittogamica Universita di
Pavia 2 & 3:161. (1877) 1879. Nom. illegit.
Art. 63.1. =Sphaeria vitis J.L. Castagne,
Catalogue Des Plantes qui Croissent
Naturellement aux Environs de Marseille, p.
166. 1845. Nom. illegit. Art. 64.1. On dried
vines of Vitis vinifera L., Vitaceae. France.

vitis S. Schulzer v. Miiggenburg, Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wien
20:642-643. 1870. On vines of Vitis vinifera
L., Vitaceae. Austria.

volkartiana E. Miiller, Sydowia. Annales Mycol-
ogici, Horn, N.O. 4(1-6):210. 1950.
=Phaeosphaeria volkartiana (E. Miiller) A.
Hedjaroude, Sydowia. Annales Mycologici,
Horn, N.O. 22(1-4):84. (1968) 1969. On
dead culms of Trisetwm distichophyllum
(Vill.) Beauv., Tristeum spicatum (L.) K.
Richter, Gramineae. Switzerland.

vrieseae W. Siemaszko, Acta Societatis Botani-
corum Poloniae, Warszawa 1:21—22. 1923.
On leaves of Vriesea sp., Bromeliaceae.
U.S.S.R.

waghorniana H. Rehm, Hedwigia, Dresden 39:324.
1900. On cortex? of Betula sp., Betulaceae.
Canada.

weberi C.A. Oudemans, K. Akademie van Weten-
schappen Amsterdam, Afdeeling Natuurkund

March 1991

Verslagen en Mededeelingen 3 Reeks, Deel
2:156. 1885. On leaves of Ranunculus nivalis
L. var. sulphureus Wahlenberg, Ranuncu-
laceae. U.S.S.R.

weddellii (J.P. Montagne) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:86. 1883.
=Sphaeria weddellii J.P. Montagne, Annales
des Sciences Naturelles, Paris, Botanique,
Series 4, 5:372-373. 1856. On culms of
Bambusa sp., Gramineae. Bolivia.

wegeliniana P.A. Saccardo & P. Sydow, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 14:567. 1899.
=Leptosphaeria rumicis H. Wegelin. Nec F.
Fautrey. On dry stems of Rwmex sp.,
Polygonaceae. Switzerland.

wegeliniana P.A. Saccardo & P. Sydow forma
teucrii J. Feltgen, Vorstudien zu einer Pilz-
Flora des Grossherzogthums, Luxemburg
1(3):222. 1903. On dry stems of Teucrium
scorodonia L., Labiatae. Luxembourg.

wehmeyeri R.A. Shoemaker, Canadian Journal of
Botany, Ottawa 62:2725. 1984. On stems of
Lupinus rubricaulis Greene, L. latifolius J.G.
Agardh, Thermopsis montana Nutt., Legumi-
nosae. Canada, U.S.A.

williamsii C.G. Hansford, Proceedings of the
Linnean Society of New South Wales, Sydney
82:216. 1957. On dead leaves of Dianella
revoluta R. Br., Liliaceae. Australia.

winteri G. Niessl v. Mayendorf in H.G. Winter,
Hedwigia, Dresden 22:1. 1883. =Sclero-
pleella winteri (G. Niessl v. Mayendorf) F. v.
Hohnel, Annales Mycologici, Berlin 18:76.
1920. Nom. inval. Art. 33.1. [Combination
not formally proposed.] =Wertsteinina winteri
(G. Niessl v. Mayendorf) E. Miiller, Sydowia.
Annales Mycologici, Horn, N.O. 4(1—-6):203.
1950. On dry leaves of Plantago alpina L.,
Plantaginaceae. Germany.

woodrow-wilsonii [as Woodrowi Wilsoni] L.
Garbowski, Bulletin Trimestriel de la Société
Mycologique de France, Paris 39:238. 1924.
On soft parts of dried leaves of Eryngium
campestre L., Umbelliferae. U.S.S.R.
(Crimea).

woroninii E. Docea & A. Negru in A. Negru, E.
Docea, and E. Szasz, Novitates Systematicae
Plantarum Non Vascularium (=Novosti
Sistematiki Nizshikh Rastenij, Novitates
Systematicae Plantarum Non Vascularium)
9:168. 1972. On seeds of Cannabis sativa L.,
Cannabaceae. Romania.

xerophylli J.B. Ellis, American Naturalist, Lancaster,
Pennsylvania 17(1):316—317. 1883. =Metas-
phaeria xerophylli (J.B. Ellis) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2(Ad-
denda Ad Volumen Secundem):LXI. 1883;

Nomenclator of Leptosphaeria

289

Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 9:829.
1891. On dead leaves of Xerophyllum
asphodeloides (L.) Nutt., Liliaceae. U.S.A.

xiphii G. Passerini, Rendiconti della Sedute della R.
Accademia dei Lincei, Classe di Scienze
Fisiche, Matematiche e Natural (=Atti dell’
Accademia Nazionale dei Lincei, Rendiconti,
Roma), Series 4, 3:90-91. 1887. On wilted
leaves of Iris foetidissima L., Iridaceae. Italy.

xylogena M. Curzi & M. Barbani, Atti dell’ Istituto
Botanico della Universita e Laboratorio
Crittogamica di Pavia, Milano, Series 3,
3:160. 1927. On decorticated branches of
Salix sp., Salicaceae. Italy.

yerbae C.L. Spegazzini, Anales del Museo Nacional
de Historia Natural de Buenos Aires 17(Series
3, 10):126-127. 1908; Hongos de La Yerba
Mate, No. 32. 1908. On wilting and dead
branches of //ex paraguariensis A. St. Hil.,
Aquifoliaceae. Argentina.

yulan P.A. Saccardo, Nuovo Giornale Botanico
Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:312. 1875. On wilted
leaves of Magnolia yulan Desf., Magnoli-
aceae. Italy.

zahlbruckneri P. Strasser, Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wien
57:315-316. 1907 [as Leptosphaeria
(Pocosphaeria Sacc.)]. =Acanthostigmella
zahlbruckneri (P. Strasser) F. v. Hohnel,
Sitzungsberichte der Akademie der Wissen-
schaften in Wien, Mathematisch-
naturwissenschaftliche Klasse, Abt. I,
118:1503. 1909. On dry stems of Mentha
sylvestris L. [=Mentha longifolia (L.) Huds.],
Labiatae. Austria.

zeae G.L. Stout. Mycologia, Lancaster, Pennsylvania
22:277. 1930. On leaves of Zea mays L.,
Gramineae. U.S.A. (Illinois).

zeae-maydis A.M. Saccas, Revue de Pathologie
Végetale et D’Entomologie Agricole de
France, Paris 30:179-180. 1951. On dead
leaves of Zea mays L., Gramineae. French
Equatorial Africa.

zeicola A.M. Saccas, Revue de Pathologie Végétale
et D’Entomologie Agricole de France, Paris
30:176-177. 1951. On living leaves of Zea
mays L., Gramineae. French Equatorial
Africa.

zingiberis (K. Sawada) V. Dhar, L.S. Srivastava &
H.S. Sahambi, Current Science, Bangalore
50:540. 1981. =Phaeosphaeria zingiberis K.
Sawada, Descriptive Catalogue of Taiwan
(Formosan) Fungi, Part XI. (Special Publica-
tion of the National Taiwan University
Taipaei No. 8), p. 67. 1959. On leaves of
Zingiber officinale Rosc., Zingiberaceae.
India.

Illinois Natural History Survey Bulletin

zizaniaecola (M.J. Berkeley & M.A. Curtis) P.A.

Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:86. 1883. =Sphaeria zizaniaecola M.J.
Berkeley & M.A. Curtis, Grevillea,

London 4:145. 1876. On stems of Zizania sp.,
Gramineae. U.S.A.

Vol. 34 Art. 3

zizannivora K. Hara, A List of Japanese Fungi

Hitherto Known, p. 402(2). 1954 [as ziza-
ninaevora). [There are two pages numbered
402; the description is on the second page.]
On living plant of Zizania latifolia Turcz.,
Gramineae. Japan.

Host Index

Abies balsamea Miller
L. faulii
Abies excelsa (Lam.) Poiret
L. vagabunda
L. vagabunda forma abietis
Abutilon avicennae Gaertner
L. abutilonis
Abutilon indicum (L.) Sweet
L. abutilonis
Acacia kempeana F. Muell.
L. clelandii
Acantholimon acerosum Boiss.
L. kotschyana
Acantholimon melananthum
Boiss.
L. kotschyana
Acer campestre L.
L. vagabunda
Acer laetum C.A. Mey.
L. aceris
Acer negundo L.
L. diana
Acer pseudoplatanus L.
L. dioica
L. obesula
Acer saccharum auct.
L. inquinans
Acer sp.
L. controversa
L. leucoplaca
L. muelleri
Achillea millefolium auct.
L. achilleae
L. compressa
L. dolioloides
L. millefolii
L. ogilviensis forma
achilleae
L. staritzii
Achillea nana L.
L. nanae
Achillea ptarmica L.
L. marginalis
L. passerinii
L. ptarmicae
Achillea sp.
L. doliolum
L. tanaceti
Aconitum compactum Rchb.
L. nigromaculata

Aconitum lycoctonum L.

L. lasiosphaerioides
Aconitum nappelus L.

L. aconiti

L. anthostomoides

L. napelli

L. nigromaculata
Aconitum paniculatum Lam.

L. nigromaculata
Aconitum sp.

L. scotophila
Acorus calamus L.

. acorella

acori

densa

. microscopica subsp.
calami

L. typharum forma acori
Actaea spicata L.

L. umbrosa
Adenostyles albifrons (L. fil.)
Reichenb.

L. nitschkei forma

adenostylidis
Adesmia sp.

L. adesmicola
Adhatoda sp.

L. acanthi
Adonis pyrenaica DC.

L. doliolum var. pachy-

spora

booo

Aeluropus littoralis (Gouan.) Parl.

L. aeluropodis
Agastache urticifolia (Benth.)
Ktze.

L. brightonensis

L. darkeri
Agave ferox C. Koch

L. obtusispora forma

agaves
Agave rigida Mill. var. sisalana
(Perrine) Engelm.

L. agaves
Agave striata Zucc.

L. obtusispora
Agropyron repens (L.) Beauv.

L. anisomeres

L. avenaria

L. linearis

Agrostis vulgaris With.
L. poae var. agrostidis
L. rhizomatum
Agrostis sp.
L. sparsa
Ailanthus glandulosa Desf.
L. ailanthi
L. endophaena
L. glandulosae
Aira alpina L.
L. insignis
Aira cespitosa L.
L. anarithma
L. insignis forma airae-
cespitosae
L. lineolaris
L. personata
L. pleurospora
L. quinta
Albizia julibrissin Durz.
L. marginalis
Algae
L. fluviatilis
L. lemaneae
L. marina
L. mirabilis
L. mirandae
Alhagi sp.
L. alhagii
Alisma plantago L.
L. rivularis
Alliaria officinalis Andrz ex Bieb.
L. alliariae
Allium validum §. Wats.
L. lassenensis
Alnus glutinosa (L.) Gaertner
L. coniothyrium
L. vagabunda
Alnus sp.
L. leucoplaca
L. lonicerina
Aloe affinis A. Berger
L. verwoerdiana
Aloe arborescens Miller
L. aloes
Aloe striatula Haw.
L. verwoerdiana
Alopecurus borealis Trin.
L. alopecuri

Alopecurus ovatus Knapp

L. insignis

L. microscopica
Aloysia citriodora Ort. ex Pers.

L. octophragmia
Alpinia speciosa K. Sch.

L. alpiniae
Amaranthus sp.

L. eriophora
Ambrosia trifida L.

L. drechsleri

L. fulgida
Ambrosia sp.

L. doliolum
Ammophila arenaria (L.) Link

L. ammophilae

L. ammophilae forma

calamagrostidis-
arenariae

L. littoralis

L. perforans
Ammophila arundinacea Host

L. sabuletorum
Ammophila sp.

L. marram
Ammothamnus lehmannii Bunge

L. ammothamni
Amorpha fruticosa L.

L. amorphae
Ampelopsis heterophylla (Thunb.)
Sieb. & Zucc.

L. vagabunda
Anacylus radiatus Loisel.

L. anacycli
Anaphalis sp.

L. doliolum
Anarrhinum bellidifolium (L.)
Willd.

L. anarrhini
Andromeda tetragona L.

L. andromedae

L. hyperborea
Andromeda sp.

L. sublanosa
Andropogon glomeratus (Walt.)
B.S.P.

L. subcompressa
Andropogon ischaemum L.

L. eustomoides

L. ischaemi

L. trimera
Andropogon muricatus Retz.

L. muricata
Andropogon saccharoides Swartz

L. puiggarii
Andropogon sp.

L. herpotrichoides

L. latebrosa

L. michotii
Androsace lactea L.

L. pachyasca

Illinois Natural History Survey Bulletin

Anemone narcissiflora L.

L. raphidophora
Anemone sylvestris L.

L. anemones
Anemone virginiana L.

L. platypus
Angelica sylvestris L.

L. conoidea

L. conoidea forma

angelicae
Angelica sp.

L. obesula

L. setosa
Anthemis tinctoria L.

L. dolioloides

L. tanaceti
Anthericum ramosum L.

L. antherici
Anthriscus sylvestris (L.) Hoffm.

L. dolioloides var. inops
Antirrhinum majus L.

L. thuemeniana
Antirrhinum siculum Miller

L. galiorum subsp.

antirrhini

L. insulana
Apios fortunei Maxim.

L. apios

L. apios-fortunei
Apocynum sp.

L. doliolum
Aquilegia vulgaris L.

L. aquilegiae
Arabis alpina L.

L. arabidis

L. johansonii
Aralia sp.

L. doliolum
Araucaria imbricata Pav.

L. californica
Arbutus unedo L.

L. arbuti
Arctium sp.

L. doliolum
Areca sapida Soland. ex Hook. f.

L. arecae
Arenaria ciliata L.

L. auerswaldii
Armeria vulgaris Willd.

L. staritzii
Arrhenatherum avenaceum
Beauv.

L. arrhenatheri
Arrhenatherum elatius (L.)
Beauv. ex J. Presl. & C. Presl.

L. arrhenatheri
Artemisia absinthium L.

L. gloeospora
Artemisia annua L.

L. sydowiana

Vol. 34 Ar. 3

Artemisia austriaca Jacq.
L. compressa
Artemisia californica Less.
L. tumefaciens
Artemisia campestris L.
. artemisiae
. Caespitosa
. compressa
. phaeospora
. sydowiana
. valesiaca
Artemisia camphorata Vill.
L. camphorata
Artemisia frigida Willd.
L. tetonensis
Artemisia herba-alba Asso
L. rothomagensis var.
artemisiae
Artemisia tridentata Nutt.
L. lasioderma
Artemisia vulgaris L.
L. doliolum var. subdisti-
cha
. grammodes
. hispanica
. kalmusii
. purpurea
. uncinata
Artemisia sp.
L. cervispora
L. crustacea
L. owaniae
L. tumefaciens
Aruncus dioicus Ferm.
L. umbrosa
Aruncus silvestris Kostel.
L. arunci
Arundinaria sp.
L. eumorpha
Arundo donax L.
L. donacina
L. recessa
L. rhodophaea
Asclepias syriaca L.
L. dearnessii
Asclepias sp.
L. doliolum
L. russellii
Asparagus officinalis L.
. asparagi
. asparagina
. passeriniana
. portoricensis
. praeclara
. punctoidea
L. socialis
Asparagus sp.
L. comatella
L. indica
Asperella japonica Hack.
L. asperellae

Teall alll Soll all alll

alll Sal wall ool So

‘eal sll soll wall nl oo

March 1991

Asperula sp.
L. politis
Aspicilia calcarea (L.) Korb
L. crozalsti
Aspidistra elatior Blume
L. aspidistrae
Asplenium septentrionale (L.)
Hoffm.
L. asplenii
Aster multiflorus Ait.
L. astericola
Aster sagittifolius Wedem. ex
Willd.
L. asteris
Aster salignus Willd.
L. conoidea forma asteris
Aster sp.
L. doliolum
L. heliopsidis
Astrantia major L.
L. umbrosa
Atraphaxis muschketovii Krasnov
L. atraphaxidis
Atriplex hortensis L.
L. calvescens
Atriplex verrucifera Bieb.
L. atriplicis
Atriplex sp.
L. echinella
Avena sativa L.
L. korrae
Avena sp.
L. avenae
Avicennia marina (Forsk.) Vierh.
var. resiniferae (Forst.) Bakh.
L. australiensis
Avicennia nitida Jacq.
L. avicenniae
Axonopsus compressus (Swartz)
Beauv.
L. korrae
Azalea sp. (=Rhododendron sp.)
L. azaleae
Baccharis sp.
L. bicuspidata
Baeomyces rufus (Huds.) Rebent.
L. baeomycearia
L. neottizans
Baldingera arundinacea (L.)
Dumort.
L. baldingerae
L. larseniana
Ballota acetabulosa (L.) Bentham
L. ballotae :
Ballota nigra L.
L. slovacica
Bambusa sp.
L. amphiloga
L. bambusae
L. bambusicola

Nomenclator of Leptosphaeria

L. scabrispora

L. schneideriana

L. tigrisoides

L. weddellii
Baptisia sp.

L. comatella
Bardana sp.

L. bardanae
Barkhousia taraxacifolia (Thuill.)
DC.

L. modesta var. rubellula
Berberis ilicifolia Forst.

L. berberidicola
Berberis petiolicola Wall.

L. punjabensis
Berberis vulgaris L.

L. berberidis forma

berberidis

L. coniothyrium
Berberis sp.

L. inconspicua
Berteroa incana (L.) DC.

L. submaculans
Betula verrucosa Ehrh.

L. betulina
Betula sp.

L. betulina

L. waghorniana
Biebersteinia emodii Jaub. &
Spach.

L. hollosiana
Blysmus compressus (L.) Panzer
ex Link

L. petkovicensis var. elymi
Bocconia sp.

L. bocconiae
Bomarea caldasii (HBK) Asch. &
Graebn.

L. bomareae
Borreria leucomela (L.) Mudd

L. leucomelaria
Brachypodium sylvaticum
(Hudson) Beauv.

L. brachypodii

L. microscopica forma

brachypodii

L. sylvatica
Brachypodium sp.

L. culmifraga var. linearis
Brassica campestris L.

L. alliariae

L. maculans
Brassica crispa Rafin.

L. salebrosa
Brassica napus L.

L. napi
Brassica oleifera Moench

L. maculans forma

denudata
Brassica oleracea L.
L. cesatiana

293

Brassica rapa L.

L. nigrella
Brassica sp.

L. olericola
Braya sp.

L. norvegica
Briza media L.

L. brizae

L. media
Bromus asper Murray

L. culmifraga var.

bromicola

Bromus inermis Leysser

L. avenaria
Broussonetia sp.

L. coniothyrium
Buddleja davidii Franch.

L. buddlejae

L. davidii

L. polini
Buphthalmum salicifolium L.

L. septemcellulata
Bupleurum falcatum L.

L. agnita var. bupleuri
Bupleurum petraeum L.

L. bupleuri
Bupleurum ranunculoides L.

L. ranunculoides
Buxus sempervirens L.

L. buxina

L. melanommoides

L. revocans
Cacalia hastata L.

L. doliolum var. cacaliae
Cacalia sp.

L. nitschkei
Calamogrostis epigejos (L.) Roth

L. culmorum forma epigeii
Calamagrostis montana Host

L. fuckelii
Calamagrostis sylvatica Host

L. quinta
Calamagrostis sp.

L. coccodes

L. graminum

L. intermedia

L. sparsa
Calligonum sp.

L. calligoni
Calluna vulgaris (L.) Hull

L. ericae
Calopogonium mucunoides Desv.

L. calopogonii
Camellia japonica L.

L. camelliae-japonicae
Camellia theifera Griff.

L. depressa

L. tornatospora
Camellia sp.

L. camelliae

294

Campanula zoysti Wulfen

L. pachyasca

L. plemeliana
Camphorosma monspeliaca L.

L. serbica
Camphorosma perennis Pall.

L. camphorosmae
Canna indica L.

L. cannae
Cannabis sativa L.

L. cannabina

L. woroninii
Capparis aphylla Roth

L. ahmadii

L. capparidicola
Capparis horrida L.

L. simillima
Capparis spinosa L.

L. capparidis
Cardamine sylvatica Link

L. trollii
Carduus carlinoides Gouan

L. carlinoides
Carduus defloratus L.

L. centaureae
Carduus nutans L.

L. carduina
Carduus scopulorum Greene

L. chrysanthemi
Carduus sp.

L. carduorum
Carex acutiformis Ehrh.

L. sparsa
Carex arenaria L.

L. cariciphila
Carex baldensis L.

L. kochiana
Carex firma Host

L. caricis-firmae
Carex folliculata L.

L. folliculata
Carex glauca Scop.

L. striolata var. caricis-

glaucae

Carex gracillima Schwein.

L. folliculata var. oxyspora
Carex hirta L.

L. caricis

L. occulta
Carex hyperborea Drejer

L. macrotheca
Carex leporina L.

L. culmorum var. paleicola
Carex paludosa Good.

L. gigaspsora

L. puccinioides

L. rivalis
Carex panicea L.

L. caricina
Carex pendula Hudson

L. caricicola

L. caricis

Illinois Natural History Survey Bulletin

Carex pulla Good.

L. caricinella

L. consobrina
Carex rigida Good.

L. macrotheca
Carex riparia Curtis

L. caricicola
Carex silvatica auct.

L. caricina
Carex vaginata Tausch

L. caricis
Carex vesicaria L.

L. paludosa
Carex vulpina L.

L. caricis-vulpinae

L. microscopica

L. microscopica var.

caricis-vulpinae
Carex sp.

L. clara

L. cumana

L. epicarecta

L. hemicrypta
Carlina acaulis L.

L. anthophila

L. cynaracearum
Carlina vulgaris L.

L. doliolum

L. doliolum forma

carlinae-vulgaris
Carpinus sp.

L. depressa

L. subsimilis
Carya sp.

L. cacuminispora

L. lejostega

L. leucoplaca
Cassia sp.

L. cassiaecola
Cassinia aculeata R. Br.

L. plagia
Castanea sp.

L. involucralis
Castilleja miniata Doug. ex
Hook.

L. castillejae

L. concinna
Castilleja pallida (L.) Sprengel

L. castilleiae
Catabrosa algida Fr.

L. algida
Ceanothus sp.

L. ceanothi
Cecropia peltata L.

L. cecropiae
Centaurea jacea L.

L. jaceae
Centaurea scabiosa L.

L. centaureae
Centranthus sp.

L. galiicola var. brachy-

spora

Vol. 34 Art. 3

Cephalaria uralensis (Murray)
Roemer & Schultes

L. cephalariai-uralensis

L. taurica
Cerastium biebersteinii DC.

L. biebersteinii
Cercocarpus ledifolius Nutt. ex
Torr. & Gray

L. cercocarpi
Cereus peruvianus (L.) Miller

L. cerei-peruviani
Chaerophyllum aureum L.

L. agnita var. major
Chamaerops excelsior Boj

L. spatharum
Chamaerops humilis L.

L. algarbiensis

L. chamaeropis

L. debeauxii

L. magnusiana
Chamaerops sp.

L. pruni forma plurivora
Cheiranthus annuus L.

L. impressa
Chelidonium majus L.

L. chelidonii
Chenopodium album L.

L. calvescens

L. chenopodii-albi

L. eutypoides

L. inculta

L. nigricans
Chenopodium anthelminticum
Bert. ex Steud.

L. anthelmintica
Chenopodium sp.

L. eriophora
Chimonanthus sp.

L. coniothyrium
Chondrilla juncea L.

L. bella

L. eburnea

L. mirabilis

L. trichostoma
Chondrus crispus J. Stackhouse

L. chondri

L. danica

L. marina
Chrysanthemum corymbosum L.

L. agnita var. chrysanthemi
Chrysanthemum leucanthemum L.

L. tanaceti
Chrysanthemum vulgare (L.)
Bernh.

L. dolioloides

L. tanaceti
Chrysothamnus graveolens
(Nutt.) Greene

L. arthrophyma

March 1991

Chusquea serrulata Pilger

L. chusqueae

L. saginata
Chusquea sp.

L. consociata

L. stellata
Cicclidotus fontinaloides (Hedw.)
P. Beauv.

L. cinclidoti
Cinnamomum camphora T. Nees
ex Eberm.

L. cinnamomi
Cinnamomum zeylanicum Blume

L. almeidae
Cirsium altissimum (L.) Spreng.

L. compressa
Cirsium arvense (L.) Scop.

L. cirsii-arvensis
Cirsium lanceolatum (L.) Scop.

L. corynispora
Cirsium muticum Michx.

L. mesoedema
Cirsium palustre (L.) Scop.

L. galiorum forma

cirsiorum
Cirsium sp.

L. dolioloides var. cirsii

L. doliolum
Cistus albidus L.

L. cisti
Cistus ladanifer L.

L. cistina
Cistus monspeliensis L.

L. cisticola
Citrus aurantium L.

L. bondari

L. papulosa
Citrus grandis Osb.

L. bondari
Citrus limonia Osb.

L. bondari
Citrus limonum Risso

L. vagabunda forma citri-

limonii
Citrus medica L.

L. bondari
Citrus sinensis Osb.

L. bondari
Citrus sp.

L. bondari

L. citricola
Cladium articulatum R. Br.

L. aliena
Cladium mariscus (L.) Pohl

L. cladii

L. crucheti
Clematis recta L.

L. aucta
Clematis vitalba L.

L. agminalis

Nomenclator of Leptosphaeria

. agminalis forma minor
. grignonnensis
. haematites
. Incruenta
. nectrioides
. pleosporoides
. pyrenopezizoides
rimalis
. vagabunda
. vitalbae
. vitalbae var. sarmenti-
cola
Clerodendrum foetidi Bunge
L. clerodendri
Coccothrinax argentata (Jacq.)
Bailey
L. coccothrinacis
Cocos romanzoffiana Cham.
L. cocoes
L. molleriana
Coffea arabica L.
L. coffeigena
Coffea canephora Pierre ex
Froehn.
L. canephorae
L. coffeigena var. longiros-
trata
Coffea excelsa Cheval.
L. centrafricana
L. excelsa
L. lobayensis
L. longispora
Coffea robusta L. Linden
. africana
. cylindrospora
. gigaspora
. macrorostra
. oubanguiensis
. tetraspora
Coffea sp.
L. coffaeicida
L. pusilla
L. tonduzi
Coix lacryma-jobi L. var.
susutama Honda
L. coicis
Coleosanthus reniformis (Gray)
Rydb.
L. coleosanthi
Collinsonia canadensis L.
L. collinsoniae
Collomia squarrosa Nutt.
L. collumiae
Colocasia antiquorum Schott
L. colocasiae
Conium maculatum L.
L. conii
L. coniigena
L. rubicunda
Convallaria majalis L.
L. smarodsii

loll all soll all Soll soll call wall call oll oa

Teall Soll Soll Soll soll

tN
‘Oo
wn

Convallaria multiflora L.
L. convallariae
Convallaria polygonatum L.
L. bellynckii
Cora pavonia (Swartz) Fries
L. corae
Corallorrhiza multiflora Nutt.
L. corallorhizae
Cordyline dracaenoides Kunth
L. cordylines
Cornus alba L.
L. limitata
Cornus florida L.
L. olivaespora
Cornus sanguinea L.
L. corticola
L. fiedlaeri
L. limitata
L. platycarpa
L. vagabunda
Cornus stolonifera Michx.
L. rugosa
Cornus sp.
L. mamillana
Coronilla coronata L.
L. dumetorum var.
coronillae
Coronilla emerus L.
L. riofrioi
Coronilla sp. [as C. comata L.]
L. coronillae
Corvus cornix [Animalia]
L. corvina
Corylus avellana L.
L. avellanae
L. vagabunda
Corylus sp.
L. leucoplaca
Cosmos sulphureus Cav.
L. cosmicola
Coumarouna punctata S.F. Blake
L. coumarounae
Cousinia multiloba DC.
L. shahvarica
Crataegus monogyna Jacq.
L. pomona forma tran-
silvanica
Crataegus oxyacantha L.
L. saccardiana
Crepis sibirica L.
L. sibirica
Crepis vesicaria L. subsp.
haenseleri (Boiss. ex DC.) P.D.
Sell
L. rubellula
Crithmum maritimum L.
L. helminthospora forma
crithimi-maritimi
L. rhopalispora
Cucumis sativus L.
L. alexandrinis

296

Cucurbita pepo L.
L. cucurbitae
Cycas revoluta Thunb.
L. cycadis
L. irrepta
Cynanchum sp.
L. modesta var. cibostii
L. scotophila
Cynodon dactylon (L.) Pers.
L. cynodontis-dactyli
L. korrae
L. narmari
L. rhizomatum
Cynodon transvaalensis Burtt-
Davy
L. narmari
Cynodon sp.
L. pontiformis
Cynoglossum officinale L.
L. cynoglossi
L. vindobonensis
Cynosurus cristatus L.
L. cynosurt
Cyperus flavescens L.
L. cyperi
L. cypericola
Cyperus monti L.
L. cyperina
Cyperus papyrus L.
L. papyri
Cyperus schoenoides Griseb.
L. ceballosi
Cytisus alpinus Miller
L. corrugans
Cytisus laburnum L.
L. lucina
Dactylis glomerata L.
L. caricis
L. culmicola var. nigrans
L. dactylina
L. nigrans
L. recutita
Danthonia frigida Vickery
L. gaubae
Daphne gnidium L.
L. daphnes
L. vagabunda forma
daphnes
Daphniphyllum macropodium
Mig.
L. daphniphylli
Dasylirion junceum Zucc.
L. dasylirii
Dasystoma sp.
L. doliolum
Datura stramonium L.
L. caballeroi
Daucus carota L.
L. longipedicellata
L. rostruptt

Illinois Natural History Survey Bulletin

Daucus sp.

L. comatella

L. modesta forma dauci
Daviesia latifolia R. Br.

L. daviesiae
Desmodium sp.

L. dissiliens

L. distributa
Desmoncus sp.

L. desmonci
Deutzia scabra Thunb.

L. dichroa
Dianella revoluta R. Br.

L. williamsii
Dianthus atrorubens All.

L. leptospora
Dianthus caryophyllus L.

L. dianthi
Dianthus prolifer L.

L. proliferae?

Dianthus superbus L.

L. dianthi
Dianthus sp.

L. dianthi
Dichosciadium ranunculaceum
(F. v. Mueller) Domin

L. dichosciadti
Didymella vincetoxici (de Not.)
Sacc.

L. didymellae-vincetoxici
Digitalis lutea L.

L. modesta forma digitalis-

luteae
Digitalis purpurea L.

L. valdiviensis
Digitalis sp.

L. digitalis
Dipsacus sylvestris Hudson

L. bractearum var.

bractearum

L. carpophila

L. modesta forma sylvestris
Dodonaea viscosa (L.) Jacq.

L. dodonaeae
Dorycnium suffruticosum Vill.

L. cucurbitarioides
Draba alpina L.

L. drabae
Dracaena draco (L.) L.

L. convallariae

L. convallariae forma

dracaenae

L. dracaenae

L. draconis
Dryas octopetala L.

L. dryadis

L. rostrupii
Drypis spinosa L.

L. rehmiana
Dulcamara sp.

L. vagabunda

Vol. 34 An. 3

Dupontia fisheri R. Br.

L. insignis

L. vagans

L. vagans forma scirpi
Dupontia psilosantha (Rupr.)
Griseb.

L. insignis
Echium vulgare L.

L. cesatiana

L. echiella

L. echii
Eichhornia crassipes (C.F.P.
Mart.) Solms-Laub.

L. eichhorniae
Elaeis guineensis Jacq.

L. elaeidicola

L. elaeidis
Eleocharis sp.

L. norfolcia
Elymus arenarius L.

L. arenaria

L. elymi

L. larseniana

L. quinta
Elymus? barbatus F. Kurtz.

L. lagenoides
Elymus glaucus Buckley

L. elongata
Elymus sp.

L. elymi
Elyna spicata Schrader

L. sparsa var. elynae
Elytrigia juncea (L.) Nevski

L. ammophilae
Empetrum nigrum L.

L. empetri
Endiusa hirsuta Alef.

L. endiusae
Ephedra distachya L.

L. ephedrae
Epichloe typhina (Pers.:Fr.) Tul.

L. associata
Epilobium angustifolium L.

L. cylindrospora
Epilobium dodonaei Vill.

L. cadubriae
Epilobium fleischeri Hochst.

L. epilobii

L. multiseptata forma

alpina

Epilobium hirsutum L.

L. tritorulosa
Epilobium montanum L.

L. epilobii
Epilobium trigonum Schrank

L. epilobii
Equisetum arvense L.

L. arvensis
Equisetum hyemale L.

L. berlesei

L. hiemalis

March 1991

Equisetum limosum L.

L. limosa
Equisetum variegatum Schleicher
ex Weber & Mohr

L. equiseti

L. equiseticola
Equisetum sp. [as E. pallidum]

L. larvalis
Eranthemum sp.

L. eranthemi
Eremochloa ophiuroides (Munro)
Hack.

L. korrae
Erianthus alopecuroides (L.) Ell.

L. orthogramma
Erica carnea L.

L. subtecta
Erigeron canadensis L.

L. agnita var. erigerontis

L. canadensis

L. congesta
Erigeron viscidus Rydb.

L. erigerontis
Erigeron sp.

L. doliolum

L. plurisepta
Eriobotrya japonica (Thunb.)
Lindley

L. eriobotryae

L. miyakeana

L. puttemansii
Eryngium agavifolilum Griseb.

L. vinosa
Eryngium campestre L.

L. eryngii

L. nigrella

L. woodrow-wilsonii
Erysimum cheiranthoides L.

L. submaculans
Erysimum diffusum Ehth.

L. acutispora
Erysimum erysimoides (L.)
Fritsch ex Janchen

L. acutispora
Erysimum hieracifolium L.

L. fusispora forma erysimi
Erythrina crista-galli L.

L. aerea
Erythrina sp.

L. erythrinae
Espeletia neriifolia Sch. Bip. ex
Wedd.

L. jahnii
Espeletia sp. (?E. schultzii>
Wedd.)

L. espeletiae
Eucalyptus globulus Labill.

L. molleriana
Euonymus europaeus L.

L. castagnei

L. constricta

Nomenclator of Leptosphaeria

Euonymus sp.

L. californica
Eupatorium cannabinum L.

L. agnita var. ambigua

L. cerastii
Eupatorium capillifolium (Lam.)
Small

L. clavispora
Eupatorium coronopifolium
Willd.

L. mesoedema
Eupatorium maculatum L.

L. jacksonii
Eupatorium purpureum L.

L. jacksonii
Eupatorium saucechicoense
Hieron.

L. preandina
Eupatorium sp.

L. doliolum
Euphorbia cyparissias L.

L. euphorbiae
Euphorbia dendroides L.

L. sacculus
Euphorbia esula L.

L. euphorbiae forma esulae
Euphorbia osyridea Boiss.

L. euphorbiicolla
Euphorbia oxyroidea [authority
unknown]

L. depressa
Euphorbia pilosa L.

L. euphorbiaecola
Euphorbia sibthorpii Boiss.

L. sibtorpii
Euphorbia wulfenii Hoppe ex
Koch

L. bresadolaeana
Euphorbia sp.

L. tolgorensis
Euphrasia lutea L.

L. coniformis
Eustephia sp.

L. australis
Fagus sylvatica L.

L. valdobbiae
Fagus sp.

L. faginea

L. seminuda
Farsetia incana (L.) R. Br.

L. conferta
Feijoa sp. [as F. feijoicola}

L. feijoae
Ferula badra-kema K. Pol.

L. ferulicola
Festuca arenaria Osbeck

L. arenaria
Festuca arundinacea Schreber

L. sabuletorum
Festuca kerguelensis Hook. f.

L. kerguelensis

Festuca sp.
L. crastophila
L. eustomella
L. stromatoidea
L. taminensis
Ficus elastica Roxb.
L. fici-elasticae
Filipendula ulmaria Maxim.
L. obesula
L. rustica
Flotovia excelsa DC.
L. flotoviae
Foeniculum vulgare Miller
L. foeniculacea
L. foeniculi
Foeniculum sp.
L. doliolum
L. doliolum var. pinquicula
Francoa sonchifoliae Cav.
L. francoae
Frasera speciosa Dougl. ex
Griseb.
L. fraserae
Frasera? sp.
L. harknessianna
Fraxinus americana L.
L. fraxini
Fraxinus sp.
L. controversa
L. leucoplaca
Fungi
. associata
. caucana
. cryptica
didymellae-vincetoxici
. fungicola
. geasteris
phyllachoricola
. phyllachorivora
. platychorae
. Stereicola
. tungurahuensis
Furcraea bedinghausti C. Koch
L. rusci forma fourcroyae
Furcraea longaeva Karw. &
Zucc.
L. infernalis
Furcraea sp.
L. translucens
Galeobdolon luteum Hudson
L. galeobdolonis
Galeopsis speciosa Miller
L. galeopsidicola
Galium aparine L.
L. aparines
L. scitula
Galium boreale L.
L. dumetorum var. galii-
borealis
Galium mollugo L.
L. galiicola

Toll Soll Sail Soll Sal all Sal all Sal Soll

L. molluginis
L. plectrospora
Galium sylvaticum L.
L. galii-silvatici
Galium sp.
L. galii
L. galiorum
Gaultheria shallon Pursh
L. gaultheriae
Geaster pseudolimbatus Hollos
L. geasteris
Genista anglica L.
L. genistae
Genista ferox Poir.
L. phiala
Genista tinctoria L.
L. fusispora
L. genistae var. microspora
Gentiana asclepiadea L.
L. umbrosa
Gentiana lutea L.
L. galiorum forma
gentianae
L. salebrosa
L. sapeyensis
Geranium richardsonii Fisch. &
Trautv.
L. modestula
Geranium sp.
L. subcaespitosa
Gerardia sp.
L. doliolum
Ginkgo biloba L.
L. ginkgo
Glyceria aquatica (L.) Wahlenb.
L. microscopica forma
glyceriae
L. recutita
Glyceria fluitans (L.) R. Br.
L. glyceriae
Glyceria plicata (Fries) Fries
L. glyceriae-plicatae
Glyceria spectabilis Mert. &
Koch
L. culmorum var. fla-
vobrunnea
Gnaphalium sp.
L. galiorum var. gnaphali-
ana
L. gnaphalii
Gossypium sp.
L. gossypit
Grindelia? sp.
L. nigricans var. grindeliae
Guazuma sp.
L. guazumae
Gymnadenia conopsea (L.) R. Br.
L. orchidearum
Gynerium argenteum Nees
L. gynerti
L. subiculifera?

Illinois Natural History Survey Bulletin

Gypsophila paniculata L.

L. gypsophilae
Haloxylon aphyllum (Minkw.)
Ijin

L. haloxyli
Hamemelis virginiana L.

L. hamamelidis
Hardenbergia violacea
(Schneev.) Stearn

L. hardenbergiae
Hedera helix L.

L. hederae

L. hedericola

L. helicicola

L. papulosa
Hedysarum obscurum L.

L. chochrjakovii
Heleocharis palustris (L.) Roem.
& Schult.

L. sowerbyi
Helianthemum sp.

L. helianthemi
Helianthus annuus L.

L. drechsleri

L. helianthi

L. lindquistii
Helianthus giganteus L.

L. helianthi
Helianthus grosserratus Martens

L. drechsleri
Helianthus sp.

L. consessa

L. doliolum

L. heliopsidis
Heliopsis sp.

L. drechsleri

L. heliopsidis
Helleborus foetidus L.

L. lathonia
Helleborus viridis L.

L. lathonia var. hellebori-

foetidi
Helonias bullata L.

L. heloniaefolia
Hemerocallis sp.

L. hemerocallidis
Heracleum lanatum Michx.

L. simmonsii
Heracleum palmatum Baumg.

L. doliolum var. dissimilis
Heracleum sphondylium L.

L. massarioides
Heracleum sp.

L. doliolum
Hermodactylus tuberosus (L.)
Miller

L. hermodactyli
Hesperis dinarica G. Beck

L. hesperidicola

Vol. 34 Art. 3

Hesperis sp.
L. galiicola var. brachy-
spora
Hevea brasiliensis Mill. Arg.

L. heveae
Hicoria sp.

L. exocarpogena
Hieracium murorum L.

L. brauni
Hierochloé alpina (Willd.)
Roemer & Schultes

L. hierochloae
Hierochloé éantarctica R. Br.

L. fuegiana
Hippophaé rhamnoides L.

L. fuscella var. hippophaes

L. fuscella var. sydowiana

L. hippophaes
Homo sapiens

L. senegalensis

L. tompkinsii
Hordeum arenarium Ascherson

L. arenaria
Hordeum distichon L.

L. recutita
Hordeum turkestanicum Nevski

L. horde
Hordeum vulgare L.

L. narmari

L. nodorum form sp. hordei
Humulus lupulus L.

L. dumetorum
Hura crepitans L.

L. hurae
Hydrangea arborescens L.

L. hydrangeae
Hydrocotyle asiatica L.

L. nesodes
Hyparrhenia sp.

L. hyparrheniae
Hypericum alpestris Stev. ex
Ledeb.

L. hypericola
Hypericum calycinum L.

L. vagabunda
Hypericum perforatum L.

L. cesatiana

L. dematium

L. hyperici

L. lankeana

L. ocellata
Ilex paraguariensis A. St. Hil.

L. paraguariensis

L. yerbae
Inula crithmoides L.

L. helminthospora

L. rhopalispora
Inula salicina L.

L. derasa var. franconica

L. franconica

March 1991

Ipomoea batatas (L.) Lam.

L. bataticola

L. ferruginea
Iris arenaria Waldst. & Kit.

L. iridis
Tris foetidissima L.

L. iridicola

L. vectis

L. xiphii
Tris germanica L.

L. heterospora

L. parvula var. iridis-

germanicae
Tris pseudacorus L.

L. iridigena

L. larseniana

L. longispora

L. microthyrioides

L. parvula
Isatis tinctoria L.

L. allorgei
Jambosa malaccensis DC.

L. eustomoides var.

punctata
Jasminum fruticans L.

L. emiliana
Jubaea spectabilis Humb.,
Bonpl., & Kunth

L. briosiana

L. jubaeae
Juglans regia L.

L. petiolaris
Juglans sp.

L. leucoplaca
Juncus acutiflorus Ehrh. ex
Hoffm.

L. juncorum
Juncus acutus L.

L. junci-acuti
Juncus articulatus L.

L. therophila
Juncus biglumis L.

L. junciseda
Juncus drummondii E. Mey.

L. solheimii
Juncus effusus L.

L. defodiens

L. hydrophila

L. junci

L. juncina

L. petkovicensis

L. pseudo-diaporthe

L. riparia

L. viridella
Juncus filiformis L.

L. sepalorum
Juncus glaucus Sibth.

L. junci

L. junci-glauci

L. subriparia

L. vitensis

Nomenclator of Leptosphaeria

Juncus hostii Tausch

L. juncicola
Juncus lampocarpus Ehth. ex
Hoffm.

L. lamprocarpi

L. monilispora
Juncus maritimus Lam.

L. maritima

L. neomaritima
Juncus roemerianus Scheele

L. neomaritima
Juncus trifidus L.

L. juncicola

L. sepalorum
Juncus sp.

L. michotii

L. norfolcia

L. variabilis
Juniperus communis L.

L. juniperi
Juniperus nana Willd.

L. juniperina

L. saprophila
Kalmia angustifolia L.

L. kalmiae
Kerria japonica (L.) DC.

L. vagabunda
Kerria sp.

L. coniothyrium

L. vagabunda var.

divergens
Kigelia pinnata DC.
L. dryadea subsp.
lussoniensis
Knautia arvensis (L.) Coulter

L. phyteumatis
Koeleria albescens DC.

L.-clavata
Koeleria cristata (L.) Pers.

L. herpotrichoides
Koeleria gracilis Pers.

L. clavata
Koerberiella wimmeriana
(Koerber) B. Stein

L. koerberi
Lactuca muralis (L.) Gaertner

L. quadriseptata
Lactuca sp.

L. doliolum
Lagotis minor (Willd.) Standley

L. holmii
Lamium album L.

L. parietariae forma lamii
Lantana camara L.

L. isocellula

L. rajasthanensis
Lappa sp.

L. eriophora

L. galiicola var. brachy-

spora

L. modesta forma lappae

299

Lapsana communis L.

L. galiorum var. lapsanae
Laserpitium angustifolium Georgi

L. oreophiloides
Laserpitium halleri Crantz

L. ladina
Laserpitium latifolium L.

L. ettalensis
Laserpitium siler L.

L. cornuta

L. sileris
Laserpitium sp.

L. spectabilis
Lathyrus latifolius L.

L. lathyrina

L. niessleana
Lathyrus niger (L.) Bernh.

L. dolioloides var. lathyri
Lathyrus sylvestris L.

L. lathyri

L. multiseptata
Laurus nobilis L.

L. lauri

L. nobilis

L. paoluccii
Laurus sp.

L. coniothyrium vat.

foliicola
Lavandula stoechas L.

L. lavandulae
Lavandula vera DC.

L. salviae forma minor
Lavandula sp.

L. cayanillesii
Lecanora frustulosa (Dicks.) Ach.

L. steinii
Lecidea excentrica Roehling

L. advenula
Lecidea vernalis (L.) Ach.

L. consocians
Leersia oryzoides (L.) Swartz

L. leersiae

L. leersiana
Leleba sp. [as L. simplex]

L. lelebae
Lemanea pluviale C. Aq.

L. fluviatilis

L. lemaneae
Leonurus cardiaca L.

L. doliolum var. leonuri
Lepidium campestre (L.) R. Br.

L. ogilviensis forma lepidii
Lepidium virginicum L.

L. virginica
Leptorhaphis leptogiophila Minks
ex Wint.

L. leptogiophila
Lepus europaeus

L. fimiseda
Lespedeza bicolor Turez.

L. lespedezae

300

Lespedeza capitata Michx.
L. microspora
Liatris scariosa (L.) Willd.
L. trimerioides
Libanotis montana Crantz
L. foliicola
L. libanotis
Libanotis sp.
L. longchampsi
Lichenes
. advenula
. apocalypta
. arnoldii
. baeomycearia
. caninae
clarkii
. consocians
corae
. crozalsti
. galligena
. geographicola
. inarensis
. koerberi
. leptogiophila
. leucomelaria
. lichenicola
. maheui
. mamillula
. neottizans
. oligospora
. parmeliarum
. peltigerarum
. peltigerea
. polaris
. pycnostigma
. ramalinae
. rivana
. rivana forma solorinae
. sphyridiana
. Steinii
. stereocaulorum
. tartarina
. umbilicariae
. usneae
Lilium martagon L.
L. martagoni
Lilium superbum L.
L. lilii
Lilium sp.
L. lilicola
Linum tenuifolium L.
L. hrubyana
Lippia (Aloysia) citriodora Kunth

ileal aioli ela sl ale al lalallala al alalalal lal elelel el ola oll elo l

L. octophragmia var. major

Lippia seriphioides A. Gray
L. basalduai
Liriodendron tulipifera L.
L. halima
Liriodendron sp.
L. stictoides

Illinois Natural History Survey Bulletin

Lobelia excelsa Lesch.
L. lobeliae
Lobelia tupa L.
L. tupae
Lolium perenne L.
L. eustomoides
L. lolii
Lomatia obliqua R. Br.
L. chilensis
Lonicera alpigena L.
L. albulae
Lonicera canadensis Bartr.
L. lonicerae
Lonicera caprifolium L.
L. caprifolii
L. corticola
Lonicera periclymenum L.
L. periclyment
Lonicera tatarica L.
L. frondis
L. periclymeni vat. tatarica
L. surculorum
Lonicera xylosteum L.
L. lonicerae
Lonicera sp.
L. aulica
L. lonicerae
Lophanthus sp.
L. lophanthi
Lunaria rediviva L.
L. lunariae
Lupinus sp.
L. barriae
L. byssincola
L. foeniculacea subsp.
lupina
L. frigida
L. lupini
L. lupinicola
Lupinis latifolius J.G. Agardh
L. wehmeyeri
Lupinus obtusilobus
L. castrensis
Lupinus rubricaulis Greene
L. wehmeyeri
Lupunus albicaulis var. shastensis
L. shastensis
Luzula albida (Hoffm.) DC.
L. culmorum forma
hungarica
L. epicalamia var.
pleosporoides
L. juncina forma macro-
spora
Luzula lutea (All.) DC.
L. epicalamia
L. microscopica var. alpina
Luzula maxima (Reichard) DC.
L. luzulae

Vol. 34 Art. 3

Luzula nemorosa (Pollich) E.H.F.
Meyer

L. epicalamia

L. petrakii
Luzula sylvatica (Hudson) Gaudin

L. caricis

L. epicalamia
Lycium vulgare Dunal.

L. lyciophila
Lycopodium annotinum L.

L. campisilii

L. crepini

L. lycopodina

L. marcyensis
Lycopodium clavatum L.

L. lycopodiicola
Lycopodium selago L.

L. marcyensis
Lycopodium selago L. var.
appressum Desv.

L. arctalaskana
Lycopus europaeus L.

L. cesatiana
Lygeum spartum L.

L. spartit
Lythrum alatum Pursh

L. lythri
Lythrum salicaria L.

L. salicaria
Maclura aurantiaca Nutt.
L. maclurae
Magnolia grandiflora L.

L. magnoliae
Magnolia yulan Desf.

L. yulan
Malus domestica Borkh.

L. mandshurica

L. pomona

L. ternata
Malus sp.

L. concentrica
Manihot utilissima Pohl

L. petri
Maranta arundinacea L.

L. marantae
Marattia sp.

L. caffra
Marrubium vulgare L.

L. dumetorum var.

marrubii
Matisia paraensis Hub.

L. matisiae
Medicago falcata L.

L. blumeri
Medicago sativa L.

L. circinans

L. medicaginicola

L. medicaginis

L. pratensis

ene i ———_s eer errr
———— ee —————eeeeeeeeeeeeee

March 1991

Medicago sp.

L. medicaginis

L. medicaginum
Melampyrum pratense L.

L. suffulta
Melandrium apetalum (L.) Fenzl

L. vanhoeffeniana
Melandrium triflorum J. Vahl. ex
Liebm.

L. vahlii
Melica altissima L.

L. culmicola forma melicae

L. fuscidula
Melica inaequiglumis Boiss.

L. melicae
Melica magnolii Gren. & Godron

L. fuscidula forma

magnolii
Melica uniflora Retz.

L. melicae
Melilotus alba Medicus

L. dumetorum forma

meliloti

L. meliloti
Melilotus sp.

L. viridella
Menispermum canadense L.

L. coniothyrium
Mentha piperita L.

L. substerilis
Mentha sylvestris L.

L. menthae

L. zahlbruckneri
Mertensia sp.

L. mertensiae
Mesogloia leveillei (J. Ag.)
Mengh.

L. mirandae
Metasequoia glyptostroboides Hu
& Cheng

L. metasequoiae
Mikania sp.

L. agnita var. acheniarum

L. mikaniae
Milium effusum L.

L. linearis
Minuartia sedoides (L.) Hiern.

L. dennisiana
Monotes dasyanthus Gilg.

L. baumii
Moriera sp.

L. morierae
Morus alba L.

L. fallax

L. massariella var. disticha
Morus sp.

L. japonica
Muehlenbeckia platyclados
Meissn.

L. muehlenbeckiae

Nomenclator of Leptosphaeria

Muehlenbergia filiformis Rydb.
L. georgius-fischeri
Muehlenbergia racemosa
(Michx.) Britton, Stern, & Pogg.
L. muehlenbergiae
Musa cavendishii Lamb. ex
Paxton
L. musae
L. musigena
Musa sp.
L. musarum
L. taichungensis
Muscari comosum (L.) Miller
L. muscari
Musci
L. bryophila
L. cinclidoti
L. heufleri
L. polytrichina
Myrica gale L.
L. myricae
Myrrhis odorata (L.) Scop.
L. ogilviensis forma
myrrhis-odorata
Myrtus communis L.
L. myrti
L. myrticola
Nandina domestica Thunb.
L. nandinae
Nardus stricta L.
L. nardi
L. pachytheca
Obione portulacoides (L.) Moq.
L. obiones var. evolutior
Obione sp.
L. obiones
Ochrolechia tartarea (L.) Massal
L. tartarina?
Ocimum sanctum L.
L. ocimicola
Oenothera biennis L.
L. capsularum
L. ellisiana
Oenothera longiflora L.
L. argentina
Onagra strigosa Rydb.
L. onagrae
Onobrychis sativa Link
L. onobrychidicola
L. onobrychidis

Ophiopogon japonicus Ker-Gawl.

L. ophiopogonis
Opuntia ficus-indica (L.) Miller

L. sicula
Opuntia lindheimeri Engelm.

L. opuntiae
Ornithogalum pyrenaicum L.

L. ornithogali
Orthrosanthus chimboracensis
Baker

L. orthrosanthi

301

Oryza sativa L.
. Cattanei
. inecola
. wamotoi
korrae
. narmari
oryzae
. oryzicola
. oryzina
. Salvinit
Oryzopsis miliacea (L.) Bentham
& Hooker ex Ascherson &
Graebner
L. papillosa
Oxalis stricta L.
L. aglaja
Oxyria digyna (L.) Hill
L. oxyriae
Paeonia moutan Sims.
L. moutan

eal Soll wall Soll all Sail Soll all

Palm

L. trochus
Pandanus odoratissimus Nor.

L. pandanicola
Pandanus utilis Bory

L. pandani
Panicum crus-galli L.

L. occidentalis
Panicum grumosum Nees

L. anthostomella

L. melanommoides

L. subsuperficialis
Panicum palmaefolium Koen.

L. panici
Papaver radicatum Rottb.

L. papaveris
Parietaria officinalis L.

L. muralis

L. parietariae
Parmelia saxatilis (L.) Arch.

L. parmeliarum
Parmelia sp. {as P. atrata
Zahlbr. |

L. galligena
Paspalum giganteum Baldw. ex
Vasey

L. proteispora
Pastinaca sativa L.

L. clivensis
Pastinaca sp.

L. doliolum

L. setosa
Pedicularis brachyodonta
Schlosser & Vuk. var. fallacis
(Beck) Hayek

L. malyi
Pedicularis bracteosa Benth. ex
Hook.

L. jacksonensis

302

Pedicularis debilis Franch. ex
Maxim.
L. doliolum var. angustis-
pora
Pelargonium capitatum (L.)
L’Her.

L. elaoudi
Pelargonium sp.

L. pelargonii
Peltigera aphthosa (L.) Willd.
forma complicata Th. Fries

L. peltigerarum
Peltigera canina (L.) Willd.

L. caninae

L. mamillula

L. rivana
Peltigera malacea (Ach.) Funck.

L. arnoldii
Peltigera cf. rufescens (Weis.)
Humb.

L. clarkii
Peltigera sp.

L. peltigerea
Pennisetum clandestinum Hochst.
ex Chiov.

L. narmari
Pennisetum purpureum Schum.

L. penniseti

L. penniseticola
Penstemon confertus Dougl. ex
Lindl.

L. lethalis
Periclymenum sp.

L. vagabunda forma

lonicerae
Persea americana P. Mill.
L. gratissima
L. gratissima var. longis-
pora
Persea lingue Nees

L. lingue
Petasites sp.

L. vagabunda var. caulium
Petroselinum hortense auct.

L. melanommoides
Peucedanum cervaria (L.)
Lapeyr.

L. penicillus
Peucedanum longifolium Waldst.
& Kit.

L. doliolum forma

syndoliola
Phaca alpina L.

L. phacae
Phalaris arundinacea L.

L. fuckelii forma filamentif-

era
Phaseolus vulgaris L.
L. phaseoli
L. phaseolorum

Illinois Natural History Survey Bulletin

Phleum arenarium L.

L. arenaria
Phleum boehmeri Wibel.

L. rousseliana

L. sequana
Phleum pratense L.

L. culmifida
Phlox decussata Lyon

L. phlogis
Phoenix canariensis Hort. ex
Chabaud

L. batumensis
Phoenix dactylifera L.

L. phoenicis

L. pinnarum

L. pinnarum vat. rachidis
Phoradendron townsendii Trel.

L. phoradendri
Phormium sp.

L. phormicola

L. phormii

L. roumegueri
Phragmites communis Trin.
. albo-punctata
. arundinacea
. arundinacea vat. godini
. clavicarpa
. culmicola var. hispalen-

sis
. culmorum forma
phragmitis
elaeospora
. graminis
. larseniana
nigrans forma arundinis
. perpusilla
. phragmiticola
. subalpina
. typharum subsp.
phragmatina

L. vaginae
Phragmites sp.

L. arundinacea

L. culmicola var. rhizoma-

tum

L. cumulata

L. mosana
Phyllachora disseminata Syd.

L. phyllachoricola
Phyllachora graminis (Pers.:Fr.)
Fuckel

L. phyllachorivora
Phyllachora tungurahuensis
Petrak

L. tungurahuensis
Phyllachora sp.

L. caucana

L. cryptica
Phyllostachys bambuoides
Siebold & Zuce.

L. bambusae

SN Sail Soll Soll call

SoU all Soll all Soll soll all

Vol. 34 Art. 3

Phyllostachys bambusoides Sieb.
& Zucc. var. bambusoides
L. minoensis

L. phyllostachydis
Physalis pubescens L.

L. physalidis
Physostegia virginiana (L.)
Benth.

L. physostegiae
Phyteuma scheuchzeri All.
L. psilospora

Phyteuma spicatum L.

L. phyteumatis
Phytolacca americana L.

L. clavigera

L. variegata
Phytolacca decandra L.

L. phytolaccae
Picris hieracioides L.

L. picridis
Pilularia sp.

L. pilulariae
Pimpinella anisum L.

L. pimpinellae
Pinecenectria sp.

L. almeidana
Pinus sylvestris L.

L. acicola

L. picastra

L. pini
Pinus sp.

L. squamata
Piper capense L.

L. piperis
Pircunia dioica Moq.

L. argentinensis
Pistacia terebinthus L.

L. emiliana
Placodium teicholytum (Ach.)
DG:

L. crozalsii
Plantago alpina L.

L. winteri
Plantago cynops L.

L. cynops
Plantago macrostachys Decne.

L. austro-americana
Plantago maritima L.

L. vagabunda subsp.

alvarensis
Plantago sp.

L. plantaginicola
Platanus occidentalis L.

L. platanicola
Platanus sp.

L. platanicola
Platychora ulmi (Schleich.:Fr.)
Petrak

L. platychorae

March 1991

Pleioblastus hindsii (Munro)
Nakai

L. bambusae
Pleioblastus simoni (Carriere)
Nakai

L. bambusae
Pleurospermum austriacum (L.)
Hoffm.

L. ogilviensis var.

pleurospermi

Plocama pendula W. Aiton

L. plocamae
Plumbago sp.

L. plumbaginis
Poa alpina L.

L. microscopica
Poa annua L.

L. muirensis
Poa aquatica L.

L. culmifraga var.

propinqua

Poa colpodea Th. Fries

L. microscopica
Poa nemoralis L.

L. linearis

L. poae
Poa pratensis L.

L. avenaria
Poa sudetica Haenke

L. culmifraga forma poae
Poa sp.

L. sparsa
Poa sp.?

L. anarithmoides
Polygonatum sp.

L. polygonati
Polygonum bistorta L.

L. altaica
Polymmis sp.

L. doliolum
Polytrichum formosum Hedw.

L. heufleri
Polytrichum juniperinum Willd.
ex Hedw. var. alpinum Schimp.

L. polytrichina
Populus alba L.

L. aegira

L. alcides

L. leucoplaca
Populus balsamifera L.

L. borealis var. populi
Populus nigra L.

L. salicinearum
Populus tremula L.

L. immunda
Populus sp.

L. decaisneana
Porella sp.

L. porellae
Potentilla argentea L.

L. minima

L. superficialis

Nomenclator of Leptosphaeria

Potentilla argyrophylla Wallich
ex Lehm.

L. hollosiana
Potentilla caulescens L.

L. pulchra
Potentilla fragarioides L.

L. hollosiana
Potentilla maculata Pour.

L. oligotheca
Potentilla sp.

L. doliolum
Poterium sanguisorba L.

L. poterii
Primula elatior (L.) Hill

L. primulana
Primula latifolia Lapeyr.

L. primulaecola
Primula sikkimensis Hook.

L. delawayi
Protea melaleuca R. Br.

L. protearum
Prunella vulgaris L.

L. brunellae

L. hesperia
Prunus avium L.

L. rimicola
Prunus baccata Borkh.

L. corticola
Prunus domestica L.

L. corticola

L. pruni
Prunus dulcis (Mill.) D.A. Webb

L. pachytheca
Prunus mume Sieb. & Zucc.

L. mume
Prunus spinosa L.

L. corticola

L. lycii
Pseudosasa japonica (Siebold &
Zucc. ex Steudel) Makino

L. bambusae
Pteridium aquilinum (L.) Kuhn

L. coorgica
Preris aquilina L.

L. aquilina
Pterocelastrus tricuspidatus
Walp.

L. pterocelastri
Punica granatum L.

L. puniciae
Pyrola secunda L.

L. marginata
Pyrus communis L.

L. lucilla

L. subcutanea
Pyrus sinensis Hort.

L. nashi
Quamoclidion multiflorum Torr.
ex A. Gray

L. quamoclidii

303

Quercus pedunculata Ehrh.

L. dryadea

L. vagabunda
Quercus robur L.

L. alcides forma quercina
Quercus suber L.

L. scolecosporarum
Quercus virens Ait.

L. janus
Quercus sp.

L. dryophila

L. leucoplaca

L. puteana
Radicula sp. (=Rorippa sp.)

L. reidiana
Ramalina sp.

L. ramalinae
Ranunculus affinis R. Br.

L. ranunculi
Ranunculus nivalis L. var.
sulphureus Wahlenberg

L. weberi
Ranunculus polyanthemos L.

L. ranunculi-polyanthemi
Ranunculus thora L.

L. thorae
Ranunculus villarsii DC.

L. andrijevicensis
Raphanus maritimus Sm.

L. raphani
Reseda lutea L.

L. resedae
Restio australis R. Br.

L. restionis
Rhamnus sp.

L. limitata
Rhinanthus sp.

L. affinis

L. dolioloides var.

rhinanthi
Rhizocarpon geographicum (L.)
DC.

L. geographicola

L. polaris
Rhodiola rosea L.

L. rhodiolicola
Rhododendron ferrugineum L.

L. rhododendri
Rhododendron sp.

L. californica

L. rhododendri
Rhus glabra L.

L. rhoina
Rhus typhina L.

L. curta
Ribes alpinum L.

L. cladophila

L. ribis
Ribes grossularia L.

L. grossulariae

304

Ribes nigrum L.
L. anceps
Ricinus communis L.
L. torrendii
Rinodina collectica (F\lk.) Am.
L. maheui
Robinia pseudacacia L.
L. eustoma forma legumi-
nosa
L. lyndonvillae
L. petiolicola
Rosa canina L.
L. aculeorum
Rosa sp.
L. coniothyrium
L. corticola
L. fuscella
L. hazslinszkit
L. lejostega
L. sepincola
Rubia peregrina L.
L. bornmuelleri
L. icositana
Rubus fruticosus L.
L. coniothyrium
L. notarisii
Rubus idaeus L.
L. oxyspora
L. praetermissa
Rubus odoratus L.
L. praetermissa
Rubus sp.
. abbreviata
. controversa
. doliolum
. hendersoniae
. osculanda
. sepincola
. thomasiana
. vagabunda var. sarmenti
Rudbeckia amplexicaulis Vahl.
L. rudbeckiae
Rulingia prostrata Maiden &
Betche
L. rulingiae
Rumex acetosa L.

L. quadriseptata
Rumex nepalensis Sprengel
L. rumicicola

L. rumicis
Rumex patientia L.
L. rumicis
Rumex scutatus L.
L. scutati
Rumex sp.
L. rumicis
L. sepincola
L. wegeliniana
Ruscus aculeatus L.
L. convallariae forma rusci
L. glauco-punctata

[opil Soll Sol Soll Soll all soll

Illinois Natural History Survey Bulletin

L. rusci

L. rusci forma caulina

L. ruscicola forma

cladodiicola
Ruscus androgynus L.

L. rusci
Ruscus hypoglossum L.

L. rusci

L. rusci var. rusci-

hypoglossi
Ruscus hypophyllum L.

L. rusci var. hypophyllum
Ruscus sp.

L. ruscicola
Ruta graveolens L.

L. bulgarica
Sabal serrulata Roem & Schult.

L. sabalicola
Sabal sp.

L. sabaligera
Saccharum officinarum L.
. bicolor
. Kuangfuensis
. sacchari
. saccharicola
. Spegazzini
. Spegazzini var. minor
. taiwanensis
. tucumanensis
Sagittaria sp.

L. duplex
Salicornia peruviana H.B.K.

L. promontorii
Salix alba L.

L. salicinearum
Salix aurita L.

L. cinerea
Salix caprea L.

L. vagabunda forma

salicis-capreae
Salix purpurea L.

L. purpurearum

L. vagabunda
Salix vitellina L.

L. coniothyrium
Salix sp.

. baggei

. borealis

. consimilis

. gillotiana

. hendersoniae
. leucoplaca

. lonicerina

. malojensis

L. xylogena
Salsola kali L.

L. kali

L. lecanora

L. salsolae
Salvia glutinosa L.

L. montana

Toot Sail all Soll Soll all Sal Sa

all all call Soll Soll Soll Soll

Vol. 34 An. 3

Salvia officinalis L.
L. nicolai
L. salviae
Salvia pratensis L.
L. caespitosa forma salviae
Sambucus ebulus L.
L. derasa forma macro-
spora
L. dumetorum forma ebuli
L. megalospora
L. ruthenica
Sambucus melanocarpa A. Gray
L. sambucina
Sambucus nigra L.
L. coniothyrium
L. sambuci
Sambucus racemosa L.
L. dumetorum vat.
dolichospora
L. hirta
L. luxemburgensis var.
dolichospora
L. tiroliensis
Sanguisorba officinalis L.
L. sanguisorbae
Sanguisorba polygama F. Nyl.
L. sanguisorbae
Sansevieria ehrenbergii
Schweinf. ex Baker
L. baldratiana
Sarothamnus scoparius (L.)
Wimmer ex Koch
L. fuscella forma micro-
spora
L. sarothamni
Sarothamnus sp.
L. californica
Sarracenia purpurea L.
L. scapophila
Sasa paniculata (F. Schmidt)
Makino
L. bambusae
Sasa sp.
L. sasacola
L. sasae
Sasaella ramosa (Makino)
Makino
L. bambusae
Satureja hortensis L.
L. fallaciosa
Saxifraga oppositifolia L.
L. brachyasca
Saxifraga sp.
L. monotis
Scabiosa columbaria L.
L. modesta
L. passerinii
Scabiosa urceolata Desf.
L. obesa
Scheuchzeria palustris L.
L. bacillifera

March 1991

Schoenocaulon officinale A. Gray

L. schoenocauli
Schoenoplectus lacustris (L.)
Palla

L. lacustris

L. sowerbyi
Scirpus holoschoenus L.

L. apogon

L. holmii

L. micropogon

L. striolata
Scirpus lacustris L.

L. culmicola var. aquatica

L. maculans

L. scirpina

L. sowerbyi
Scirpus maritimus L.

L. saxonica
Scirpus sylvaticus L.

L. nardi var. dubiosa
Scirpus sp.

L. littoralis

L. maculans

L. michotii
Scrophularia aquatica L.

L. camilla

L. scrophulariae
Scrophularia canina L.

L. torbolensis
Scrophularia nodosa L.

L. oreophiloides subsp.
scrophulariae
Scutellaria brevibracteata Stapf

L. davisiana
Secale cereale L.

L. secalina

L. secalis

L. setulosa
Secale sp.

L. pontiformis
Selaginella arenicola subsp.
acanthonota (Underwood) R.
Tryon

L. rostrata
Selaginella helvetica (L.) Spring

L. helvetica

L. helvetica forma major

L. lycopodiicola var. major

L. major
Selaginella rupestris (L.) Spring

L. rostrata
Selaginella underwoodii Hieron.

L. mellispora
Semele androgyna (L.) Kunth

L. semelina
Semiarundinaria tatebeana Muroi

L. bambusae
Senecio atratus Greene

L. clivensis

L. clivensis var. constricta

Nomenclator of Leptosphaeria

Senecio cordatus Koch
L. ogilviensis var.
senecionis-cordata
Senecio crassulus A. Gray

L. octoseptata
Senecio erucifolius L.

L. rothomagensis
Senecio fuchsti C.C. Gmelin

L. dumetorum var.

coniformis
Senecio jacobaea L.

L. derasa

L. modesta forma jacobae

L. ogilviensis
Senecio nemorensis L.

L. macrospora

L. robusta

L. senecionis
Senecio rupestris Waldst. & Kit.

L. robusta
Senecio vulgaris L.

L. dolioloides
Senecio sp.

L. clivensis

L. doliolum
Seseli montanum L.

L. montis-bardi
Seseli sp.

L. niessleana var. staritzii
Sieversia turbinata (Rydb.)
Greene

L. sieversiae
Silene acaulis (L.) Jacq.

L. hausmanniana

L. hausmanniana var.

cherleriae

L. sabauda

L. silenes-acaulis
Silene arvatica Lag.

L. sabauda forma arvaticae
Silene inflata Sm.

L. silvestris
Silene rosulata Soyer-Willem. &
Godr.

L. indeprensa
Sinapis arvensis L.

L. sinapis
Sisymbrium alliaria (L.) Scop.

L. alliariae
Sisymbrium loeselii L.

L. submaculans
Sisymbrium strictissimum L.

L. acutispora
Smallanthus uvedalia (L.)
Mackenzie

L. longipedicellata
Smilax aspera L.

L. aetnensis

L. catalaunica

L. smilacis

305

Smilax excelsa L.

L. dobrogica
Smilax mauritanica Poiret

L. papulosa
Smilax pseudo-china L.

L. nervisequa
Smilax sp.

L. doliolum

L. smilacis
Sobolewskia lithophila Bieb.

L. lithophilae
Solanum dulcamara L.

L. opizit

L. sarmenticia

L. solani

L. solanicola

L. umbrosa

L. vagabunda vat.

dulcamarae
Solanum sodomeum L.

L. sodomaea
Solanum sp.

L. comatella
Solidago caesia L.

L. longipedicellata
Solidago virgaurea L.

L. ogilviensis forma

megalospora

L. planiuscula
Solidago sp.

L. aulica

L. doliolum

L. perplexa

L. solidaginis
Solorina crocea (L.) Ach.

L. lichenicola

L. oligospora

L. rivana forma solorinae
Sonchus sp.

L. doliolum
Sophora jabonica L.

L. sophorae
Sophora tetraptera J. Mill.

L. martinianum
Sorbaria sorbifolia A. Br.

L. spiraeae
Sorbus aucuparia L.

L. sorbi
Sorbus sp.

L. leucoplaca
Sorghum arundinaceum Roem &
Schult.

L. sorgho-arundinacei
Sorghum vulgare Pers.
.amphibola
. culmifraga
. eustoma
. grisea
L. septovariata

foal all soll

306

Sparganium erectum L.
L. sparganii
Sparganium ramosum Hudson
L. huthiana
L. sparganii
L. sparsa var. meizospora
Sparganium sp.
L. clara
Spartina alterniflora Loisel.
L. neomaritima
Spartina juncea auct.
L. borziana
Spartina townsendii H. Groves &
J. Groves
L. neomaritima
Spartina sp.
L. discors
L. duplex
L. incarcerata
L. macrosporidium
L. marina
L. neomaritima
L. pelagica
L. spartinae
L. sticta
Spartium junceum L.
L. lusitanica
Spartium sp.
L. dioica
Sphyridium fungiforme Flotow
L. sphyridiana
Spiraea sorbifolia L.
L. spiraeae
Sporobolus depauperatus Torr. ex
Hemsl.
L. sporoboli
Staphylea trifolia L.
L. rubrotincta
Statice occidentalis Lloyd
L. staticicola
Steironema ciliatum (L.) Rafin.
L. steironematis
Stellaria graminea L.
L. isariphora
L. salebricola
Stellaria humifusa Rottb.
L. stellariae
Stellaria media (L.) Vill.
L. richoni
L. stellariae
Stellaria uliginosa Murray
L. uliginosa
Stenotaphrum secundatum
(Walter) O. Kuntze
L. narmart
Stereocaulon alpinum Laur.
L. apocalypta
L. stereocaulorum
Stereum bicolor (Pers.) Pers.
L. stereicola

Illinois Natural History Survey Bulletin

Stereum subpileatum Berkeley &
Broome

L. fungicola
Stipa capillata L.

L. stipae
Stipa pennata L.

L. matritensis
Stipa tenacissima L.

L. macrochloae

L. pampaniniana

L. stipae

L. stipae-minor
Stratiotes aloides L.

L. stratiotis
Suaeda australis Moq.

L. suaedae
Succisa pratensis Moench.

L. planiuscula
Succisa sp.

L. modesta forma succisae

L. morthieriana
Swertia sp.

L. swertiae
Symphytum caucasicum Bieb.

L. dumetorum var.

symphyti
Syringa vulgaris L.

L. trematostoma
Tamarix gallica L.

L. hollosii

L. meridionalis

L. tamaricis
Tamarix germanica L.

L. tamaricis
Tamarix sp.

L. crozalsiana
Tanacetum vulgare L.

L. dolioloides

L. tanaceti

L. thielensii
Taxus canadensis Marsh.

L. taxicola
Tecoma radicans Juss.

L. carpophila
Tellima grandiflora (Pursh)
Douglas ex Lindley

L. deficiens
Tephrosia virginiana (L.) Pers.

L. tephrosiae
Teucrium scorodonia L.

L. teucrii

L. wegeliniana forma

teucril
Thalictrum aquilegifolium L.

L. tenuis

L. thalictri

L. thalictrina
Thalictrum collinum Wallr.

L. thalictricola
Thalictrum dioicum L.

L. houseana

Vol. 34 Art. 3

Thalictrum flavum L.

L. cruenta
Thalictrum minus L.

L. quadriseptata
Thea sinensis L.

L. cavarae

L. hottai
Theobroma cacao L.

L. theobromicola
Thermopsis montana Nutt.

L. wehmeyeri
Thuja orientalis L.

L. thujaecola
Tilia sp.

L. leucoplaca

L. priuscheggiana
Tillandsia bicolor Brongn.

L. aerea
Tofieldia calyculata (L.)
Wahlenb.

L. crastophila forma

tofieldiae

L. oreophila

L. submodesta

L. tofieldiae
Tortula tortuosa Ehrh. ex Hedw.

L. bryophila
Tragopogon sp.

L. ophioboloides
Trevoa trinervia Gill. & Hook.

L. trevoae
Trichopteryx sp. [as T. afroflam-
mida|

L. trichopterygis
Trifolium alpestre L.

L. trifolii-alpestris
Trifolium angustifolium L.

L. agnita var. trifolii
Trifolium kingii S. Wats.

L. monticola
Trifolium medium L.

L. trifolii
Trifolium trichocephalum Bieb.

L. balcarica
Triglochin maritima L.

L. hollosiana

L. juncaginearum

L. maritima
Triglochin palustris L.

L. juncaginearum

L. monilispora forma

triglochinis

L. triglochinicola

L. triglochinis
Trisetum distichophyllum (Vill.)
Beauv.

L. volkartiana
Trisetum longiglume Hackel
L. mendozana
Trisetum spicatum (L.) K. Richter

L. holiosiana
L. volkartiana

March 1991

Trisetum subspicatum (L.) Beauv.

L. microscopica
Triticum aestivum L.
L. korrae
L. narmari
Triticum durum Desf.
L. avenaria form sp.
triticea
Triticum repens L.
L. rubelloides
Triticum vulgare Vill.
L. avenaria form sp.
triticea
L. nodorum
L. recutita
L. tritici
Triticum sp.
L. pontiformis
Trollius europaeus L.
L. trollii
Typha angustata Bory & Chaub.
L. aquatica
Typha angustifolia L.
L. dematiicola
L. iridigena var. typhae
L. lacustris
L. typharum
L. typhiseda
L. typhiseda forma
sodoloci
Typha latifolia L.
. bispora
. caricis
. grandispora
. kunzeana
. lacustris
. licatensis
. licatensis forma
rupefortensis
. maculans var. typhicola
. palustris
. perpusilla var. typhae
. pseudohleria
. punctillum
. typhae
. typharum
. typhicola
Typha sp.
L. duplex
L. mucosa
Ulmus campestris auct.
L. massariella
L. ulmicola
Ulmus sp.
L. leucoplaca
L. ramulicola

Soil oall soll all Soll all

Tool Soll all Soll Soll all all

Nomenclator of Leptosphaeria

Umbellularia sp.

L. odora
Umbilicaria pustulata (L.)
Hoffm.

L. umbilicariae
Urtica dioica L.

. acuta

. acuta forma insignis

. acuta forma urticae

. atropurpurea

. coniformis

. conoidea forma
macrospora

Urtica sp.

L. acutiuscula

L. doliolum
Usnea florida (L.) Web.

L. usneae
Vaccinium sp.

L. lejostega
Valeriana dubia Turcz.

L. dubia
Valeriana officinalis L.

L. agnita

L. carneomaculans

L. quadriseptata
Veratrum sp.

L. veratri
Verbascum phlomoides L.

L. echinops
Verbesina virginica L.

L. drechsleri
Veronica latifolia auct.

L. striata
Veronica urticifolia Jacq.

L. carpogena
Veronica sp.

L. doliolum
Viburnum tinus L.

L. tint
Vicia cracca L.

L. viciae
Vicia sativa L.

L. niessleana forma viciae
Vinca major L.

L. maderensis
Vinca media Hoffmanns. & Link

L. demissa
Vinca minor L.

L. vincae
Vincetoxicum officinale Moench

L. molybdina
Vincetoxicum sp.

L. scotophila
Vitex agnus-castus L.

L. casta

Tall wall all soll Soll oa!

307

Vitis vinifera L.
. ampelina
. cerlettii
. chaetostoma
. cirricola
cookei
gibelliana
. pampini
socia
. vinealis
. viticola
. vitigena
. vitis
Vitis vinifera L. subsp. sylvestris
L. sclavonica
Vitis sp.
L. pampini
Vriesea sp.
L. vrieseae
Xanthoxylum sp.
L. brasiliensis
Xerophyllum asphodeloides (L.)
Nutt.

Toll oll Soll soll all all Soll soll Soll call all oe

L. xerophylli
Xerophyllum tenax (Pursh) Nutt.
L. hysterioides
Yucca elephantipes
L. setosa
Yucca filamentosa L.
L. filamentosa
Yucca gloriosa L.
L. obtusispora
Yucca sp. [as Y. colusplei|
L. pacifica
Zea mays L.
L. bubakii
L. ceratispora
L. hyalospora
L. interspersa
L. luctuosa
L. maydis
L. patellaeformis
L. punctiformis
L. seriata
L. variiseptata
L. zeae
L. zeae-maydis
L. zeicola
Zingiber officinale Rosc.
L. zingiberis
Zizania latifolia Turez.
L. zizannivora
Zizania sp.
L. zizaniaecola

Host Family Index

Acanthaceae
L. acanthi
L. eranthemi
Aceraceae
L. aceris
L. controversa
L. diana
L. dioica
L. inquinans
L. leucoplaca
L. muelleri
L. obesula
. vagabunda
Agavaceae
. agaves
. almeidana
. baldratiana
. convallariae
. convallariae forma
dracaena
. cordylines
. dasylirii
. dracaenae
. draconis
. filamentosa
. infernalis
. obtusispora
. obtusispora forma
agaves
. pacifica
. phormicola
. phormit
. roumegueri
. rusci forma fourcroyae
. Setosa
Alismataceae
L. duplex
L. rivularis
Alstroemeriaceae
L. bomareae
Amaranthaceae
L. eriophora
Amaryllidaceae
L. australis
Anacardiaceae
L. brasiliensis
L. curta
L. emiliana
L. rhoina

ll

Poo oS

loatl Soll all Soll all Soll allo

ll all soll call soll

Apocynaceae
L. demissa
L. hardenbergiae
L. maderensis
L. vincae
Aquifoliaceae
L. paraguariensis
L. yerbae
Araceae
. acorella
-acort
. colocasiae
. crucheti
. densa
. microscopica subsp.
calomi
L. typharum
Araliaceae
L. doliolum
L. hederae
L. hedericola
L. helicicola
L. papulosa
Araucariaceae
L. californica
Arthropyreniaceae
L. leptogiophila
Asclepiadaceae
L. asclepiadis
. dearnessii
. doliolum
. modesta var. cibostii
. molybdina
. russellii
. scotophila
Aspleniaceae
L. asplenii
Avicenniaceae
L. australiensis
L. avicenniae
Baeomycetaceae
L. baeomycearia
L. neottizans
L. pycnostigma
L. sphyridiana
Berberidaceae
L. berberidicola
L. berberidis
L. coniothyrium forma
berberidis

foil all soll all Soll oa

(all ool Soll Soll Sal Sa

L. inconspicua

L. nandinae

L. punjabensis
Betulaceae
. avellanae
. betulina
. betulina
. coniothyrium
. depressa
. leucoplaca
. lonicerina
. subsimilis
. vagabunda
. waghorniana
Bignoniaceae

L. carpophila

L. dryadea subsp.

lussoniensis

Boraginaceae

L. cesatiana

L. cynoglossi

L. dumetorum vat.

symphyti

L. echiella

L. echii

L. mertensiae

L. vindobonensis
Bromeliaceae

L. aerea

L. vrieseae
Buxaceae

L. buxina

L. melanommoides

L. revocans
Cactaceae

L. cerei-peruviani

L. opuntiae

L. sicula
Calycanthaceae

L. coniothyrium
Campanulaceae

L. lobeliae

L. pachyasca

L. phyteumatis

L. plemeliana

L. psilospora

L. tupae
Cannabaceae

L. cannabina

L. woroninii

Teall Sail Soll Soll Soll Soll all Soll Sal

March 1991

Cannaceae

E.

cannae

Capparaceae

L.
L.
L.
L.

ahmadii
capparidicola
capparidis
simillima

Caprifoliaceae

E.
E.
L.
L.
E.
L.

L.
E.

L.
L.
LE.
E

loll Sol

albulae

aulica

caprifolii
coniothyrium
corticola

derasa forma macro-
spora

dumetorum forma ebuli
dumetorum var.
dolichospora
frondis

hirta

lonicerae
luxemburgensis var.
dolichospora

. megalospora
. periclymeni
. periclymeni var.

tartarica

L. ruthenica
L. sambuci
L. sambucina
/ a
if
iE
E

surculorum

. tint
. tiroliensis
. vagabunda forma

lonicerae

Caryophyllaceae

[all oll soll all Soll all sa

call soll Soll Sail Soll Soll all Sol Soll Soll Sal all soll all Soll

. auerswaldii

. biebersteinii

. dennisiana

. dianthi

. gypsophilae

. hausmanniana

. hausmanniana var.

cherleriae

. indeprensa

. isariphora

. leptospora

. proliferae?

. rehmiana

_richoni

. Sabauda

. Sabauda forma arvaticae
. salebricola

. Sarraziniana

. Silenes-acaulis

. silvestris

. Stellariae

. Stellariae

. uliginosa

. vahlii

B.

vanhoeffeniana

Nomenclator of Leptosphaeria

Celastraceae

E..
E.
L.
E.
L.

californica

castagnei

constricta
dolioloides var. inops
pterocelastri

Chenopodiaceae

oll call Soll all Soll Soll all oll oll oll oll oll oll oll oll oll oll oll

. anthelmintica

. atriplicis

. calvescens

. camphorosmae
. chenopodii-albi
. echinella

. eriophora

. eutypoides
-haloxyli

. inculta

. kali

. lecanora

. nigricans

. obiones

. obiones var. evolutior
. promontorit

. Salsolae

. serbica

. Suaedae

Chondriellaceae

Ib,

mirabilis

Chordariaceae

Ib

mirandae

Cistaceae

|p
Ee.
i.

cist
cisticola
cistina

Clavicipitaceae

Ibe

associata

Compositae

Tall Soll Soll all Soll Gall Sail Sal Soll Soll Sal Soll Sail Soll Soll Soll all call sal all call call all oa

. achilleae

. agnita var. acheniarum
. agnita var. ambigua

. agnita var. chrysanthemi
. agnita var. erigerontis
. anacycli

. anthophila

. artemisiae

_ arthrophyma

. astericola

. asteris

. aulica

. bardanae

. bella

. bicuspidata

. braunt

. Caespitosa

. camphorata

. canadensis

. carduina

. carduorum

. carlinoides

. centaureae

. cerastil

Toll Soll Soll Soll all all Soll Soll Soll Soll Soll Soll Soll call Soll all wall Soll coll wall all wall Soll

[oo

Toptl Soll Sail Sail Soll all all Soll Sai Soll

fb

é

‘eoll call soll soll Soll soll Soll soll all Soll ll all call

309

. cervispora

. chrysanthemi

. cirsti-arvensis

. Clavispora

. clivensis

. Clivensis var. constricta
. coleosanthi

. compositarum

. compressa

. congesta

. conoidea forma asteris
. Consessa

. corynispora

. cosmicola

. crustacea

. cynaracearum

. derasa

. derasa forma alpestris
. derasa forma robusta
. derasa var. franconiea
. dolioloides

. dolioloides var. cirsii

. doliolum

. doliolum forma

carlinae-vulgaris

. doliolum var. cacaliae
. doliolum var. subdisti-

cha

. drechsleri
. dumetorum var.

coniformis

. eburnea

. erigerontis

. eriophora

. espeletiae

. filiformis

. flotoviae

. franconica

. fulgida

. galiicola var. brachy-

spora

. galiorum forma

cirsiorum

. galiorum var. gnaphali-

ana

. galiorum var. lapsanae
. gloeospora

. gnaphalii

. grammodes

. helianthemi

- helianthi

. heliopsidis

. helminthospora

. hispanica

. jaceae

. jacksonii

. jahnii

. kalmusii

. lasioderma

IE
E,

lindquistii
longipedicellata

310

loot Sal all Soll all So call Soll all all ll

SN Soll ll

oS

Toatl Soll Soll Sal Soll Soll Soll all Soll all Goll Soll all Sol all cal

Sail Soll all Sal Soll Sal Sall Sail Soll all Soll Soll Gall Soll Soll Soll all calls)

. macrospora
. marginalis

. mesoedema

. mikaniae

. millefolii

. modesta forma jaco-

baeae

. modesta forma lappae

. modesta var. rubellula

. nanae

. nigricans var. grindeliae
_ nitschkei

_ nitschkei forma

adenostylidis

. octoseptata
. ogilviensis
. ogilviensis forma

achilleae

. ogilviensis forma

megalospora

. ogilviensis vat.

senecionis-cordati

. ophioboloides
. owaniae

. passerinit

. perplexa

. phaeospora

. picridis

. plagia

. planiuscula

. plurisepta

. preandina

. ptarmicae

. purpurea

. quadriseptata
. rhopalispora

. robusta

. rothomagensis
. rothomagensis vat.

artemisiae

. rubellula

. rudbeckiae

. Senecionis

. septemcellulata
. Shahvarica

. sibirica

. Simmonsit

. Solidaginis

. Staritzit

. Sydowiana

. tanaceti

. fetonensis

. thielensii

. trichostoma

. trimerioides

. tumefaciens

. uncinata

. vagabunda var. caulium
. valesiaca

Illinois Natural History Survey Bulletin

Convolvulaceae
L. bataticola
L. ferruginea
Cornaceae
L. fiedlaeri
L. limitata
L. mamillana
L. olivaespora
L. platycarpa
L. rugosa
L. vagabunda
Corvidae [Birds]
L. corvina
Crassulaceae
L. rhodiolicola
Cruciferae
. acutispora
. alliariae
. allorgei
. arabidis
. cesatiana
. conferta
. drabae
. fusispora forma erysimi
. galiicola var. brachy-
spora
. hesperidicola
- Johansonii
. lithophilae
. lunariae
. maculans
. maculans forma
denudata
. morierae
. nap
_ nigrella
. norvegica
. ogilviensis forma lepidii
. olericola
. planiuscula forma
cruciferarum
. raphani
. reidiana
. Salebrosa
. sinapis
. submaculans
. trollii
. virginica
Cucurbitaceae
L. alexandrinis
L. cucurbitae
L. obesa
Cupressaceae
L. juniperi
L. juniperina
L. saprophila
L. thujaecola
Cycadaceae
L. cycadis
L. irrepta

Toll Soll all Soll all Sail SoU Soll all ll Soll all oN Soll Soll call coll all oll oll all

(eal Soll all Soll Soll all

Vol. 34 Art. 3

Cyperaceae
. aliena
. apogon
. caricicola
. caricina
. caricinella
. cariciphila
. Caricis
. Caricis-firmae
. caricis-vulpinae
. ceballosi
. Cladii
. Clara
. consobrina
. culmicola var. aquatica
. culmorum var. paleicola
. cumana
. cyperi
. cypericola
. Cyperina
. epicarecta
. folliculata
. folliculata var. oxyspora
. glgaspsora
. hemicrypta
- holmii
. kochiana
. lacustris
. littoralis
. littoralis forma
calamagrostidis-
arenariae
. macrotheca
. maculans
. michotii
. micropogon
. microscopica
. microscopica vat.
caricis-vulpinae
. occulta
. paludosa
. papyrt
. petkovicensis var. elymi
. pinnarum
. pinnarum var. rachidis
. puccinioides
. rivalis
. Saxonica
. Scirpina
. Sowerbyi
. Sparsa var. elymae
. Sparsa var. meizospora
. Striolata
. Striolata var. caricis-
glaucae
L. viridella
Daphniphyllaceae
L. daphniphylli
Dennstaedtiaceae
L. coorgica

oath Soll Sail Soll Soll oll Soll all Sal call coll Soll call call Soll call Soll all coll all coll oll oll coll oll alll all wall

oll call Soll Sol ool

[sail sll Soll all Soll all Soll Soll Soll Soll all all coll wall

March 1991

Dipsacaceae

L.
L.

Tool all call Soll Soll Soll Soll

L.

bractearum
carpophila var.
bractearum

. cephalariai-uralensis

. modesta

. modesta forma succisae
. modesta forma sylvestris
. morthieriana

. passerinii

. phyteumatis

. planiuscula forma

succisae
taurica

Dipterocarpaceae

L.

baumii

Elaeagnaceae
L. fuscella var. hippophaes
L. fuscella var. sydowiana

L.

hippophaes

Empetraceae

E:

empetri

Ephedraceae

Es.

ephedrae

Equisetaceae

[oll all soll soll soll all Se

. arvensis

. berlesei

. equiseti

. equiseticola
. hiemalis

. larvalis

. limosa

Ericaceae

‘ool Soll Soll Sall Soll Soll Soll all all Soll Soll

. andromedae

arbuti
azaleae

. californica

ericae

. gaultheriae

hyperborea
kalmiae

. lejostega

. rhododendri
. sublanosa

. subtecta

Euphorbiaceae

pil soll Soll Soll Soll Sail Sal Sal Se

. bresadolaeana

. depressa

. euphorbiae

. euphorbiae forma esulae
. euphorbiicolla

heveae

. hurae

. petri

. Sacculus
i.
is
Ee

sibtorpii
tolgorensis
torrendit

Fagaceae

L.
iE.

alcides forma quercina
dryadea

Nomenclator of Leptosphaeria

L. dryophila

L. faginea

L. involucralis

L. janus

L. leucoplaca

L. puteana

L. scolecosporarum
L. seminuda

L. vagabunda

L. valdobbiae

Geastraceae

L. geasteris

Gentianaceae

L. fraserae

L. galiorum

L. galiorum forma
gentianae

. harknessianna

. Salebrosa

. Sapeyensis

. Swertiae

. umbrosa

loll Soll wall all Se

Geraniaceae

L. elaoudi

L. hollosiana

L. modestula

L. pelargonii

L. subcaespitosa

Ginkgoaceae

L. ginkgo

Gramineae

. aeluropodis

. albo-punctata
algida

. alopecuri
ammophilae
amphibola
.amphiloga
anarithma

. anarithmoides
anisomeres

. anthostomella
arenaria
arenaria

. arrhenatheri

arundinacea

asperellae
avenae
avenaria
avenaria
bambusae
bambusicola
. beaumontii
bicolor

. borziana

. brachypodii
brizae

. bubakii

Si pial al al al al al al al al al al al al sl sl al al al ol al al al al al al ala

arrhenatheri var. italica

. arundinacea var. godini

poh

ll

| be
| by

all Soll Sai Soll Soll Soll call Soll Soll all Soll call call all call soll all we

Si al al pial ai al al al al al al al ai al al al a

caricis

. cattanet
. ceratispora
. chusqueae

clavata
clavicarpa
coccodes
coicis

. conimbricensis
. consociata

crastophila

. culmicola

. culmicola forma major

. culmicola forma melicae
. culmicola var. hispalen-

SIS

. culmicola var. minor
. culmicola var. migrans
. culmicola var. rhizoma-

tum

. culmifida
. culmifraga
. culmifraga forma

majuscula

. culmifraga forma

manuscula

. culmifraga forma poae
. culmifraga var. alpestris
. culmifraga var.

bromicola

. culmifraga var. linearis
. culmifraga var.

propinqua

. culmorum
. culmorum forma epigeti
. culmorum forma

phragmitis

. culmorum var. fla-

vobrunnea

. cumulata
. cynodontis-dactyli
. cynosurt

dactylina
discors
disseminata

. donacina

duplex
elaeospora
elongata
elymi
eumorpha
eustoma

. eustomella

. eustomoides

. eustomoides forma lolii

. fuckelii

. fuckelii forma filamentif-

era
fuegiana
fuscidula

L. fuscidula forma
magnolii

. gaubae

. georgius-fischert

. glyceriae

. glyceriae-plicatae

. graminis

. graminum

. grisea

. gynerit

. hazslinskyana

. herpotrichoides

. hierochloae

. hollosiana

. hordei

. hyalospora

. hyparrheniae

. Incarcerata

. inecola

. insignis

. insignis forma airae-
cespitosa

. intermedia

. interspersa

. ischaemi

. wamotoi

. kerguelensis

. korrae

. kuangfuensis

. lagenoides

. larseniana

. latebrosa

. leersiae

. leersiana

. lelebae

. linearis

. lineolaris

. littoralis

_lolit

. luctuosa

. macrochloae

. macrosporidium

. marina

.marram

. matritensis

. maydis

. media

. melanommoides

. melicae

. mendozana

. microscopica

. microscopica forma
brachypodii

. microscopica forma
glyceriae

. minoensis

. mosana

. moutoniana

. muehlenbergiae

_muirensis

aU atl Soll cll Soll Soll call all call all coll wall all all coll ll all allo

oll Soll Sail Soll all Sail all all Soll all Sal all Gall Soll Soll Soll Soll call Soll cll Sal all call all call cal all call allo

é

Tall Soll all Soll Sol

Illinois Natural History Survey Bulletin

loail Sal all Soll Soll Soll Soll all Soll all Soll all oe

Topil Soll all Soll Soll all Sol Soll Soll Soll all Soll Sal nll Sal all Soll Soll Soll Soll all all all call wall wall all Sol all call call call Soll Sol Soll call call call wall call Sol col call call call

. muricata

. nardi

. nardi var. dubiosa

. narmari

. neglecta

. neomaritima

. nigrans

. nigrans forma arundinis
. nodorum

. nodorum form sp. hordei
. norfolcia

. occidentalis

. ophiopogonis var.

graminum

. orthogramma
. oryzicola
. oryzae

oryzina

. pachycarpa

. pachytheca

. pampaniniana
. panict

. papillosa

. patellaeformis
. pelagica

. penniseticola
. penniseti

. perforans

. perpusilla

. personata

. phragmiticola
. phyllostachydis
. pleurospora

. poae

. poae var. agrostidis
. pontiformis

. proteispora

. puiggarit

. punctiformis

. quinta

. recessa

. recutita
_rehmii

. rhizomatum

. rhodophaea

. rousseliana

. rubelloides

. Sabuletorum

. Sacchari

. Saginata

. salvinii

. Sasacola

. Sasae

. Scabrispora

. Schneideriana
. secalina

. secalis

. Septovariata

. Sequana

. Seriata

Vol. 34 Art. 3

. setulosa

. sorghi-arundinacei

. Sparsa

. Spartit

. Spartinae

. Spegazzini

. Spegazzini var. minor

. sporoboli

. stellata

. Stipae

. Stipae-minor

. Straminis

. Stromatoidea

. subalpina

. subcompressa

. subiculifera?

. subsuperficialis

. sylvatica

. taiwanensis

. faminensis

. thurgoviensis

. tigrisoides

. trichopterygis

. trimera

. tritici

. tucumanensis

. typharum var. phrag-
matina

. vagans

. vagans forma scirpi

. vaginae

. variiseptata

. volkartiana

. weddellii

. zeae

. zeae-maydis

. zeicola

. zizaniaecola

. Zizannivora

[oath Soll all Soll Soll all Soll Soll Sal all all Soll all wall all call all call Soll all cal all voll all call all

(oll soll all Soll call Soll all all Soll all

Guttiferae

feel

. cesatiana

. dematium

. hyperict

. hypericola
. lankeana

. ocellata

. vagabunda

loll all sll all Sal Sad

Hamamelidaceae

L. hamamelidis

Hominidae

L. senegalensis
L. tompkinsii

Hydrocharitaceae

Iridace

L. stratiotis

ae

L. hermodactyli
L. heterospora
L. iridicola
L. iridigena
L. iridis

March 1991

L. larseniana

L. longispora

L. microthyrioides

L. orthrosanthi

L. parvula

L. parvula var. iridis-
germanicae

L. vectis

L. xiphii

Juglandaceae

L. cacuminispora
L. exocarpogena
L. lejostega

L. leucoplaca

L. petiolaris

Juncaceae

L. caricis

L. culmorum forma
hungarica

L. defodiens

L. epicalamia

L. epicalamia var.
pleosporoides

L. frigida

L. hollosiana

L. hydrophila

L. junci

L. juncicola

L. juncina

L. juncina forma macro-
spora

L. junciseda

L. junci-acuti

L. junci-glauci

L. juncorum

. lamprocarpi

. luzulae

. maritima

. michotit

. monilispora

. neomaritima
. norfolcia

. petkovicensis
. petrakii

. pseudo-diaporthe
. riparia

. Sepalorum

. solheimii

. subriparia

. therophila

. variabilis

. vitensis

‘all soll all Soll Soll Sal Soll Soll all Sal Sall Soll Sal all Soll Sal all Se

Juncaginaceae

L. juncaginearum

L. maritima

L. monilispora forma
triglochinis

L. triglochinicola

L. triglochinis

. microscopica var. alpina

Nomenclator of Leptosphaeria

Labiatae

. ballotae

. brightonensis

. brunellae

. caespitosa forma salviae
. cavanillesii

. cesatiana

. collinsoniae

. darkeri

. davisiana

. doliolum var. leonuri
. dumetorum var.

(ail Soll Soll Soll Soll all Soll all Soll Soll

[all sll pall Soll al Soll Soll all call Soll call Soll Soll Soll all call wall wall wall

EE.

marrubii

. fallaciosa

. fiumana

. galeobdolonis

. galeopsidicola

. hesperia

. lavandulae

. lophanthi

. menthae

. montana

_ nicolai

. ocimicola

. parietariae forma lamii
. physostegiae

. Salviae

. Salviae forma minor
. Slovacica

. substerilis

. feucril

. translucens

. wegeliniana forma

teucril
zahlbrucknert

Lauraceae

L.
Ib.
Ib

hone

almeidae
cinnamomi
coniothyrium var.
foliicola

. gratissima
. gratissima var. longis-

pora

. lauri

. lingue
_ nobilis
. odora
iL,

paoluccii

Lecanoraceae

L.

steinit

Lecideaceae

|G
E.

advenula
consocians

Leguminosae

opi Soll Sal Soll Sal Sal So

. adesmicola

. aerea

. agnita var. trifolit
. alhagii
-ammothamni

. amorphae

. aplos

. apios-fortunei

. balcarica

. barriae

. blumeri

. byssincola

. californica

. calopogonii

. cassiaecola

. castrensis

. chochrjakovii

. circinans

. clelandii

. comatella

. coronillae

. corrugans

. coumarounae

. cucurbitarioides

. daviesiae

. dioica

. dissiliens

. distributa

. dolioloides var. lathyri

. dumetorum forma
meliloti

. dumetorum var.
coronillae

. endiusae

. erythrinae

. eustoma forma legumi-
nosa

L. foeniculacea subsp.

lupina

L. fuscella forma macro-
spora

. fusispora

. genistae

. geniStae Var. microspora

. lathyri

. lathyrina

. lespedezae

. lucina

. lupini

. lupinicola

. lusitanica

. lyndonvillae

. marginalis

. martinianum

. medicaginicola

. medicaginis

. medicaginum

. meliloti

. microspora

. monticola

. multiseptata

. niessleana

. niessleana forma viciae

. onobrychidicola

. onobrychidis

. petiolicola

. phacae

. phaseoli

Topil all Soll Soll all all Soll Soll Gall Soll all call Soll call wall call call call all wall wall call

é

noo

[soll soll soll Soll Soll all Soll Soll Soll Sal Soll Soll Sol all Soll Soll all wall wall all coll call coll call all call oa

314

L. phaseolorum
L. phiala
L. pratensis
. riofriot
sarothamni
shastensis
sophorae
. tephrosiae
. trifolii
. trifolii-alpestris
. viciae
. viridella
. wehmeyeri
Lemaneaceae

L. fluviatilis

L. lemaneae
Leporidae

L. fimiseda
Liliaceae

SM all coll coll coll wall call all call on

aloes
. antherict
asparagt
asparagina
. aspidistrae
. bellynckii
. comatella
. convallariae
. crastophila forma

tofieldiae
. dobrogica
. heloniaefolia
. hemerocallidis
. hysterioides
indica
lassenensis
lilicola
lilii
martagoni
muscart
nervisequa
oreophila
. ornithogali
papulosa
passeriniana
polygonati
. portoricensis
praeclara
. punctoidea
schoenocauli
. semelina
. smarodsii
socialis
submodesta
. tofieldiae
veratri
. verwoerdiana
. williamsii
. xerophylli
Linaceae

L. hrubyana

[pil Soll Soll Soll Soll all Soll wall oa

Si aial aint ai al ol ala Slat alal ala al al al ai al al al al al al ai al a

Illinois Natural History Survey Bulletin

Loganiaceae

L. buddlejae

L. davidii

L. polini
Lycopodiaceae

L. arctalaskana

L. campisilti

L. crepini
L. lycopodiicola
L. lycopodina

L. marcyensis
Lythraceae

L. lythri

L. salicaria
Magnoliaceae

L. halima

L. magnoliae

L. stictoides

L. yulan
Malvaceae

L. abutilonis

L. gossypit
Marantaceae

L. marantae
Marattiaceae

L. caffra
Marsileaceae

L. pilulariae
Matoniaceae

L. matisiae
Menispermaceae

L. coniothyrium
Moraceae

L. coniothyrium

L. fallax

L. fici-elasticae

L. japonica

L. maclurae

L. massariella var. disticha
Musaceae

L. musae

L. musarum

L. musigena

L. taichungensis
Myricaceae

L. myricae
Myrtaceae

L. eustomoides var.

punctata

L. feijoae

L. molleriana

L. myrti

L. myrticola
Nyctaginaceae

L. quamoclidii
Oleaceae

L. controversa

L. emiliana

L. fraxini

L. leucoplaca

L. trematostoma

Vol. 34 Art. 3

Onagraceae
L. argentina
L. cadubriae
L. capsularum
L. cylindrospora
L. ellisiana
L. epilobii
L. multiseptata forma
alpina
L. onagrae
L. tritorulosa
Orchidaceae
L. corallorhizae
L. ophiopogonis
L. orchidearum
Oxalidaceae
L. aglaja
Paeoniaceae
L. moutan
Palmae
. algarbiensis
. arecae
. batumensis
. briosiana
. chamaeropis
. coccothrinacis
cocoes
. debeauxti
. desmonci
. doliolum var. pachy-
spora
. elaeidicola
. elaeidis
. jubaeae
. magnusiana
. molleriana
. phoenicis
. pinnarum
. pruni
. sabalicola
. Sabaligera
. Spatharum
. trochus
Pandanaceae
L. pandani
L. pandanicola
Papaveraceae
L. bocconiae
L. chelidonii
L. papaveris
Parmeliaceae
L. gallingena
L. parmeliarum
Peltigeraceae
. arnoldii
. caninae
clarkii
. lichenicola
.mamillula
. oligospora
. peltigerea

SI all coll ool all call all call ool oa!

opll Soll all Sal all ll Soll all Soll all call

‘all all call Soll call all

March 1991

L. rivana

L. rivana forma solarinae
Pertusariaceae

L. tartarina
Phaeosphaeriaceae

L. didymella-vincetoxici
Phyllachoraceae

L. caucana

L. cryptica

L. phyllachoricola

L. phyllachorivora

L. tungurahuensis
Physciaceae

L. leucomelaria

L. maheui
Phytolaccaceae

L. argentinensis

L. clavigera

L. phytolaccae

L. variegata
Pinaceae

L. acicola

L. faulii

L. picastra

L. pini

L. squamata

L. vagabunda

L. vagabunda forma abietis
Piperaceae

L. piperis
Plantaginaceae

L. austro-americana

L. cynops

L. plantaginicola

L. vagabunda subsp.

alvarensis

L. winteri
Platanaceae

L. platanicola
Plumbaginaceae

L. kotschyana

L. plumbaginis

L. staritzii

L. staticicola
Polemoniaceae

L. collumiae

L. phlogis
Polygonaceae
_ altaica
. atraphaxidis
. calligoni
. muehlenbeckiae
. oxyriae
. quadriseptata
. rumicicola
rumicis
. SCutati
. sepincola
. wegeliniana

‘Sail soll Sal Sail Soll all Sal Sol Soll Soll Se

Nomenclator of Leptosphaeria

Polypodiaceae

L. impressa
Polytrichaceae

L. heufleri

L. polytrichina
Pontederiaceae

L. eichhorniae
Porellaceae

L. porellae
Porpidiaceae

L. koerberi
Primulaceae

L. delawayi

L. pachyasca

L. primulaecola

L. primulana

L. steironematis
Proteaceae

L. chilensis

L. protearum
Pteridaceae

L. aquilina
Punicaceae

L. puniciae
Pyrolaceae

L. marginata
Ramalinaceae

L. rahmalinae
Ranunculaceae
. aconitt
. agminalis
. agminalis forma minor
. andrijevicensis
. anemones
anthostomoides
. aquilegiae
aucta
cruenta
doliolum var. pachy-
spora
grignonnensis
. haematites
. houseana
. Incruenta
. lasiosphaerioides
lathonia
. lathonia var. hellebori-
foetidi
L. napelli
L. nectrioides
L. nigromaculata
L. platypus
L. pleosporoides
L. pyrenopezizoides
Er.
|b
Ls:
L.
E.
| i

ail all coll all call oll all all ll

feat ool soll all soll wall a!

quadriseptata
ranunculi
ranunculi-polyanthemi
raphidophora

rimalis

scotophila

315

. tenuis
. thalictricola
. thalictrina
. thalictri
. thorae
. trollii
. umbrosa
. vagabunda
. vitalbae
. vitalbae var. sarmenti-
cola

L. weberi
Resedaceae

L. resedae
Restionaceae

L. restionis
Rhamnaceae

L. ceanothi

L. limitata

L. trevoae
Rhizocarpaceae

L. geographicola

L. polaris
Rosaceae
. abbreviata
.aculeorum
. ArUNCL
. cercocarpt
concentrica
coniothyrium
controversa
corticola
doliolum
dryadis
. eriobotryae
. fuscella
hazslinszkii
. hendersoniae
hollosiana
lejostega
. leucoplaca
lucilla
mandshurica
minima
miyakeana
mume
nashi
. notarisil
obesula
oligotheca
osculanda
. oxyspora
pachytheca
pomona
pomona forma tran-
silvanica
L. poterii
L. praetermissa
L. pruni
L. pruni forma plurivora

Topil Soll Soll Gall Soll all Sail Sal Sail Se

PER RRP REP RPP PSP PP RP Pe

316

lh,
If
_ rimicola

. rostrupit

. rustica

. saccardiana

. Sanguisorbae

. sepincola

. Sleversiae

. sorbi

. Spiraeae

. subcutanea

. superficialis

. ternata

. thomasiana

. umbrosa

. vagabunda

. vagabunda var.

Teall all Soll Soll all wall soll Soll ool all wall wall Soll all ll on

IU,

pulchra
puttemansit

divergens
vagabunda var. sarmenti

Rubiaceae

Test Soll Soll Soll Soll all all

leh le)

Todd Soll Soll ll Sail Soll Sail Sall Sal all Soll sll all Sal all all Gall ll

. africana

. aparines

. bornmuelleri

. canephorae

. centrafricana

. coffaeicida

. coffeigena

. coffeigena var. longiros-

trata

. cylindrospora
. dumetorum var. galii-

borealis

. excelsa

. galit

. galticola

. galii-silvatict
. galiorum

. glgaspora

. icositana

. lobayensis

. longispora

. macrorostra
. molluginis

. oubanguiensis
. plectrospora
. plocamae

. politis

. pusilla

. scitula

. tetraspora

. tonduzi

Ruscaceae

lool Soll Soll Soll Soll

é

. convallariae forma rusci
. glauco-punctata

.rusct

_rusci forma caulina

_ rusci var. hypophylli

. FUSCL Var. rusci-

hypoglosii

. ruscicola

Illinois Natural History Survey Bulletin

Rutaceae

Ee
L.
6,
L.
i.

bondari

bulgarica

citricola

papulosa

vagabunda forma citri-
limonii

Salicaceae

Tail Sal all Soll Soll Soll Soll Soll Soll all Soll Soll call alll call coll call call

L

. aegira

. alcides

. baggel

. borealis

. borealis var. populi
. cinerea

. coniothyrium

. consimilis

. decaisneana

. gillotiana

. hendersoniae

. Immunda

. leucoplaca

. lonicerina

. malojensis

. purpurearum

. Salicinearum

. vagabunda

. vagabunda forma

salicis-capreae

. xylogena

Sapindaceae

Ih

dodonaeae

Sarraceniaceae

Ib

scapophila

Saxifragaceae

Ib.
. brachyasca

. Cladophila

. deficiens

. dichroa

. francoae

. grossulariae

. hydrangeae

. monotis

. ramsaugiensis
. ribis

[all Soll Soll Soll Soll Soll Sal Sal Sal Sa

anceps

Scheuchzeriaceae

LE.

bacillifera

Scrophulariaceae

Toll all Soll all Soll Soll Soll all Soll Sal

é

. affinis
_anarrhini
aquilana

. camilla

. carpogena
. castilleiae
. castillejae
. concinna

. coniformis
. digitalis

. dolioloides var.

rhinanthi

. doliolum

SI Soll Soll Soll all coll oo

oath cal Soll Soll Soll oa!

Vol. 34 Art. 3

. doliolum var. angustis-

pora

. echinops
. galiorum subsp.

antirrhini

_holmii

. insulana

. jacksonensis

. lethalis

_malyt

. modesta forma digitalis-

luteae

. oreophiloides subsp.

scrophulariae

. scrophulariae
. Striata

. suffulta

. thuemeniana
. torbolensis

. valdiviensis

Selaginellaceae

Todd Soll Soll ll Soll

helvetica

. helvetica forma major

. lycopodiicola var. major
. major

. mellispora

. rostrata

Simaroubaceae

E.
Ee
E.

ailanthi
endophaena
glandulosae

Smilacaceae

EL.
E.
Ib
EE.

aetnensis
catalaunica
doliolum
smilacis

Solanaceae

Tool soll soll alll call wall coll soll wall Soll all call oo

. caballeroi

. comatella

. lyeti

. lyciophila

. opizii

. physalidis

. Sarmenticia
. sSodomaea

. solani

. solanicola

. umbrosa

. vagabunda
. vagabunda var.

dulcamara

Sparganiaceae

L.
L.
E..
L.

clara

huthiana

sparganii

sparsa var. meizospora

Staphyleaceae

L.

rubrotincta

Sterculiaceae

Es

Ruazumae

March 1991

L. rulingiae

L. theobromicola
Stereaceae

L. fungicola

L. stereicola
Stereocaulaceae

L. apocalypta

L. stereocaulorum
Tamaricaceae

L. crozalsiana

L. hollosii

L. meridionalis

L. tamaricis
Taxaceae

L. taxicola
Taxodiaceae

L. metasequoiae
Teloschistaceae

L. crozalsii
Theaceae
. camelliae
. camelliae-japonicae
. cavarae
. depressa
. hottai
. tornatospora
Thelephoraceae

L. corae
Thymelaeaceae

L. daphnes

L. vagabunda var. daphnes
Tiliaceae

L. leucoplaca

L. priuscheggiana
Typhaceae
. aquatica
. bispora
. caricis
. dematiicola
duplex
grandispora
. iridigena var. typhae
. kunzeana
. lacustris
. licatensis
. licatensis forma

rupefortensis
. maculans var. typhicola
. mucosa
. palustris
. perpusilla var. typhae
. pseudohleria
. punctillum
typhae
typharum
. typharum forma acori
. typhicola
. typhiseda
. typhiseda forma
sodoloci

Teall soll all soll soll

oll soll soll all Soll call all call all all oa

‘all Soll Soll Soll Soll Soll Soll Sal all Soll Soll Se

Nomenclator of Leptosphaeria

Ulmaceae
L. leucoplaca
L. massariella
L. ramulicola
L. ulmicola
Umbelliferae
. agnita
. agnita var. bupleuri
. agnita var. major
bupleurt
cibostii
clivensis
comatella
cont
. conligena
. conoidea
. conoidea forma
angelicae
. cornuta
. diaporthoides
. dichosciadii
. doliolum
.doliolum forma
syndoliola
. doliolum var. dissimilis
doliolum var. pinquicula
. eryngil
. ettalensis
. ferulicola
. foeniculacea
. foeniculi
. foliicola
. helminthospora forma
crithimi-maritimi
ladina
. libanotis
. longchampsi
. longipedicellata
. massarioides
melanommoides
. modesta forma dauci
modesta var. cibostii
. montis-bardi
nesodes
niessleana var. staritzil
_nigrella
. obesula
. ogilviensis forma
myrrhis-odorata
. ogilviensis var.
pleurospermi
. oreophiloides
. penicillus
. pimpinellae
. ranunculoides

iE
1b
L
L
L. rhopalispora
Ji,
E
Ib
i

[oath Soll Soll call So Soll Soll all Soll Soll all coll all call wall oa!

all all Soll all Soll call soll all a

[oll all Soll Soll Soll all Soll call call coll oll call ll

é

. rhopographoides
. rostruput

. rubicunda

. Setosa

317

L. sileris

L. spectabilis

L. umbrosa

L. utahensis

L. vinosa

L. woodrow-wilsonii
Umbilicariaceae

L. umbilicariae
Urticaceae
. acuta
. acuta forma insignis
. acuta forma uticae
. acutiuscula
. atropurpurea
. cecropiae
. coniformis
. conoidea forma

macrospora

L. doliolum
L. dumetorum
L. muralis

L. parietariae
Usneaceae

L. usneae
Valerianaceae
. agnita
. carneomaculans
. dubia
. galiicola var. brachy-

spora

L. quadriseptata
Venturiaceae

L. platychorae
Verbenaceae
. baldingerae
. basalduai
casta
. clerodendri
. isocellula
. octophragmia
. octophragmia var. major
. rajasthanensis
Viscaceae

L. phoradendri
Vitaceae
. ampelina
. cerlettii
. chaetostoma
. cirricola
cookei
. gibelliana
. pampini
sclavonica
socia
vagabunda
vinealis
viticola
viligena
Vitis
Zingiberaceae

L. alpiniae

L. zingiberis

oil all Soll all soll call all

oil all alls

loath call all oll Soll Soll Soll Sal

ll al ol all all all all all ail oll aad ll

Substrate Index

Achenes

L
Ib

. agnita var. acheniarum

. Inconspicua

Apothecia

Mi
L.

Ashes

fb

Bark

Teall Sail Soll Soll all all call soll

Berries

wh

Bracts

pooh

Toll Soll all Solon!

Branch

oO

Tall call all oll Soll Soll Soll Sal Soll Sal Soll Sal Sl Sl Sal Sal Sa

consocians
maheui

. inconspicua

. akagiensis

. avicenniae
. buxina

. clerodendri
. Inquinans

. Inspersa

. leucoplaca
_muelleri

. squamata

. saprophila

. arctalaskana

. bractearum

. carduina

. carpophila var.

bractearum

. crepini

. lamprocarpi
. lycopodina

. spatharum

. vitensis

S

. adesmicola
. aerea

. aetnensis

. africana

. ahmadii

. ailanthi

. alhagii

. ambiens
.ammothamni
. amorphae

. anceps

_ arbuti

. avellanae

. azaleae

. baggei

. berberidis
. betulina

. basalduai

. biebersteinii

. borealis var. populi

. borziana

. capparidicola

. capparidis

. castagnei

. Catataunica

. cavanillesii

. chilensis

. cinerea

- cinnamomi

. CISti

. cisticola

. clelandii

. coffeigena var. longiros-
trata

. consimilis

. constricta

. controversa

. crozalsiana

. crucheti

. cylindrospora

. daviesiae

. derasa forma macro-
spora

. desmonci

. dichroa

. dioica

. dobrogica

. dodonaeae

. dumetorum var.
dolichospora

. elaoudi

. emiliana

. endophaena

. ephedrae

. euphorbiicolla

. excelsa

L. fiedlaeri

L. fuscella forma micro-

spora

L. fuscella var. hippophaes

L. fuscella var. sydowiana

L. fusispora forma erysimi

L. gibelliana

L. gigaspora

L. gillotiana

L. ginkgo

foot soll Soll Soll Soll Soll call coll oll all oll Soll coll coll wal oa

[Soll all Soll Soll Soll Sain Soll Soll all Soll all Sal wal

loll soll soll call Soll oe

[ool Soll all Soll Soll all Soll all voll coll oll Soll all

Tooth soll all soll

sath Soll Sol all Soll all Soll soll Sal call Soll all voll Soll all call all all coll coll soll Soll coll call wall wall coll

. gratissima

. grossulariae
. hazslinszkii

. hendersoniae
. hippophaes

. hirta

. hollosii

. hydrangeae

. kuangfuensis
. lejostega

. limitata

. longispora

. lonicerae

. luxemburgensis var.

dolichospora
. macrorostra
. malojensis
. mamillana
. massariella
. massariella var.
brasiliensis
. melanommoides
. meridionalis
. muehlenbergiae
_mume
. myricae
_myrti
. myrticola
. nigrella
. odora
. olivaespora
. osculanda
. pachytheca
. periclymeni
. phiala
. platycarpa
. polini
. preandina
. puniciae
. revocans
. rhododendri
. ribis
. rimicola
. riofriort
. Salviae
. Sambuci
. Sarothamni
. Sclavonica
. simillima

LO LT

March 1991

. smilacis

. Spiraeae

. stictoides

. suaedae

. subcutanea

. subsimilis

. tamaricis

. ternata

. tetraspora

. tiroliensis

. torrendii

. frematostoma

. revoae

. trichostoma

. tumefaciens

. vagabunda

. vagabunda forma
daphnes

. vagabunda var.
divergens

. vitigena

. xylogena

. yerbae

ets

. almeidana

bella

. cistina

. coniothyrium

. corticola

. daphnes

. fusispora

. Juniperi

. lavandulae

. papillata

. platanicola

. rulingiae

Toa Soll Soll soll all all Soll soll wall cll all oll wall wall call oll all

sail Soll

Branch

all Soll all all Soll all Soll Soll Soll Sal all Sa

Broom

oat

. sorghophila
Canes
L. hendersoniae
L. hippophaes
L. hydrophila
L. micropogon
L. praetermissa
Capsules
. capsularum
. carpogena
. carpophila
. ogilviensis
. scrophulariae
Carpel

alll Soll wall soll Soll se

. triglochinicola
Cones

6

. pint

Culms

. aliena
.amphiloga

_ anthrostomella
. apogon

. arenaria

all Soll call Soll Se

Nomenclator of Leptosphaeria

[Soil all Soll all Soll call oll Soll all call wall all wall wall all wall call

é

é

Ioail Soll Soll Sail Soll Sal all Se

Toll Soll Soll Soll Soll Soll Soll Soll all Soll Soll call call Soll call call all all call

[all Soll Soll Sal Soll

. arrhenatheri

. arundinacea

. baldingerae

. bambusae

. bambusicola

. beaumontii

. bryzae

. Cattanet

. CIrsil-arvensis

. clavicarpa

. coccodes

. conimbricensis

. culmicola

. culmicola var. aquatica
. culmicola var. minor
. culmifida

. culmifraga

. culmifraga forma

majuscula

. culmorum forma

hungarica

. culmorum var. fla-

vobrunnea

. culmorum var. paleicola
. cumulata

. cynosuri

. dactylina

. discors

. donacina

. epicalamia

. epicalamia var.

pleosporoides

. eumorpha

. eustomella

. eustomoides

. graminis

. grisea

. hazslinskyana

. helianthi

. herpotrichoides
_holmii

. ischaemi

. Junct

. Juncicola

. Juncina

. Junci-acutt

. larvalis

. lelebae

. linearis

. lineolaris

. littoralis

. littoralis forma

calamagrostidis-
arenariae

_lolii

. luctuosa

. marina

. maritima

. matritensis

. melanommoides

lai

etl ce ell tl sell soll ell coll Stl sol cll coll a cll ool al oll oa Ul tll coll ool oll wall all call coll cl call call coll coll call coll coll call call call cal all coll all all call Call coll Soll all all all all all all all al ll

. michotii
. microscopica forma

brachypodii

. monilispora

. mosana

. moutoniana

. nardi

. neomaritima

_ nigrans forma arundinis
. nodorum

. occidentalis

. oryzae

. pachycarpa

. pachytheca

. papillosa

. patellaeformis

. pelagica

. penniseti

. perpusilla var. typhae
. petkovicensis

. petkovicensis var. elymi
. petrakit

. phacae

. phragmiticola

. pleurospora

. poae

. pontiformis

. proteispora

. punctiformis

. raphani

. reidiana

. rhodophaea

. riparia

. rubelloides

. FUSCL

.rusci forma caulina
. rusci var. hypophylli
. sabauda

. sanguisorbae

. Saxonica

. scabiens

. scabrispora

. scirpina

. secalina

. setulosa

. solheimii

. sowerbyi

. Sparsa

. Spartii

. Spartinae

. Spegazzini var. minor
. sporoboli

. Sticta

. Straminis

. stromatoidea

. subcompressa

. subsuperficialis

. taminensis

. therophila

. thurgoviensis

loath Sal all Soll all oa

. trichopterygis
. tucumanensis
. typhae

. variabilis

. volkartiana

. weddellii

Driftwood

Ie:
Exocarp
lie
Feathers
Ihre
Florets
Ib
Fronds

LL.
ie
iE
Ihe
Ib
Ls

Fruits

hot

Glumes
IE,

orae-maris
exocarpogena
corvina
lamprocarpi

aquilina
asplenii
caffra
elaeidicola
matisiae
pinnarum

. dryadis
. genistae
. pomona forma tran-

silvanica

oryzina

Hymenium

L.

consocians

L. fungicola

L.
L.

lichenicola
stereicola

Involucre

Ih
Leaves

ail Sail all Soll all Sol Soll Soll Sal all Soll all Soll Soll Soll Soll all call all call wal call call

involucralis

. abutilonis
-aceris

. acicola
.acorella
.acori

. aegira

. aeluropodis
. aerea

. agaves

. aglaja

. alcides

. alcides forma quercina
. alexandrinis
. algarbiensis
. algida

. aliena

. almeidae

_ aloes

. alopecuri

. alpiniae
.ammophilae
. anarithma

. anarithmoides
. andromedae

Illinois Natural History Survey Bulletin

Teall Soll Sol Soll all all Sa

SoU call Soll Soll all Soll Soll all Sal call Soll wall all call coll oll coll oll call coll oll coll call call coll all coll all call coll all coll oll wall coll oll coll all all coll all coll oll oll coll ll all coll all wall wall

. antarctica

. apios

. apios-fortunei

. aquatica

. arecae

. arenaria

. arenaria form sp.

triticea

. arrhenatheri var. italica
. asperellae

. aspidistrae

. auerswaldii

. australis

. austro-americana
. avenae

. avenaria

. bacillifera

. baldratiana

. bambusae

. bataticola

. batumensis

. baumii

. berberidicola

. betulina

. bicolor

. bispora

. bomareae

. bondari

. bornmuelleri

. brachyasca

. brachypodti

. brasiliensis

. briosiana

. buddlejae

. californica

. calopogonii

. camelliae

. camelliae-japonicae
. camilla

. camphorosmae
. campisilii

. canephorae

. cannabina

. cannae

. caricicola

. caricina

. caricinella

- caricis

. caricis-firmae
. caricis-vulpinae
. Cattanei

. cavarae

. ceballosi

. cecropiae

. cercocarpt

. cChamaeropis

. chenopodii-albi
. Chochrjakovii

. chusqueae

. citricola

Tall oll oall Soll Soll call wall Soll Soll wall all wa

all Sat Soll all all Soll call co oll wall wall

call all

ladl oad all Soll soll Sell all Soll Soll all Soll Sal cll Soll Sal all all Sn

‘ood sal soll soll Sal oo!

Vol. 34 Art. 3

. clara

. Clavata

. coccothrinacis
. COCOeS

. coffaeicida

. coffeigena

. coIcis

. collumiae

. colocasiae

. concentrica

. coniothyrium

. coniothyrium var.

foliicola

. consobrina

. consociata

. convallariae

. convallariae forma

dracaenae

. convallariae forma rusci
. cordylines

. corrugans

. coumarounae

. crastophila

. crastophila forma

tofieldiae

. cucurbitae
. culmicola var. hispalen-

SIS

. culmifraga var. alpestris
. culmifraga var.

propinqua

. culmorum
. culmorum forma epigeii
. culmorum forma

phragmitis

. cumana
. cycadis

. cynaracearum

. cynodontis-dactyli
. cypert

. cypericola

. daphniphylli

. dasylirii

. debeauxii

. decaisneana

. dennisiana

. densa

. diana

. dichosciadii

. dracaenae

. draconis

. dryadea

. dryadea subsp.

lussoniensis

. dryadis

. dryophila

. duplex

. eichhorniae
. elaeidis

. elaeospora

— --_—-_—_——-- -

March 1991

L. elymi

L. empetri

L. epicarecta

L. eriobotryae

L. erythrinae

L. eustomoides var.
punctata

L. faulii

L. feijoae

L. ferruginea

L. fici-elasticae

L. filamentosa

L. foliicola

L. folliculata

L. fraxini

L. frigida

L. fuegiana

L. fuscidula

L. fuscidula forma
magnolii

L. gaubae

. georgius-fischeri

. gigaspsora

. glandulosae

glauco-punctata

. glyceriae

. glyceriae-plicatae

gossypil

. graminum

grandispora

. gratissima

. gratissima var. longis-

pora

guazumae

gynerii

. hardenbergiae

hausmanniana

hausmanniana var.

cherleriae

hederae

. hedericola

. helicicola

. heloniaefolia

. helvetica

. hemicrypta

hermodactyli

hesperia

heveae

honiaraensis

huthiana

hyperborea

. hysterioides

immunda

. indeprensa

indica

. infernalis

. insignis

. insignis forma airae-

cespitosae

iridicola

Food oll soll oll soll soll soll oll call Soll Soll Soll Soll all Soll Soll Gall call Sad Soll all all Soll IEEE call oll all call oll all all all al all

Nomenclator of Leptosphaeria

oll soll soll all

Toll Soll Soll Soll wall coll all call all we

Pee Ee

Teall soll all Soll

[atl all Soll all Sal Sail all all all Soll all Soll all Soll call all call all call call call coll call call call cal all call

. iridigena

. iridigena var. typhae
. iridis

. isariphora

. wamotoi

Jahnii
Janus
jubaeae

Junci

. juncina forma macro-

spora

. Juniperina

kerguelensis

. kotschyana
. lacustris

larseniana
lasioderma

lathonia var. hellbori-
foetidi

lauri

. leersiae

. leersiana

. licatensis

. licatensis forma

rupefortensis
lilii

. lingue
. livida

lobeliae

. lucilla

lucina

. luzulae

. maclurae

. macrochloae
. macrotheca
. maculans

. magnoliae

. Mmagnusiana

major

. malojensis
. mandshurica

marantae
marcyensis
marginalis
marginata

. maritima

matritensis
maydis
media
melicae

. mellispora

. mendozana

. mertensiae

. microscopica subsp.

calami

. microscopica Var. alpina
. mikaniae

. minima

. miyakeana

. molleriana

Ww
nN

. monticola

. morthieriana

. moutan

. mucosa

muirensis

muricata

musae

musarum

musigena

nardi var. dubiosa

nashi

. neglecta

. nervisequa

nesodes

. niessleana

. nobilis

. nodorum form sp. hordei

. obtusispora

. obtusispora forma
agaves

. occulta

. oligotheca

. ophiopogonis

. ophiopogonis var.
graminum

. orthrosanthi

oryzae

. pachyasca

. pachytheca

. pacifica

L. paludosa

L. pampaniniana

L. pandani

L. pandanicola

L. panict

L. paoluccii

L. papulosa

L. paraguariensis

L. parvula

L. parvula vat. iridis-

germanicae

L. passerinit

L. pelargonit

L. penniseticola

L. perforans

L. petri

L. phlogis

L. phoradendri

L. phormii

L. physalidis

L. pilulariae

L. piperis

L. plantaginicola

L. plemeliana

L. pomona

L. portoricensis

L. primulaecola

L. priuscheggiana

L. protearum

L. pruni

fool call all call oll coll wall Soll all all ol call all oll oll oll oll all

hoof

Sol coll coll coll

322

L. pruni forma plurivora

L. pterocelastri

L. puccinioides

L. puiggarit

L. pulchra

. punctillum

. purpurearum

. pusilla

. puttemansit

. recutita

. rehmiana

_rehmit

. rhododendri

. rivalis

. rostrata

. rostrupu

. FUSCI

. rusci forma fourcroyae

. USCI Var. FUSCI-
hypoglossi

. ruscicola

. Sabauda

. sabauda forma arvaticae

. sabuletorum

. sacchari

. saccharicola

. Saginata

. salebricola

. Salicinearum

. Sasae

. scolecosporarum

. seriata

. Silenes-acaulis

. smarodsit

. smilacis

. solani

. sorbi

. sorghi-arundinacet

. Sparsa

. Sparsa var. meizospora

. Spegazzini

. Staticicola

. Stellariae

. Stipae

. Stipae-minor

. Stratiotis

. Striolata var. caricis-
glaucae

. subiculifera

. substerilis

. Subtecta

. Swertiae

. sylvatica

. taichungensis

. taiwanensis

. tanaceti

. taxicola

. theobromicola

. thorae

. thujaecola

wielel eal aial al olal ale al alae ala alalalel alee al olelelalolalalolole lolol,

Teall Soll all Soll Soll all Soll all Soll Soll soll

Illinois Natural History Survey Bulletin

fall all oa

featl soll Soll Soll Soll all Soll Soll Soll col all call ll oll oll ol oll all oll al lll a

Nuts

reel

Paper

Toot Soll all sa

Pedicels

Ib
E..

Toot Soll Soll call all soll wal

tint

. tofieldiae

. tonduzi

. translucens

. rrifolii-alpestris
. tritict

. typharum

. typharum subsp.

phragmatina

. typhicola
. typhiseda
. typhiseda forma

sodoloci

. uliginosa

. ulmicola

. vagans

. vagans forma scirpi
. valdobbiae

. vanhoeffeniana
. varliseptata

. vectis

. verwoerdiana

. vincae

. vinosa

. vrieseae

. weberi

. williamsii

. wintert

. woodrow-wilsonii
. xerophylli

. xiphii

. yulan

. zeae

. zeae-maydis

. zeicola

. zingiberis

. cacuminispora

. fibrincola

. papyricola

. (ritict var. papyricola
. typharum subsp.

papyrogena

eranthemi
eustoma

Peduncles

L.
Es.
E.
. lycopodiicola
. papyri

. raphidophora
. scapophila

. typhiseda

. vitensis

bryzae
espeletiae
francoae

. anemones
. cinclidoti

Pods

Vol. 34 Art. 3

. eryngil

. monticola

. petiolaris

. petiolicola

. phoenicis

. primulaecola
. typhiseda

call soll all all all all

. endiusae

. eustoma forma legumi-
nosa

. Impressa

. lunariae

. lyndonvillae

ho

hot

Rachis

L. culmifraga forma poae
L. eustomoides forma lolii
L. sabalicola

L. trochus

Rhizomes

Seeds

Sepals

L. culmicola var. rhizoma-
tum

. heterospora

. littoralis

. pontiformis

. rhizomatum

. vagabunda subsp.
alvarensis

loath Soll all Soll a

L. australiensis
L. capparidis
L. circinans
L. cladti

L. lycii

L. maculans forma

denudata
L. septovariata

. cookei

. grignonnensis
. haematites

. nectrioides

. notarisii

. pampini

. pleosporoides
. rhizomatum

. rimalis

. Sarmenticia

. thomasiana

. vagabunda var. sarmenti
. vinealis

. vitalbae

[aol all Soll all Soll ool ll Soll all all Soll Soll snl ol

. albo-punctata

. microscopica forma
glyceriae

L. sequana

L. woroninii

(alll

L. auerswaldii
L. sepalorum

March 1991

Sheaths

E:

hot

[optl call Soll Soll Soll all Soll Soll all Soll oll Soll all call coll oll wall oll oll coll o

altaica

. arenaria
. avenaria form sp.

triticea

. cariciphila

. cattanei

. cookei

. culmicola forma melicae
. culmicola var. hispalen-

sis

. culmicola var. nigrans
. culmorum
. fuckelii forma filamentif-

era

. interspersa
. korrae

. lagenoides

. latebrosa

. matritensis
. media

. monilispora
. narmari

. neomaritima
. nigrans

. nodorum form sp. hordei
. notarisii

. pandani

. rivalis

. rousseliana
. sacchari

. Salvinit

. Spartit

. Spegazzini

. subalpina

. vaginae

Sporangia

Stems

1 oF

L
L
L
E
L
L
Ie
E
L
E;
E.
Z,
18
L
iE
L
L
I
L
E
L
E.

heufleri

. abbreviata

. acanthi

. achilleae

- aconiti

. aculeorum

. acuta

. acuta forma insignis
. acuta forma urticae
. acutispora

. acutiuscula

affinis

. agminalis

. agnita

. agnita subsp. labens

. agnita var. ambigua
_agnita var. bupleuri

. agnita var. chrysanthemi
. agnita Var. erigerontis

. agnita var. major

. agnita var. trifolii

. albulae

alliariae

Nomenclator of Leptosphaeria

Sivlelvinlelalalelalalalalelalalelalelelalalelalalolelalelalalolelalelalalelalelalelalalelalolelelalelelalalolelela lola lo

. allorgei
.amphibola

. anacycli

- anarrhini

. andrijevicensis
.anomala

. anthelmintica
. antherici

. anthostomoides
. aparines

. aquilana

. aquilegiae

. arabidis

. argentina

. artemisiae

. arthrophyma
. AFUuNcl

. arvensis

. asclepiadis

. asparagi

. asparagina

. astericola

. asteris

. atropurpurea
. aucta

. aulica

. balcarica

. ballotae

. bardanae

. bellynckii

. berlesei

. blumeri

. bocconiae

. boucera

. brachyasca

. brachysperma
. brauni

. bresadolaeana
. brightonensis
. brunellae

. bryophila

. bubakii

. buddlejae

. bulgarica

. bupleuri

. caballeroi

. cadubriae

. Caespitosa

. caespitosa forma salviae
. calvescens

. camphorata

. canadensis

. carduorum

. carlinoides

. carneomaculans
. cassiaecola

. castilleiae

. castillejae

. caulincola

. centaureae

. centrafricana

Teall Soll all soll call call all Soll wall call a ll

oil ool soll all

[eal soll Soll Soll Soll Soll So

all Soll Soll Soll Soll cll coll all call Goll coll call Soll all ll coll coll oll oll all coll oll ll oll oll oll oll ll oll ll all

. cephalariai-uralensis
. cerastil
. ceratispora

cerei-peruviani

. cervispora

. cesatiana

. chelidonii

. chrysanthemi
. cibostit

. circinans

. cirricola

. cladophila

. clavata

. clavigera

. clavispora

. clivensis

. clivensis var. constricta
. coleosanthi

. collinsoniae

. comatella

. complanata

. compositarum
. compressa

. concinna

. conferta

. congesta

. coniformis

- cont

. coniigena

. coniothyrium
. conoidea

. conoidea forma

angelicae

. conoidea forma asteris
. conoidea forma

macrospora

. CONSeSSA
. controversa

. convallariae

. coorgica

. corallorhizae

. cornuta

. coronillae

. corticola

. corynispora

. cosmicola

. crastophila forma

tofieldiae

. cruenta

. crustacea

. cucurbitarioides

. culmicola forma major
. culmifraga var.

bromicola

. culmifraga var. linearis
. curta

. cylindrospora

_ cylindrostoma

. cynoglossi

. cynops

. darkeri

324

all all ell all all ell al ala l el ella lel eel ela l elo lo

fai

hoof

le [es

oil

Wes ites teste

. davidii

. davisiana

. dearnessii

. deficiens

. delawayi

. dematiicola

. dematium

. demissa

. depressa

. derasa forma alpestris
. derasa forma robusta
. derasa vat. franconica
. desciscens

. dianthi

. diaporthoides

. digitalis

. dissiliens

. dolioloides

. dolioloides var. cirsti

. dolioloides var. inops
. dolioloides var. lathyri
. dolioloides var.

rhinanthi

. doliolum
. doliolum forma

carlinae-vulgaris

. doliolum forma

syndoliola

. doliolum var. angustis-

pora

. doliolum var. cacaliae
. doliolum var. dissimilis
. doliolum var. leonuri

. doliolum var. pachy-

spora

. doliolum var. pinquicula
. doliolum var. subdisti-

cha

. drabae
. dryadea subsp.

lussoniensis

. dryadis

. dubia

. dumetorum

. dumetorum forma ebuli
. dumetorum forma

meliloti

. dumetorum var.

coniformis

. dumetorum var.

coronillae

. dumetorum var. galii-

borealis

. dumetorum var.

marrubii

. dumetorum var.

symphyti

. eburnea
. echiella
-echii

Illinois Natural History Survey Bulletin

. echinella
. echinops
. ellisiana
. elongata
. endiusae
. epilobit
. equiseti
. equiseticola
. erigerontis
. eriophora
. eryngil
. ettalensis
. euphorbiae
. euphorbiae forma esulae
. euphorbiaecola
. eutypoides
L. faginea
L. fallaciosa
L. fallax
L. ferruginea
L. ferulicola
L. filiformis
L. fimbriata
L. fiumana
L. foeniculacea
L. foeniculacea subsp.
lupina
L. foeniculi
L. fraserae
L. frondis
L. fuckelii
L. fuegiana
L. fulgida
. galeobdolonis
. galeopsidicola
. galii
. galiicola
. galiicola var. brachy-
spora
L. galii-silvatici
L. galiorum
L. galiorum torma
cirsiorum
L. galiorum forma
gentianae
L. galiorum subsp.
antirrhini
. galiorum var. gnaphali-
ana
. galiorum var. lapsanae
. gaultheriae
. genistae var. microspora
. gigaspora
. gloeospora
. gnaphalii
. grammodes
. gypsophilae
. haematites
. harknessianna
.helianthemi

Fel ll all oll Soll Soll Soll all Soll call all all wall wall ull

Tall Soll Soll Soll

ld

all Soll all Soll all Soll all Soll Soll all Se

Vol. 34 Art. 3

. heliopsidis

. helminthospora forma
crithmi-maritimi

. hemerocallidis

. hesperidicola

. hiemalis

_hirta

. hollosiana

_holmii

. houseana

. hrubyana

. hurae

. hyalospora

. hyparrheniae

. hyperict

. Icositana

. incarcerata

. Incruenta

. inculta

. indica

. Insulana

L. jaceae

L. jacksonensis

L. jacksonii

L. johansonii

L. juncaginearum

L. junci-glauci

L. juncorum

. kali

. kalmiae

. kalmusti

. kochiana

. kunzeana

. lacustris

. ladina

. lankeana

. larseniana

. lasiosphaerioides

. lassenensis

. lathyri

. lathyrina

. lecanora

. leptospora

. lespedezae

. lethalis

. libanotis

. limosa

. lindquistii

. lithophilae

. lobayensis

. longchampsi

. longipedicellata

. longispora

. lonicerina

. lophanthi

. lupini

. lupinicola

. lusitanica

. lyciophila

. lythri

ail

Teall soll Soll Soll Soll coll coll col call oll all call wall call call ll wall oa

[all Soll Soll Soll all all all Soll Sal Sal all all voll Soll Soll Soll all all call wall all oll Soll cll call voll wall call oll all call on

March 1991

f

[eoil coll soll Soll Soll Sain coll Soll all all soll all Soll all Soll wa

sell Soll all Soll Soll Sail all Sal Sal Sal a

[oo tl all call all Soll all all Soll Soll Soll all all Soll Soll wall call Soll all all wall wall oll all all call wall call

. macrospora

. macrosporidium

. maculans

. maculans var. typhicola
. maderensis

.malyi

. maritima

.marram

. martagoni

. massariella var. disticha
. massarioides

media

. medicaginicola
. medicaginis

. medicaginum

. megalospora

. melanommoides
. melicae

. meliloti

. menthae

. mesoedema

. Metasequoiae

. microspora

. microthyrioides
. millefolii

. mirabilis

. modesta

. modesta forma digitalis-

luteae

. modesta forma jaco-

baeae

. modesta forma lappae

. modesta forma succisae
. modesta forma sylvestris
. modesta var. cibostii

. modesta var. rubellula

. modestula

. molluginis

. molybdina

. monilispora

. monilispora forma

triglochinis

. montana

. montis-bardi

. morierae

. muehlenbeckiae

. multiseptata

. multiseptata forma

alpina

. muralis

. muscari

. nanae

. napelli

. napti

. nicolai

. niessleana

. niessleana forma viciae
. niessleana var. Staritzil
_nigrella

. nigricans

Nomenclator of Leptosphaeria

feat ool call call Soll oll Sal call coll Soll all wall co all call wa

las

é

Toot coll soll Soll Soll Soll Soll Soll Soll Soll all Soll soll all call all

Toll all Soll Soll Soll all Soll call Soll call all oa

. nigromaculata
_ nitschkei
. nitschkei forma

adenostylidis

. norfolcia

. norvegica

. obesa

. obesula

. obiones

. obiones var. evolutior
. ocellata

. ocimicola

. octophragmia

. octophragmia var. major
. octoseptata

. ogilviensis

. ogilviensis forma

achilleae

. ogilviensis forma lepidii
. ogilviensis forma

megalospora

. ogilviensis forma

myrrhis-odorata

. ogilviensis var.

pleurospermi

. ogilviensis var.

senecionis-cordati

. olericola

. olivacea

. onagrae

. onobrychidicola
. onobrychidis

. ophioboloides

opizit

. opuntiae

. orchidearum

. oreophila

. oreophiloides

. oreophiloides subsp.

scrophulariae

. ornithogali

. orthogramma

. oubanguiensis

. owaniae

. oxyriae

. oxyspora

. palustris

. papaveris

. parietariae

. parietariae forma lamii
. passeriniana

. passerinit

. pellita

. penicillus

. perplexa

. perpusilla

. personata

L.
L.
E.

phaeospora
phaseoli
phaseolorum

Ee.
. phyteumatis
. phytolaccae
. pimpinellae
. planiuscula
. planiuscula forma

ool soll soll Soll

NN etl el etl et ll Bl ol co coll a aa all oa oll oll cal ca wall call coll call call call call coll oll col all Call all all wall all Sal all all all all all

[all Soll Soll all Sol all all

fen

Ww
i)
a)

physostegiae

cruciferarum

. platanicola

. platypus

. plectrospora

. plumbaginis

. plurisepta

. politis

. polygonati

. pomiformis

. poterii

. praeclara

. pramontorii

. pratensis

. primulana

. proliferae

. pseudo-diaporthe
. pseudohleria

. psilospora

. ptarmicae

. pulchra

. punctoidea

. punjabensis

. purpurea

. quadriseptata
. quamoclidii

. ramsaugiensis
. ranunculi

. ranunculi-polyanthemi
. ranunculoides
. recessa

. resedae

. Festionis

. rhodiolicola

. rhopalispora

. rhopographoides
. richoni

. riofrioi

. rivalis

. rivularis

. robusta

. rostrupit

. rothomagensis
. rothomagensis var.

artemisiae

. rubellula

. rubicunda

. rudbeckiae

. rugosa

. rumicicola

. rumicis

. ruscicola
_ruscicola forma

cladodiicola

_ russellii

326

Sal al al al al al al ol al al al al al al al al al al ala! al al al al al al al al al al al al al al al al al al ol al al al al al aio ol al al nl ai aioli sl ol alain) a

. rustica
. ruthenica
. saccardiana

sacculus
salebrosa

. Salicaria

salsolae
sanguisorbae
Sapeyensis
Sarraziniana
schoenocauli
scitula
scotophila
scutati
semelina
senecionis
sepincola

. septemcellulata

serbica
setosa
shahvarica
shastensis
sibirica
sibtorpii
sieversiae
sileris

. silvestris

simmonsit
sinapts

. Slovacica

socialis
sodomaea
solani
solanicola

. solidaginis

Sparganit
spectabilis
Staritzi

. Steironematis
. stellariae

stellata
Stictostoma
striata
striolata
subcaespitosa
subconica
submaculans
submodesta
subriparia
suffulta

. surculorum

tanaceti
taurica
tenera
tenuis

. tephrosiae

. tetonensis

. teucrit

. thalictri

. thalictricola

Illinois Natural History Survey Bulletin

Si

ho

|bp
L.
L.

Stolons

Straw

Stroma

cE.
|b
IE
L.

Si alal al al al al al al al al al al al Sia al ani aia

loath call call call soll call wall coll oll call oo allo

. thalictrina
. thielensii

thuemeniana
tolgorensis

. torbolensis

torulispora
trichostoma
trifolit

. triglochinicola

triglochinis
trimera

. trimerioides
. tritorulosa

troll
tupae
typharum
uncinata
utahensis
vagabunda

. vagabunda forma abietis
. vagabunda forma citri-

limonti

. vagabunda forma

lonicerae

. vagabunda forma

Salicis-capreae

. vagabunda var. caulium
. vagabunda var.

dulcamara
vahlii

. valdiviensis
. valesiaca
. variegata

veratri

viciae
vindobonensis
vinosa

. virginica
. viridella
. vitalbae var. sarmenti-

cola

. Wegeliniana
. wegeliniana forma

teucrii
wehmeyeri
zahlbruckneri
zizaniaecola

. cookei
. korrae
. narmari
. notaristi

. culmifraga
Ibe

eustoma

caucana
cryptica

phyllachoricola
phyllachorivora

Stubble

roost

Thalli

Trunks

aia

Twigs

Si al al al al al al al al al al ol al al al al aial al al ol pial al

oll soll all Sal soll all Soll all Soll So

all soll all Soll oa

(soll soll soll wall Soll

Vol. 34 Art. 3

. platychorae
. tungurahuensis

. arundinacea

. arundinacea var. godini
. Cyperina

. recutita

advenula

. apocalypta

arnoldit
baeomycearia
caninae
clarkii
consocians
corae
crozalsii
galligena
geographicola
koerberi
lemaneae
leptogiophila
leucomelaria
lichenicola

. mamillula

mirandae
oligospora

. peltigerarum

. peltigerea

. polaris

. porellae

. pycnostigma

. pycnostigma vat.

morbosa

. ramalinae

rivana

. rivana forma solorinae

sphyridiana
Steinil
stereocaulorum

. subarticulata
. tartarina

. umbilicariae
. usneae

buxina

. coffeigena var. longiros-

trata
ericae

. hottai

. martinianum
. Salebrosa

. sicula

. bicuspidata
. bondari

. californica
. calligoni

casta

. ceanothi

March 1991 Nomenclator of Leptosphaeria 327

L. distributa Vines L. borealis
L. fuscella L. ampelina L. calligoni
L. hispanica L. caprifolii L. clerodendri
L. hyalina L. cerlettii L. contecta
L. leiostega L. chaetostoma L. halima
L. myricae L. pampini L. haloxyli
L. periclymeni var. tatarica L. phaseolorum L. isocellula
L. plagia L. socia L. paucispora
L. pyrenopezizoides L. viticola L. picastra
L. ramulicola L. vitis L. plocamae
L. rubrotincta Wood L. puteana
L. tornatospora L. abuensis L. rajasthanensis
L. vitalbae L. argentinensis L. rhoina

L. seminuda

Geographic Index

Afghanistan

L.

Algeria

Jos fest test jest test (est lest let lest Le

Angola

Ib

morierae

. crozalsiana

. debeauxii

. lcositana

. Indeprensa

. macrochloae
. obesa

. papulosa

. phiala

. Stipae

. typhiseda

baumii

Argentina

Toll Soll Sol Soll oll all Soll Sol Soll all Soll Soll Sal all Soll oll ual wall coll Soll Soll call call wall wall all all call call wall

. antarctica

. adesmicola

. aerea

. anthostomella

. argentina

. argentinensis

. austro-americana
. basalduai

. berberidicola

. CON

. conligena

. cordylines

. cylindrostoma

. fuegiana

. gynerit

. lagenoides

. lindquistii

. lycopodiicola var. major
. mendozana

. obtusispora

. preandina

. promontorit

. proteispora

. Sacchari

. Spegazzini

. Spegazzini var. minor
. Subiculifera

. subsuperficialis
. tucumanensis

. vinosa

. yerbae

Australia

E;

aliena

etl all all all Soll ll all Soll soll oll wall oll Soll call ll wall oa

Austria

foodl oall soll Soll all soll all Soll Soll all Soll all coll Soll all call call call Soll call cal call call coll all wall allo

é

. australiensis
. australis

. camelliae

. cannae

. clelandii

. daviesiae

. dichosciadti
. gaubae

. hardenbergiae
. korrae

. Narmari

. paucispora

. plagia

. restionis

. rulingiae

. suaedae

. williamsii

. acuta

. aparines

. artemisiae

. baggei

. caricis-firmae

. cattanei

. cesatiana

. cinerea

. coniformis

. coniothyrium

. corticola

. culmifraga var. alpestris
. depressa

. derasa forma robusta
. eustoma

. fusispora

. gnaphalii

. Inconspicua

. intermedia

. juncicola

. macrospora

. marginata

. medicaginis

. medicaginum

. monilispora
.napi

. nigrans

_ nitschkei forma

adenostylidis

. oreophila
. parvula

io]
is

Si

=
Toil nll all nll oll all nll all all Sal Sal = 101 Soll Soll Soll all all Soll Soll Soll Soll wall Soll all wall call call wall all call all call wall call

o

(oad all sol soll nll soll ll oall Soll Soll all sal od

. penicillus

. personata

. pleurospora

. pulchra

. recutita

_ TUSCI

. saprophila

. Scirpina

. seminuda

. Senecionis

. Sparsa var. elynae
. Spectabilis

. Sphyridiana

. suffulta

. tanaceti

. thorae

. tiroliensis

. umbrosa

. vindobonensis
. vitalbae

. vitigena

. Vitis

. zahlbruckneri

. acuta

. albo-punctata
. arenaria

. bellynckii

. carduorum

. crepini

. doliolum

. endophaena

. gnaphalii

. haematites

. Juncina forma macro-

spora

. libanotis

. longchampsi

. maculans

. mamillana

. melanommoides
. michotii

. mosana

. moutoniana

. mucosa

. nardi

. nardi var. dubiosa
. obesula

. perpusilla

March 199]

L. punctiformis
. revocans

. rivularis

. rusci

. salebricola

. sowerbyi

. subriparia

. thielensii

. typharum

Teall Soll Soll Soll Soll Soll Soll

Bolivia

L. agnita var. acheniarum
iE

. weddellii
Brazil
. alpiniae
. bondari
. brasiliensis
. desmonci
. diaporthoides
. eustomoides var.
punctata
. massariella var.
brasiliensis
. matisiae
. mikaniae
. paraguariensis
. pelargonii
. puiggarii
. puttemansit
. saccharicola
. Schneideriana
Bulgaria
L. bubakii
L. bulgarica
L. dianthi

Sa all all Soll coll all

ool call all Soll Soll all all Sa

L. andromedae
L. anisomeres
L. asclepiadis
L. associata

L. australiensis
1G,

. avenaria form sp.

triticea

. barriae

. berlesei

. brunellae

. canadensis

. caricinella

- Caricis

. corticola

. culmifraga forma
minuscula

. culmorum

. dearnessii

. doliolum )

. drechsleri

. elongata

. equiseti

. eustoma

L. faulii

L. folliculata

L. fuckelii

Teall Soll Soll call Soll Soll Soll

all soll soll all Soll all

Nomenclator of Leptosphaeria

. gaultheriae

. hesperia

. hiemalis

. Insignis

. Jacksonii

. licatensis

_ lilii

. luctuosa

. lycopodiicola
. lycopodina

. marcyensis
.marram

. microscopica
. microspora

. neomaritima
. nigrans

. nodorum

. ogilviensis

. punctillum

. rousseliana

. russellii

. rugosa

. solani

. sorgho-arundinacei
. Steironematis
. typharum

. typhicola

. waghorniana
. wehmeyeri

oll Soll all Soll Soll all Soll oll ll oll oll oll all ol ll ll ll olla olla ol lal oll all olla

Canary Islands

L. plocamae

Cape Verde Islands

L. larvalis

Central African Republic

. centrafricana
. cylindrospora
. excelsa

. gigaspora

. lobayensis

. longispora

. macrorostra

. oubanguiensis
. tetraspora

atl Sil Soll all Soll Soll all Sal!

Central America

Chile

China

L. corae

L. chilensis

L. flotoviae

L. francoae

L. jubaeae

L. lingue

. melanommoides
. phoradendri

. trevoae

. tupae

. valdiviensis

oll soll soll Soll

- acanthi

. aspidistrae

. bambusicola
. deficiens

Soll soll soll we

allo

Teall Soll Soll Soll Soll Soll Soll Se

329

. delawayi
. doliolum var. angustis-

pora

. eranthemi

. mandshurica
. miyakeana

. oryzae

. papyrt

. plumbaginis
. scabrispora
. trochus

Colombia

ES
ES

caucana
stellata

Costa Rica

Er
[
Jb.

Cuba

Jb

coffaeicida
pusilla
tonduzi

coffeigena

Czechoslovakia

Topil Soll Soll Soll all all oll all Soll Soll all Soll all all Soll all call Soll call wall Soll all call Soll all coll call call call col oll coll col all call alll call all

. aucta

. baggei

. caricis

. carneomaculans
. conferta

. corticola

. culmorum

. cylindrospora
. cynaracearum
. dumetorum

. dumetorum forma ebuli
. euphorbiae

. fici-elasticae

. galeopsidicola
. gigaspsora

. heterospora

. hrubyana

. Juncicola

. juncina

. kalmusii

. koerberi

. lycopodina

. marginata

. megalospora

. millefolii

. niessleana

. parvula

. petkovicensis

. petrakit

. phyllachorivora
. poae

. pontiformis

. rousseliana

. rudbeckiae

. setosa

. Slovacica

. Steinii

. typhicola

. uncinata

Denmark

.ammophilae
. arenaria

. bacillifera

. berlesei

. bispora

. chondri

- CON

. corvina
_culmorum

. danica

. dianthi

. marina

. microscopica
.nodorum form sp. hordei
. occulta

. oligotheca

. recutita

. rostrupit

. sSabuletorum

atl col call call call call call all oll ol olla lol lle ll on

Dominican Republic

. calopogonii

. cecropiae

. coccothrinacis
. coumarounae
. eichhorniae

. guazumae

. smilacis

. theobromicola

ROR EERO

East Africa

L. piperis

Ecuador

L. bomareae

L. consociata

L. phyllachoricola
L. plantaginicola
L. saginata

Ethiopia

L. baldratiana

Europe

L. auerswaldit
L. phaeosticta
L. pomiformis

-acort

acuta

. affinis

. ammophilae

. aSparagina

. coccodes

. culmicola var. nigrans

. culmifida

.culmorum

. dolioloides

. dolioloides var. cirsti

. dolioloides var. inops

. doliolum

. elongata

. fuckelii forma filamen-
tifera

a TE Be a to oT Tl el Bo tll cll

Illinois Natural History Survey Bulletin

Teall Soll Soll Soll Soll Gall all oo

all soll call Soll all

[atl Soll all call Soll coll cal call wall Soll coll Soll call oa

é

France

Soll all call Soll Soll Soll all call Soll all Soll Soll all Soll Soll call Soll all Soll all call Soll Soll coll Soll oa

. herpotrichoides

. immunda

. leucoplaca

. lycopodina

. maculans

. maculans var. typhicola
. microscopica

. microscopica subsp.

calami

. modesta

_ nigrans

. nigricans

. oligospora

. orchidearum

. oreophiloides subsp.

scrophulariae

. perpusilla var. typhae
. planiuscula

. praeclara

. praetermissa

. ptarmicae

. punctoidea

. thodiolicola

. ribis

. rustica

. socialis

. Spiraeae

. typhae

. typharum

. typharum subsp.

phragmatina

. vagabunda var.

dulcamarae

. acuta

. aCUta Var. insignis

. acuta Var. urticae

. agminalis
.agminalis forma minor
. ailanthi

. allorgei

. arenaria

. arundinacea

. arundinacea var. godini
. avellanae

. baldingerae

. bambusae

. berberidis

. bupleuri

. calvescens

. caprifolii

. carduorum

. caricicola

. Castagnei

. chelidonii

. cinclidoti

. cisti

. Clavata

. coniothyrium

. coniothyrium forma

berberidis

oath Soll all Soll Sa all all oll So Sol oll call co coll coll oll oll oll coll oll oll

ll

lal

~

‘sell onl all Soll all all oat

all soll all call call ll Sal all sal o

Vol. 34 Art. 3

. conoidea forma

angelicae

. conoidea forma

macrospora

. controversa

. convallariae

. crozalsit

. cCruenta

. cucurbitartoides
. culmicola

. culmicola forma

melicae

. culmicola var. aquatica
. culmicola var. nigrans
. culmicola var. rhizoma-

tum

. culmifraga

. culmifraga forma poae
. culmifraga var. linearis
. culmifraga vat.

propinqua

. curta

. cynops

. decaisneana

. derasa forma macro-

spora

. digitalis

. dioica

. dolioloides

. doliolum

. doliolum var. pachy-

spora

. doliolum var. pinquicula
. dumetorum var.

marrubii

. elaeospora
. emiliana

. empetri

. ephedrae

ericae

. eryngii

. euphorbiaecola

. eustomoides forma lolii
. foeniculacea

. fuscella forma micro-

spora

. galiorum
. galiorum forma

cirsiorum

. galiorum forma

gentianae

. galiorum var. lapsanae

. genistae Var. microspora
. gillotiana

. glauco-punctata

. grignonnensis

. haematites

. hausmanniana var.

cherleriae

March 1991

hop

CoM

lool oll wall all soll coll coll coll oll ol oll

POORER BR BARR nee

[all Soll Soll Soll Soll Soll oll all coll wall wall wall wo oll alo!

é

. hedericola
.helicicola
. helminthospora forma

crithmi-maritimi

. hippophaes
. inculta
. insignis forma airae-

cespitosae

. Iridicola

. iridigena

. isariphora
. Juncorum
. Juniperi

_ kali

. lathyri

. lauri

. lecanora

. lemoinii

. libanotis

. licatensis forma

rupefortensis

. limosa

. linearis

. lonicerae

. maculans

. maculans forma

denudata

. melanommoides

. menthae

. michotii

. microscopica forma

brachypodii

. microscopica forma

glyceriae

. minima

. modesta

. modesta forma dauci
. modesta forma digi-

talis-luteae

. modesta forma jaco-

baeae

. modesta forma lappae
. modesta forma succisae
. modesta forma

sylvestris

. modesta var. rubellula
. molybdina

. montis-bardi

. muelleri

. muralis

. nardi

. nigrans

. nigrans forma arundinis
. nigrificans

. norfolica

. obesula

. obiones

. obtusispora forma

agaves

. ogilviensis forma lepidii

Nomenclator of Leptosphaeria

BRR SERRE RR PPP PP eh Pe eB PR RRR RRR RR RRR

a)

. ogilviensis forma

megalospora

. ogilviensis forma

myrrhis-odorata

. ophiopogonis var.

graminum

. oreophiloides
. pachytheca

. pampini

. papyricola

. parietariae

. parietariae forma lamii
. perforans

. perpusilla

. phaseoli

. phragmiticola
. picridis

. pint

. pleosporoides
. polytrichina

. pratensis

. priuscheggiana
. pycnostigma

. ramalinae

. rhopalispora

_ richoni

. rothomagensis
. roumegueri

. rubellula

. FUMICIS

. FUSCL

. rusci var. hypophylli
. ruscicola

. ruthenica

. Sabauda

. Saccardiana

. Salebrosa

. Sambuci

. Sapeyensis

. Sarmenticia

. Sarothamni

. Sarraziniana

. scrophulariae
. Sequana

. smilacis

. Solidaginis

. Stellariae

. CeuCriL

. therophila

. thomasiana

. typharum

. typharum subsp.

papyrogena

. typhiseda forma

sodoloci

. vagabunda forma

abietis

. vagabunda forma

caulium

. vagabunda forma

lonicerae

é

Teall Soll all oo

331

. vagabunda forma
salicis-capreae

. vagans forma scirpi

. vincae

. viticola

. Vitis

French Equatorial Africa

ool all wall

Germa

=]
See

PRR RR ROR BRP Pe Pee Pe hehe

all all oll all all oll all lal

. heveae
. Septovariata
. Zeae-maydis
. zeicola

. achilleae

. acicola

. acuta

. acutiuscula

. agnita var. ambigua

. agnita var. chrysanthemi

. agnita var. major

. alliariae

. ammophilae

. anceps

. aquilegiae

. arabidis

. artemisiae

. asplenii

. atropurpurea

. avenae

. baggei

. bardanae

. bellynckii

. berlesei

. Caespitosa

. carduorum

. circinans

. clara

. clivensis

. coniothyrium

. contecta

. corticola

. culmorum var. paleicola

. cumulata

. dematiicola

. densa

. dolioloides

. doliolum

. dumetorum forma
meliloti

. duplex

. endiusae

. epicalamia

. ettalensis

. fibrincola

. fimiseda

. frondis

. fuckelii

. fuscella var. sydowiana

. galii-silvatici

. geographicola

. graminis

332

el

[oall Sail Soll Soll call all Soll Soll Soll Soll call call Sal

Kaan eilentlea i eats Ms tonic iilesbesil sale est leatlens| silt)

PRR E EE eee

[opi Soll all Sol all Sail Soll Soll Se

. graminum
haematites
hazslinskyana
helianthemi
helminthospora
hippophaes
hirta

huthiana
hyperici
impressa
incruenta
isariphora
Juncaginearum
Juncina
koerberi
kunzeana
lacustris
lemaneae
leptogiophila
limitata
littoralis forma
calamagrostidis-
arenariae

lolii

. lonicerae
. lonicerina

maculans
malojensis
medicaginis
medicaginum

. michotii

microscopica

. microscopica vat.

caricis-vulpinae

. multiseptata

nardi

. niessleana vat. staritzii

nigrans
nigrella

. nigromaculata

. nitschkei

. ogilviensis

. ogilviensis forma

achilleae

. ogilviensis var.

pleurospermi

. osculanda

. papillata

. penicillus

. petkovicensis
. pilulariae

. planiuscula

. pontiformis

. primulana

. psilospora

. purpurearum
. quinta

. ramsaugiensis
. recutita
_rostrupit

Illinois Natural History Survey Bulletin

Ghana

Ib

[atl Soll Soll Gall coll coll soll all call call wall wall all wall call wall call call wall

. rousseliana

. Salebricola

. salebrosa

. sanguisorbae

saxonica

. scitula

. Seminuda

. setulosa

. Sphyridiana

Staritzit

. Steinit

. Striata

. submaculans
. surculorum

. sydowiana

tanaceti
typhae

. typharum
. vincae
. winteri

Sparganit

Great Britain

‘oll Soll Soll Soll all all all Sal Sal Soll all all all Sail Sal Sal Sal Sal all all Soll Sail Soll all all all call Soll Sal Soll all call all Soll wall allo

. abbreviata
.acorella

acuta

. anarithma
. arundinacea

baeomycearia
caninae
carduorum
cesatiana
clara
clarkti
clivensis
complanata
cookei
dennisiana
derasa
dioica

. doliolum

duplex

. echinella

epicarecta

. fluviatilis
. fuscella
. glauco-punctata

gloeospora

. graminis

. hederae

. Juncina

. leucomelaria
. lunariae

. macrosporidium
. maculans

. maritima

. marram

. michotii

. mosana

nardi

Vol. 34 Art. 3

neomaritima

. neottizans

. niessleana forma viciae
nigrans

norfolcia

obiones var. evolutior
octophragmia var. major
. ogilviensis

. parmeliarum

. pelagica

. perpusilla

. petkovicensis

phormii

. pontiformis

raphani

rubelloides

ruscl

sabuletorum
sepincola

. tamaricis

. triglochinicola

. tritorulosa

. typharum forma acori
uliginosa
umbilicariae

vectis

Si alalalal al al alain al al al al al al al al al al al al nl al ol >

Greece

Green

PRE EPP eee eee eee

. ballotae
. politis
. sibtorpii

a

algida

. brachyasca

. hyperborea

. macrotheca

. oxyriae

. polaris

. ranunculi

. Stellariae

. vahlii

. vanhoeffeniana

Guadeloupe

L. hurae
L. lycii

Himalayas

L. rhododendri

Hungary

. alliariae

. anemones

. antherici

. aquilegiae

. arrhenatheri
azaleae

. berlesei

. betulina

. compressa

. constricta

. coronillae

. corticola

. culmifraga var.
bromicola

all soll Soll Soll Soll Soll Sal Soll all Sal Soll soll vo!

March

Icelan

India

1991

=

culmorum forma
hungarica

. cynoglossi
cyperi

. cypericola

. dematium
dianthi

. doliolum forma
syndoliola

. dumetorum var.
coronillae

. echinops

. equiseticola

. fiumana

. geasteris

. gypsophilae

. hollosii

iridis

irrepta

. juniperina

. lineolaris
maritima
meliloti
muscari
onobrychidicola
. onobrychidis
poae Vat. agrostidis
pontiformis
purpurea
raphidophora

. rubicunda

. Salsolae

. sinapis

. stromatoidea

. superficialis
tamaricis
ternata

. thalictricola

. thalictrina

dryadis
elymi
larseniana
. oligotheca
papaveris

Soll soll Soll soll Soliant Soll Soll Soll soll Soll Soll call Soll wall wall wall coll coll coll coll call call coll coll call “all call call “oll coll all all a SE all oll all all all

. abuensis
agaves

. aquatica
capparidicola
coorgica

. cosmicola

. eriobotryae

. helianthi
hollosiana
hyalina
indica

. isocellula

. lobeliae

L. muehlenbeckiae

Teall call Soll soll Soll Soll Sal Soll all Soll all Soll Sa

Nomenclator of Leptosphaeria

Iran

L. nigrans

L. ocimicola

L. petkovicenses var. elymi
. phoenicis

. porellae

. puniciae

. punjabensis

. rajasthanensis

. swertiae

zingiberis

all Soll soll all Soll all

L. cycadis

L. kotschyana
L. shahvarica
L. tolgorensis

Ireland

Israel

Italy

L. advenula

é

. pimpinellae

aconiti

. aculeorum
acuta
advenula

. aegira

. aetnensis

. aglaja
agminalis
agnita

alcides
alcides forma quercina
ampelina
amphibola
anarithma

. anarithmoides
anthophila

. anthostomoides
apocalypta
apogon
aquilana
arnoldii

. arrhenatheri var. italica
. arundinacea
arvensis
asparagt
bella

. bellynckii

. bornmuelleri
borziana

. brachypodii
brachysperma
bractearum
briosiana

. bryophila
brizae
cadubriae

. camilla

. camphorata

. campisilii

. canadensis

Soloist ol sist siotot aiolcls si olal asia sialal si slal alan al alal alae si alals

‘oath Soll all Soll oa

loll soll Soll Sal Soll

‘onl Soll Soll Sal soll Soll Se

pl Soll Soll all Soll sll Soll wall

Toot all Soll all Soll Goll all ool call wall call wall coll oll wall coll coll cll call cll call oll all call all call all

333

. cannabina
. capparidis

capsularum

. carduina

. carpogena

. carpophila

. carpophila var.

bractearum

. cattanei

cavarae

. cerlettii

cesatiana
chaetostoma
cibostii

. cirricola
. coniothyrium
. coniothyrium forma

berberidis

. conoidea

. corticola

. corynispora

. crastophila

. crastophila forma

tofieldiae

. crepini

. cucurbitae

. culmicola

. culmicola forma major

culmicola var. minor

. culmifraga
. culmorum

cumana

. cyperina

dactylina

dasylirii

derasa var. alpestris
diana

. dichroa

. disseminata
. doliolum

. dryadea

. epilobit

. eustoma

. eustomella

. eustomoides
. faginea

. fallaciosa

fallax

. fiedlaeri

. fuscidula

. galiicola

. galiicola var. brachy-

spora

. geographicola

. gibelliana

. glauco-punctata
. grammodes

. grandispora

. grisea

334

Teall all Soll Soll Soll Soll call all

Sialal al alslalslal al aial ola sl pial Si al>

Teall oll Soll Soll all Soll Soll Soll Soll all Soll Soll all Soll all Soll call call Soll call Soll all call Soll all call allo

hausmanniana
hederae

. helvetica

helvetica forma major
hendersoniae

. hermodactyli
. herpotrichoides

heterospora
heufleri
hiemalis

. hippophaes

hyalospora
hydrophila

. Involucralis

ischaemi

. Juncicola

kochiana
lamprocarpi

. lathonia
. lathonia var. hellebori-

foetidi

. lathyri

. lathyrina

. leersiae

. leersiana

. leptospora

. licatensis

. lichenicola

. littoralis

. littoralis forma

calamagrostidis-
arenariae
livida

. lucilla

. lucina

. luctuosa

. luzulae

. maculans

. Magnusiana

major

.mamillula

. marginalis

. marginata

. massariella

. massariella var. disticha
. massarioides

. medicaginis

. medicaginum

. melicae

. meridionalis

_ michotii

. micropogon

. microscopica

. microscopica Var. alpina
. mirandae

. modesta var. cibostii

. molluginis

. monotis

. montana

. multiseptata forma

alpina

Illinois Natural History Survey Bulletin

| bp
L.
L.
L.
E.
Ei.
E.
EE.
Ih.
E.

atl coll call soll wall wall

atl coll oll soll wall call oa!

a Busy on eu ea

esp sat sab eableeiisat ilies blesilot{eaule: Ae

soll Soll soll soll Soll call Soll Sol Soll Soll Soll Soll So

. nectrioides
. neglecta

nigrans
nobilis

. notarisit
. ogilviensis

ogilviensis var.
senecionis-cordati

. ophiopogonis

oryzae

. pampini

. pandani

. paoluccii

. parietariae

. parvula var. iridis-

germanicae
passeriniana
passerinii
patellaeformis
peltigerarum
penicillus
perforans
perpusilla
petiolicola
phytolaccae
pinnarum
pinnarum var. rachidis
platycarpa
pomona

poteri
puccinioides
punctiformis
pycnostigma vat.
morbosa
pyrenopezizoides
ranunculoides
recessa

rehmii

resedae
rhizomatum
rhododendri
rhodophaea
rivana

rivana var. solorinae
rusci

rusci forma caulina
rusci forma rusci-
hypoglossi
sacculus

. salicaria
. Salicinearum

salviae
salvinii

. sambuci

scirpina

. scotophila
. Seriata

sicula

. Silenes-acaulis

sileris

. socia

Vol. 34 Ar. 3

sodomaea

Spartit

. Spatharum

stereocaulorum

striclata

. subarticulata

subtecta

suffulta

sylvatica

thalictri

torbolensis

. trichostoma

trimera

tritici

typharum

. typhicola

. ulmicola

. vagabunda

. vagabunda forma critri-
limonii

. vagabunda var.
divergens

. vagabunda vat. sarmenti

vaginae

valdobbiae

vincae

. vinealis

. xiphii

. xylogena

. yulan

PRE REE EE RP REE REeee

ll

[aati soll call call Soll soll Soll

Ivory Coast

L. coffeigena var. longiros-
trata

L. petri

L. sorgho-arundinacei

L. peltigerea

. akagiensis
apios

. apios-fortunei
asperellae
. bambusae

. buddlejae

. cCinnamomi
hottai
inecola
Japonica
lelebae

. lilicola

. minoensis

. mume

. nandinae
nashi
oryzicola

. phyllostachydis
. Sasacola
sasae

. thujaecola
. tigrisoides
. zizannivora

all all wall soll Soll all Soll Sal Soll Sal nll Soll all call call all all Soll all Sal all Sal

March 1991

Java
L. sacchari
L. trochus
Kenya
L. bicolor
L. nodorum
L. tritici
Kerguelen Island
L. kerguelensis
Korea
L. iwamotoi
Libya
L. pampaniniana
Luxembourg
L. caespitosa forma
salviae
L. cerastii
L. culmorum var. fla-
vobrunnea
L. dolioloides var. lathyri
L. dolioloides var.
rhinanthi
L. dumetorum var.
coniformis
L. dumetorum var.
dolichospora
L. dumetorum var.
symphyti
L. echiella
L. echii
L. epicalamia var.
pleosporoides
. euphorbiae forma
esulae

ll

. galeobdolonis

. hemerocallidis

iridigena var. typhae

. Junci

. longispora

. luxemburgensis var.
dolichospora

. microthyrioides

. monilispora forma
triglochinis

. oxyspora

. paludosa

. petiolaris

. phyteumatis forma
knautiae

. planiuscula forma
succisae

. plectrospora

. proliferae

rivalis

silvestris

Sparsa Var. meizospora

. (rematostoma

. trifolii

. vitalbae var. sarmenti-

cola

Teall Soll call Sai all Sai Soll Soll Soll all wall soll we

é

oall Soll Soll Sail Soll all Soll

. fuscella var. hippophaes

Nomenclator of Leptosphaeria

L. wegeliniana forma
teucril
Madeira Archipelago
L. maderensis
Malaysia
L. scabiens
Mauritania
L. tompkinsii
Morocco
L. elaoudi
L. gratissima
L. gratissima var. longis-
pora
L. maheui
Netherlands
. ammophilae
. cariciphila
desciscens
. donacina
. galiicola
. genistae
. hemicrypta
L. junci
L. periclymeni
L. phlogis
L. pseudo-diaporthe
L. rhopographoides
L. rousseliana
L. stratiotis
New Caledonia
L. australis
New Zealand
L. martinianum
L. reidiana
L. typharum

feat all all wall call all

L. elaeidis

L. andromedae

L. caricinella

L. consobrina

L. dryadis

L. dumetorum var. galii-
borealis

. hyperborea

. insignis

. junciseda

quadriseptata

. rostrupit

sepalorum

vagans

Toil call soll soll Soll all

Pakistan
. abutilonis
. ahmadii
. depressa
. euphorbiicolla
. punjabensis
L. rumicicola
L. rumicis
Philippines
L. ambiens
L. amphiloga

‘atl Soll all Soll al

ho

Poland

Portug

mS

foal all soll call Soll oa

Toot Stl Soll Soll Soll all coll Soll call coll call all wall oll al coll coll coll coll call wall oll all oll wall ool on

. chusqueae

. dryadea subsp.
lussoniensis

. erythrinae

. marantae

oryzina

panici

. simillima

. tungurahuensis

. alliariae

. caricina

. cerei-peruviant
. coniformis

. crustacea

. lycopodina

. norvegica
opizit

. sowerbyi

. trifolii-alpestris
. triglochinis

. typhiseda

. algarbiensis

. almeidae

. almeidana

aloes

. anacycli

. anarrhini

arbuti

arecae

buxina

. cisticola

cocoes

. congesta

. conimbricensis

. coniothyrium

. convallariae forma
dracaenae

. convallariae forma

rusci

daphnes

. demissa

. diaporthoides

dolioloides

dracaenae

. foeniculi

holmii

. Infernalis

junci-acuti

juncina

. lavandulae

. lusitanica

. maculans

. michotii

molleriana

nervisequa

. papillosa

rusct

. rusci var. fourcroyae

. schoenocauli

. scolecosporarum

335

Ww
w
io)

. thuemeniana

. torrendit

. translucens

. vagabunda forma
daphnes

Sooo

Puerto Rico

L. portoricensis

Romania

. acutispora

. alexandrinis

. derasa var. franconica

. dobrogica

. doliolum forma
carlinae-vulgaris

. doliolum var. dissimilis

. doliolum var. leonuri

. fuckelii

. fusispora var. erysimi

. glyceriae-plicatae

. hazslinszkti

. pomona

. ranunculi-polyanthemi

L. salviae forma minor

L. woroninii

oat Sal Soll all oa!

Tot Soll all Sail Soll all oll oa!

Sao Tomé

L. fungicola
L. musarum

Scandinavia

L. dolioloides
L. inarensis

Senegal

L. senegalensis

Sierra Leone

L. penniseticola

Solomon Islands

L. honiaraensis

South Africa

Spain

. africana

. caffra

. cervispora

. collumiae’

. owaniae

. protearum

. pterocelastri
. verwoerdiana

oath Soll all Sail Soll Gall Soll!

. agnita var. trifolii

. caballeroi

. carlinoides

. catalaunica

. cavanillesii

. ceballosi

. CIrsti-arvensis

. cistina

. colocasiae

. culmicola var. hispalen-
AYAY

. CYNOSUTT

. dolioloides

. draconis

. fuscidula forma
magnolii

all all all oll Soll soll Soll Sal Sal Se

Sati all all Sat

Illinois Natural History Survey Bulletin

ll

bo ooe

E
L
E
I
E.
IL
L
L
5
L

glyceriae
grossulariae
hispanica
Junci-glauci
martagoni

. matritensis

. octophragmia

. riofriot

. riparia

. ruscicola forma

cladodiicola

. sabauda forma arvati-

cae

. semelina

. Staticicola
. Striolata

. variabilis

Sri Lank

L. depressa

L. lankeana

L. nesodes

L. smilacis

L. tornatospora
St. Thomas Island

L. musarum

Sweden

bo

[all Sol all Soll Sal all ll Soll all Sal Gall Sal Sal Sal

oll all Soll all Soll wall

vitensis

. associata
. bellynckii
- Caricis

. cladophila

compressa

. culmifraga
. culmifraga forma

minuscula

. culmorum
. doliolum
. doliolum var. subdisti-

cha

. duplex

. fuckelii

. hendersoniae

. hirta

. Jaceae

. lasiosphaerioides
. macrotheca

. microscopica

. nardi

. nigrans

. picastra

. Sepincola

. solani

. solanicola

. vagabunda subsp.

alvarensis

Switzerland
L. agnita var, major
L. albulae
L. blumeri
L. brauni
L. caricicola

all all all Soll Soll Soll Soll Soll Soll Soll all Soll all ll Soll call soll Soll all voll Soll all Gall Soll all Soll all call Soll all voll all call Soll call call call call wall Soll call coll call all col all oll call all wall all oll coll oll coll all oll all all

Vol. 34 Art. 3

. caricis

. centaureae

. Cladti

. compositarum
. cornuta

. corticola

. crucheti

. culmorum

. didymellae-vincetoxici
. epicalamia

. epilobti

. eustoma

. franconica

. fuckelii

. galii

. galiorum var. gentianae
. hippophaes

. Johansonii

. Juncicola

. lacustris

. ladina

. libanotis

. linearis

. lycopodina

. maculans

. morthieriana
. nanae

. napelli

. nardi

. nigrans

. nodorum

. ocellata

. ogilviensis

. ophioboloides
. palustris

. parvula

. petkovicensis
. phacae

. phaeospora

. phyteumatis

. pint

. planiuscula

. platychorae

. pleurospora

. polygonati

. primulaecola
. recutita

. rimalis

. rimicola

. robusta

. rousseliana

. rumicis

. septemcellulata
. Sowerbyi

. Sparsa

. submodesta

. sylvatica

. taminensis

. tenuis

. thurgoviensis

eccrine a

March 1991

Taiwan

Tool soll soll soll soll soll call wall wo

[Soll all all soll soll allo

. tofieldiae

. triglochinicola
trollii
typhicola
valesiaca
viciae

. vitalbae

. volkartiana

. wegeliniana

coicis

. kuangfuensis
. musae

. musigena

. pandanicola

. taichungensis
. taiwanensis

Tanzania

fi
Tunisia
E
Ve

it
Turkey
Ie
JE
Ib
L

U.S.A.

Sl aol lal i al al all al ol ol all al all ol all all al slal al al ala! ai bl alas

. elaeidicola

. macrochloae

. rothomagensis var.
artemisiae

. Stipae-minor

. davisiana

. dodonaeae
. melicae

. sylvatica

acuta

. agnita subsp. labens
. agnita var. erigerontis
anomala
anthelmintica
arctalaskana
arthrophyma
arunci

. asclepiadis
asparagi

. associata

. astericola

. asteris

. aulica

. avenaria

. avicenniae

. beaumontit

. berlesei
bicuspidata
bocconiae
borealis
borealis var. populi
boucera
brightonensis
byssincola
cacuminispora
californica
caricis

. cassiaecola

. castilleiae

. castillejae

Nomenclator of Leptosphaeria

[oll Soll Soll all oll Soll Soll soll Soll call Soll Soll call Goll coll coll coll oll coll coll wall oll coll coll all voll oll wall oll oll coll oll wall oll all call

Teall all Soll all Soll SalI Soll all vol all cal all

PRR EEE oe

. castrensis

. caulincola
. ceanothi

. ceratispora
. cercocarpi

chrysanthemi
clavicarpa

. Clavigera

clavispora

. Clivensis var. constricta
. coleosanthi
. collinsoniae

comatella
complanata
compressa

. concentrica

concinna
consessa
consimilis
corallorhizae
culmorum
darkeri
defodiens
discors

. dissiliens
. distributa
. doliolum

. drechsleri
. dryophila
. ellisiana

. elongata

elymi

. erigerontis

. eriophora

. eumorpha

. eustoma

. eustoma forma legumi-

nosa
eutypoides
exocarpogena
faulii

. filamentosa

. filiformis

. fimbriata

. foeniculacea subsp.

lupina

. folliculata var. oxyspora
. fraserae

. fraxini

. fuckelii

. fulgida

galiorum var. gnaphali-
ana

. galligena
. georgius-fischeri

halima
hamamelidis

. harknessianna
. helianthi

heliopsidis
heloniaefolia

Foot oll all oll soll Soll ll oll all oll wall ool cll oll oll oll coll cll all wal wall call call call coll call cll all call coll cll cll call col coll call all call coll call all call call oll ll all Soll call ll call all Soll call all all Soll Soll Soll al aldo

Le
Le
~

. hendersoniae
. hesperia

. hiemalis

. houseana

. hysterioides

. incarcerata

. Inquinans

. Inspersa

. interspersa

Janus
kalmiae
korrae

. larseniana

lasioderma
lassenensis
latebrosa

. leiostega

lethalis
leucoplaca

. longipedicellata

lophanthi

. luctuosa

lupini
lupinicola
lyciophila
lycopodiicola
lycopodina

. lyndonvillae

lythri
maclurae
major

. marcyensis

marina
maydis

. mellispora

mertensiae

. mesoedema

modestula

. monticola
. muehlenbergiae
. muricata

muirensis
myricae

. neomaritima

nigrans

. nigricans var. grindeliae
. occidentalis
. octoseptata

odora
ogilviensis

. olericola

olivacea
olivaespora

. onagrae
. opuntiae

orae-maris

. orthogramma
. pacifica

. papyricola

. perplexa

. phaseolorum

ive)
ww
oo

USS.

pala al lalallala al alel aia alal al alalal al alalelalalelalolalelelelolalelelale lela lolol loo toto lose io nooo no

L. phormicola

. physalidis

. physostegiae
. platanicola

. platypus

. plurisepta

. pseudohleria
. puteana

. quamoclidii
.ramulicola
rhoina

. rostrata

. rubrotincta
sabalicola
sabaligera
sambucina
scapophila
shastensis

. Sleversiae
simmonsit
solheimii

. sorgho-arundinacei
. sorghophila
Spartinae
sporoboli

. Squamata

. stereicola
sticta
stictoides
stictostoma

. Straminis

. subcaespitosa
subcompressa
. subconica

. subcutanea

. sublanosa

. Substerilis

. taxicola
tenera

. tephrosiae

. tetonensis
tini

. torulispora

. trimerioides
. tritici var. papyricola
. tumefaciens
. typharum

. utahensis
variegata
variiseptata
veratri

. virginica
viridella

. wehmeyeri

. xerophylli
zeae

. zizaniaecola

. abutilonis
-aceris
. aeluropodis

~~

Illinois Natural History Survey Bulletin

Sail Soll Sal

f

Si alalalal sl alslalslal al aioli sialsl sl sl sialolal stalls

Sal ai al al al al al allel al ai al al al al all al al al al al al oll oll

. agnita var. bupleuri

alhagii
alopecuri
ammophile
ammothamni
amorphae
atraphaxidis

. atriplicis

balcarica
bataticola
batumensis

. betulina

biebersteinti
buddlejae

calligoni
camelliae-japonicae
camphorosmae
caricis-vulpinae
casta
cephalariai-uralensis
cerei-peruviant
chamaeropis

. chenopodii-albi

. chochrjakovii

. clerodendri

. coniothyrium var.

foliicola

. consocians

. culmorum

.culmorum forma epigeil
_ culmorum forma

phragmitis

. culmorum var. hungar-

ica

. daphniphylli

davidii

dianthi

doliolum var. cacaliae
drabae

dubia

equiseti

. feijoae

. ferruginea
. ferulicola
. foliicola

. frigida

ginkgo

. glandulosae

gossypii
haloxyli
hierochloae
holmii
hordei
hydrangeae
hypericola
lespedezae
lithophilae

. lonicerae
. magnoliae
. media

Vol. 34 Art. 3

medicaginicola
metasequoiae
moutan

myrti

. myrticola
ornithogali
periclymeni var. tatarica
. pleurospora
polini

pruni

. pruni var. plurivora
. Sanguisorbae
scutati

secalina

sibirica
smarodsii
sophorae

sorbi

stipae

tanaceti

taurica
tumefaciens
typharum

usneae

vrieseae

weberi
woodrow-wilsonii

a We a Ll ae a aM cl ad ele hela te les ila

Uganda

L. hyparrheniae
L. penniseti
L. trichopterygis

Venezuela

L. cryptica

L. espeletiae
L. jahnii

L. orthrosanthi

Yugoslavia

Zaire

L. altaica

L. andrijevicensis

L. bresadolaeana

L. corrugans

L. fuckelii

L. galiorum subsp.
antirrhini

. hesperidicola

. insulana

malyi

. nicolai

pachyasca

. pachytheca

. petkovicensis

. plemeliana

. rehmiana

. sclavonica

. Serbica

. subalpina

. subsimilis

‘call sll soll Soll Soll Soll all Soll Soll Soll Soll all oo!

. canephorae
. cynodontis-dactyli

‘aol

Appendix |. Taxonomic Division of Leptosphaeria

P.A. SACCARDO (1883, 1891)

Parasites on dicotyledons.
Stem/branch inhabiting.
Leaf inhabiting.
Fruit and flower inhabiting.
Parasites on monocotyledons.
Parasites on acotyledons.
Species of unknown or doubtful spore color.

F. v. HOHNEL (1918c)

Scleropleella F. vy. Héhnel. Species with typical
pseudosphaerialean centra.

Leptosphaeria sensu F. v. Héhnel. Species with
typical dothidealean centra.

Nodulosphaeria G.L. Rabenhorst. Species with
typical sphaerialean centra.

E. MULLER (1950)

Scleropleella. This section, with few exceptions,
corresponds to Héhnel’s (1918a) generic
concept. Generally it includes smaller forms
with relatively few egg-shaped or broadly
club-shaped asci that are embedded ina
more or less well-developed cellular tissue.
The ostiole usually is poorly developed.

Eu-Leptosphaeria. This section contains the largest
number of species of the four sections. The
inner structure corresponds to a higher (more
evolved) stage than Scleropleella; the asci are
more numerous and more slender; the
paraphysoids are clearly thread-like, although
in some forms a cellular arrangement is still
noticeable; the ostiole is conspicuous, and
early in development is filled with hyaline
cells that only later give way to the pore.
Spores are thinner than in Massariosphaeria,
and are often fusiform, and they have a
mucilaginous coat.

Massariosphaeria. This section is composed of
forms whose spores (approaching the
Wettsteinina type) are relatively broad,
possess a distinct mucilaginous coat when
young, and reach maturity relatively late (at
least in part), sometimes only outside the
asci. Structure of the fruiting bodies corre-

sponds to that of section Eu-Leptosphaeria.
These forms may be regarded as transitional
to Massaria.

Nodulosphaeria. This section is composed of the
most highly evolved forms, designated by F.
v. Héhnel as “sphaerial.”” They are distin-
guished from all others by the ostiole, which
is coated with thread-like, periphyses-like
hyphae that usually diverge toward the middle
of the mouth-channel and toward the
“scheitel.” In the region of the “scheitel,”
these periphyses are often replaced by brown
bristles. Peridia of fruiting bodies usually
consist of several layers of elongate, some-
times almost rectangular cells. The numerous,
usually cylindrical-clavate asci are surrounded
by thread-like paraphysoids.

A. MUNK (1957)

Eu-Leptosphaeria. Pseudothecia sclerotioid, thickest
at the sides. Asci slender, numerous.
Ostiole, generally without a periphysoid
structure.

Para-Leptosphaeria. Pseudothecia middle-sized,
with a uniform, thin peridium; generally no
periphysoid structure in the papilla. Interas-
cicular tissue paraphysoid. This section
contains a large and heterogenous group of
species.

Scleropleella. Pseudothecia small; interascicular
tissue obsolete.

Nodulosphaeria. Pseudothecia with a complicated
structure of the papilla. It is covered with
brown, spiny hairs that extend into the
ostiole. Periphysoid tissue is distinct.

L. HOLM (1957)

Leptosphaeria sensu L. Holm. Species similar to the
type of the genus Leptosphaeria doliolum;
they occur principally on dicotyledons.

Nodulosphaeria G.L. Rabenhorst. Species similar to
the Nodulosphaeria of Miiller; they occur on
dicotyledons, especially on Compositae.

Phaeosphaeria |. Miyake. Species that correspond
essentially to the Scleropleella section of
Miiller (1950) and Munk (1957) and that
occur on monocotyledons.

340 Illinois Natural History Survey Bulletin

Entodesmium H. Riess. Species with elongated
ascospores; they intergrade with Ophiobolus
and occur on legumes.

R.A. SHOEMAKER (1984)

Leptosphaeria. Circumscribed Leptosphaeria in a
broad concept of Holm and Miiller;
however, many segregate genera were
accepted. The genus always lacks erect
setae on ascomata. The walls of the
ascocarps often have scleroplectenchyma at
least near the beak base. The physes are
broad, septate with or without guttules and
cytoplasmic accumulations and frequently
with an external gelatinous coating. Asci are
biseriate or, rarely, uniseriate. Ascospores
are fusiform, cylindric or clavate, 3-septate
or more, and the first formed septum is
constricted and near the middle. These

Vol. 34 Art. 3

ascospores frequently have globose, terminal
appendages that are rarely entirely sheathed.
No woody parts of dicotyledonous plants are
the usual susbstrates.

M.E. BARR (1987a, 1987b)

Leptosphaeriaceae. Established the family

Leptosphaeriaceae based on Leptosphaeria,
but also included Curreya, Didymolepta,
Heptameria, and Ophiobolus. The family
was segregated from the Pleosporaceae
because of the coelomycetous rather than
the hyphomycetous anamorphs and because
of the narrow, thin-walled asci. It differs
from the Phaeosphaeriaceae in having
conoid and applanate or obpyriform or
sphaeroid ascomata, whose walls are
scleroplectenchymatic.

Appendix 2. Genera Historically Allied to Leptosphaeria

Bricookea M.E. Barr, Mycotaxon. An International
Journal Designed to Expedite Publication of
Research on Taxonomy & Nomenclature of
Fungi & Lichens, Ithaca, New York 15:346.
1982. Type species: Bricookea sepalorium
(J.S. Vleugel) M.E. Barr. Family:
Phaeosphaeriaceae. Ascomata: Clustered,
immersed, subepidermal, separate or as
locules in crustose stromatic tissues, globose,
radiate arrangement of cells of peridium.
Asci: Bitunicate, oblong, short-stalked,
numerous, basal. Ascospores: Fusiform to
narrowly obovoid, 3-septate, hyaline.
Anamorph: Unknown. Remarks: Barr (1982)
erected this genus for a species of Lepto-
sphaeria described from inflorescences of
Juncus and considers it to be related to
Phaeosphaeria.

Chaetomastia (P.A. Saccardo) A.N. Berlese, Icones
Fungorum Omnium Hucusque Cognitorum
ad usum Sylloges Saccardianae Adcommoda-
tae 1:38. 1891. Lectotype species: Chaeto-
mastia hirtula (P.A. Karsten) A.N. Berlese.
Family: Dacampiaceae. Ascomata: Im-
mersed-subepidermal, becoming erumpent,
scattered or clustered, obpyriform or ovoid to
globose; apex wide, blunt, ostiole rounded or
slit-like; wall pseudoparenchymatous, cells
externally darkened and thickened in upper
regions. Asci: Bitunicate, clavate or
cylindric, 4-, 6-, or 8-spored. Ascospores:
Biseriate or uniseriate; obovoid elongate, 3-
to 11-septate; cell above first septum
enlarged; wall wide, dark, smooth or
verruculose; yellowish brown becoming dark
brown or reddish brown. Anamorph:
Coelomycetous where known; conidia
hyaline or brown, |- to 2-celled
(Aposphaeria-like or Coniothyrium-like).
Remarks: This genus is distinguished by the
obpyriform or obovoid ascomata with a wide
apical papilla, peridium of small dark cells
that is widest and 3-layered in the upper
region, and by dark asymmetric
phragmospores (Barr 1989).

Chaetoplea (P.A. Saccardo) F.E. Clements, in F.E.
Clements and C.L. Shear, The Genera of
Fungi, pp. 74, 275. 1931. =Pyrenophora
subgen. Chaetoplea P.A. Saccardo, Sylloge

Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:279. 1883. Type
species: Chaetoplea calvescens (E.M. Fries
ex J. Desmaziéres) F.E. Clements & C.L.
Shear. =Sphaeria calvescens E.M. Fries,
Scleromyceti Sueciae. Collegit, Digessit et
Evulgaviti, No. 401. Unpublished? =Py-
renophora calvescens (E.M. Fries ex J. Des-
mazieres) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:279. 1883. Family:
Phaeosphaeriaceae. Ascomata: Seated on a
subiculum, globose or depressed globose,
dark brown to black, subepidermal becoming
superficial, wall composed of polygonal
cells, surrounded by stiff dark hyphae. Asci:
Bitunicate, numerous, cylindrical, short-
stalked with thickened apical walls, 8-spored.
Ascospores: Fusoid-ellipsoid, straight or
inequilateral with three transverse septa and
with longitudinal septa, yellowish brown or
dark brown, smooth or finely verruculose.
Remarks: This genus has only recently been
accepted by Barr (1981, 1987b), who placed
it in the Phaeosphaeriaceae. The type
species, Chaetoplea calvescens, has been
placed in Pleospora by Webster and Lucas
(1959), Wehmeyer (1961), and Shoemaker
(1968). Crivelli (1983) transferred Chaeto-
plea calvescens to Leptosphaeria, and
Eriksson and Hawksworth (1986)
synonymized Chaetoplea with Lepto-
sphaeria.

Curreya P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:651. 1883. Type species:
Curreya conorum (L. Fuckel) P.A. Saccardo.
Family: Leptosphaeriaceae. Ascomata:
Scattered, immersed-subepidermal, depressed
globose; wall scleroplectenchymatic,
melanized black, surface of textura angularis;
beak absent. Asci: Many, bitunicate,
cylindrical-clavate, thick-walled in apical
region, short-stalked, containing eight
ascospores. Ascospores: Obovate, con-
stricted at septa, slightly asymmetrical, with
one or more enlarged cells, 7-septate or
more, with longitudinal septa, most cells
smooth-walled, surrounded by a gel layer.

Illinois Natural History Survey Bulletin

Remarks: The genus is included in the
Leptosphaeriaceae by Barr (1987b) and
differs from Leptosphaeria in having
longitudinally septate ascospores.

Didymolepta A. Munk, Dansk Botanisk Arkiv,

Kjobenhavn 15(2):110. 1953. Type species:
Didymolepta winteriana (P.A. Saccardo) A.
Munk. Family: Leptosphaeriaceae. Asco-
mata: Scattered, conic, glabrous, black; wall
of scleroplectenchymatic cells melanized
black; beak absent. Asci: Bitunicate,
subcylindric, sessile, thick-walled, with eight
ascospores. Ascospores: l-septate, hyaline.
Anamorph: Unknown. Remarks: This genus
has been placed in the Leptosphaeriaceae by
Barr (1987b), and it differs from other genera
in the family by having |-septate ascospores.

Entodesmium H. Riess, Hedwigia, Dresden 1:28.

1854. Type species: Entodesmium rude H.
Riess. Family: Phaeosphaeriaceae? Asco-
mata: Scattered or clustered, more or less
lageniform, slightly hairy; beaks very long.
Asci: Many, bitunicate, cylindric to narrowly
clavate, short-stalked. Ascospores: Cylindri-
cal, 4-septate or more, with a clearly
delimited apical portion and bipolar append-
ages. Anamorph: Unknown. Remarks: This
genus was recognized by Holm (1957) for a
small group of Leptosphaeria species found
on Leguminosae.

Graphyllium F.E. Clements, Studies in the

Vegetation of the State. Nebraska University.
Botanical Survey. Report on Recent
Collections, Lincoln 5:6. 1901. Type
species: Graphyllium chloes F.E. Clements.
Family: Phaeosphaeriaceae. Ascomata:
Flattened-globose, finally collapsing-
pezizoid, with more or less dark brown,
radiating hyphal tomentum about the base.
Asci: Bitunicate, stout-clavate, thick-walled.
Ascospores: Strongly flattened in one plane,
fusoid-ellipsoid to clavate-ellipsoid, with a
single vertical septum running through the
central cells but not through the end cells in
face view, yellow-brown to dark red-brown.
Anamorph: Unknown. Remarks: Graphyl-
lium is the earliest name for species formally
placed in Platyspora L.E. Wehmeyer and
Comoclathris F.E. Clements (Barr 1987b).

Heptameria H. Rehm & F. v. Thiimen, Instituto.

Revista Scientifica e Litteraria, Coimbra,
Series 2, 27:252. 1879. Type species:
Heptameria elegans H. Rehm & F. v.
Thiimen. Family: Leptosphaeriaceae.
Ascomata: Clustered, immersed-subepider-
mal, becoming erumpent, subglobose to
conical, rough-surfaced, black, short
papillate. Asci: Bitunicate, clavate, short-
stalked, with eight ascospores. Ascospores:
Fusiform, 7-septate or more, with dark

Vol. 34 Art. 3

central portion having longitudinal septa,
brown. Anamorph: Pycnidial with hyaline,
oblong, 1-celled conidia. Remarks: This
genus was included in the Leptosphaeriaceae
by Barr (1987b), and it differs from Lepto-
sphaeria by having ascospores with longitu-
dinal septa in the enlarged central cells.
Jaczewski (1894) considered that Heptameria
could be interpreted as a section of Lepto-
sphaeria, but the study of Lucas and Sutton
(1971) indicates that the genus is distinct.

Herpotrichia L. Fuckel, Fungi Rhenani Exsiccati A

Leopoldo Fuckel Collecti, Fascicle 22, No.
2771. Anno 1868. Type species: Herpotric-
hia rubi L. Fuckel. Family: Lophiostomata-
ceae. Ascomata: Globose to conic, im-
mersed, becoming erumpent, or superficial
on a subiculum, tomentose. Asci: Bitunicate,
cylindrical to clavate. Ascospores: Fusiform
to ellipsoidal, 1- to 3-septate, hyaline to dull
or dark brown, usually with gel coating
elongated beyond spore apices. Anamorph:
Coelomycetous, Pyrenochaeta, or Phoma-
like. Remarks: Differs from Leptosphaeria
in the tomentose ascocarps on a subiculum
and ascospores that can be |-septate.

Kalmusia G. Niess| v. Mayendorf, Verhandlungen

des Naturforschenden Vereins in Briinn
10:204. 1872. Type species: Ka/musia ebuli
G. Niessl v. Mayendorf. Family:
Phaeosphaeriaceae. Ascomata: Subglobose,
immersed in an effuse stroma. Asci:
Bitunicate, clavate, long stipitate. Asco-
spores: Oblong, curved, 3-septate, brown.
Anamorph: Unknown. Remarks: This genus
differs from Leptosphaeria in having a
stroma and long, stipitate asci.

Keissleriella F. v. Hohnel, Sitzungsberichte der

Akademie der Wissenschaften in Wien,
Mathematisch-naturwissenschaftliche Klasse,
Abt. I, 128:582. 1919. Type species:
Keissleriella aesculi (F. vy. H6hnel) F. v.
Hohnel. Family: Melannomataceae.
Anamorphs: Ascochyta M.A. Libert and
Dendrophoma P.A. Saccardo. Remarks:
Holm (1957) accepted the genus Tricho-
metasphaeria A, Munk but stated that there
may be earlier generic names for those
Ascomycetes with setose pseudothecia
placed in the Massarinaceae by Munk (1956),
for example, Keiss/eriella F. v. HGhnel. This
later genus differed in having l-septate
ascospores vs. several septate in Trichom-
etasphaeria. Bose (1961) united the two
genera under the earlier name Keiss/eriella
after observing variation in ascospore
septation in Keissleriella aesculi.

Lidophia J. Walker & B.C. Sutton, Transactions of

the British Mycological Society, London
62:232. 1974. Type species: Lidophia

March 199]

graminis (P.A. Saccardo) J. Walker & B.C.
Sutton. =Dilophia P.A. Saccardo, 1883, non
Dilophia T. Thomas, 1953 (Cruciferae).
Family: Leptosphaeriaceae. Ascomata:
Embedded in a stroma, spherical. Asci:
Bitunicate, cylindrical, apically thickened.
Ascospores: Narrowly fusiform, widest at the
middle and tapering gradually into an
elongated fine thread at each end, ~15-
septate, yellow, breaking into equal spore
parts at maturity. Anamorph: Unknown, but
Dilophospora is found in same stroma.
Remarks: The ascospores of Lidophia were
considered indistinct from those of Lepto-
sphaeria (Miiller 1950; v. Arx and Miiller
1975); however, Walker (1980) considers
Lidophia distinct from Leptosphaeria and
close to Ophiobolus in that the ascospores
readily break into half-spores at the central
septum. Lidophia differs from Ophiobolus in
lacking an enlarged central cell in the
ascospore.

Lophiostoma V. Cesati & G. de Notaris, nom. cons.,

Commentario della Societa Crittogamologica
Italiana, Milan 1:219. 1863. Type species:
Lophiostoma macrostoma (H.J. Tode:E.M.
Fries) V. Cesati & G. de Notaris. Based on
Sphaeria macrostoma H.J. Tode:E.M. Fries.
Family: Lophiostomataceae. Ascomata:
Immersed, subepidermal, globose to
subglobose, glabrous, cells of outer wall
melanized black, beak laterally compressed,
clypeate. Asci: Bitunicate, many, cylindro-
clavate, wall thin but thick at apex, short-
stalked, eight-spored. Ascospores: Obliquely
uniseriate, ellipsoidal to narrowly clavate,
brown, 3- to 7-phragmoseptate, | to 2
longitudinal septa present. Anamorph:
Unnamed pycnidial anamorph (Chesters and
Bell 1970). Remarks: Lophiostoma differs
from Leptosphaeria in having ascomata, with
laterally compressed papillae (Chesters and
Bell 1970).

Massaria G. de Notaris, Nuovo Giornale Botanico

Italiano e Bolletino della Societa Botanica
Italiana, Firenze 1:333. 1844. Type species:
Massaria inquinans (H.J. Tode:E.M. Fries)
G. de Notaris. Basionym: Sphaeria inqui-
nans H.J. Tode:E.M. Fries. Family:
Massariaceae. Ascomata: Immersed in
pseudostromatic tissue or a clypeus becom-
ing erumpent or superficial with bases
remaining immersed. Asci: Bitunicate,
subcylindrical with a short, stout stipe, ocular
region low and broad surrounded by a
refractive ring. Ascospores: Large, distosep-
tate, and 3-septate. Anamorph: None
reported (Miiller 1979). Remarks: The
ascospores of Massaria are generally larger
than in Leptosphaeria. Each ascus has an

Nomenclator of Leptosphaeria 343

apical cytoplasmic protrusion surrounded by
a refractive ring and usually four ascospores
at maturity. The ascospores are larger than
those in Leptosphaeria and have a thick
epispore and a mucilaginous coat (Barr
1979).

Massarina P.A. Saccardo, Sylloge Fungorum

Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:153. 1883. Lectotype
species: Massarina eburnea (L.R. Tulasne &
C. Tulasne) P.A. Saccardo. Family:
Lophiostomataceae. Ascomata: Clustered,
immersed, subepidermal, depressed,
glabrous, cells of outer ascocarp wall
melanized brown, wall tissue of textura
prismatica in face view, beak absent, ostiole
circular. Asci: Many, bitunicate, cylindric to
cylindric-clavate, short-stalked, wall thick at
apex. Ascospores: Biseriate, oblong-fusoid,
3-septate, slightly constricted at the septa,
symmetrical, hyaline to subhyaline, fre-
quently surrounded by a mucous sheath,
which may be evanescent. Anamorphs:
Anguillospora C.T. Ingold, Ceratophoma F.
v. Hohnel, Coniothyrium A.C. Corda,
Diplodia E.M. Fries, Microsphaeropsis F. v.
Hohnel, and Stagonospora P.A. Saccardo.
Remarks: Massarina is allied with Lepto-
sphaeria species having hyaline ascospores
(Metasphaeria sensu P.A. Saccardo, 1883)
but differs in formation of a clypeus and in
the ascospores, which form their secondary
septa late in development (Munk 1956; Bose
1961; Miiller 1979).

Massariosphaeria (E. Miiller) P.G. Crivelli,

Dissertation Eigendssischen Technischen
Hochschule, Ziirich, No. 7318:141. 1983.
Type species: Massariosphaeria phaeospora
(E. Miller) P.G. Crivelli. Family: Dacampi-
aceae. Ascomata: Scattered, globose to
conic, immersed, subcuticular to subepider-
mal, becoming erumpent, beak papillate,
conical to longitudinally compressed with a
circular or slit-like ostiole. Asci: Bitunicate,
numerous, clavate-cylindrical, short-stalked,
thick-walled. Ascospores: Phragmoseptate,
sometimes with longitudinal septa, hyaline to
brown, fusiform to ellipsoid or clavate.
Anamorph: Aposphaeria-like, Phoma-like.
Remarks: Miiller (1950) segregated species
of Leptosphaeria with large, thick-walled
ascospores with thick gelatinous sheaths into
the subgenus Massariosphaeria. Crivelli
(1983) elevated the subgenus to genus and
expanded it to include dictyosporous species.
It appears to be a natural grouping with easily
distinguished features.

Melanomma T. Nitschke ex L. Fuckel, Symbolae

Mycologicae, p. 159. 1870. Lectotype
species: Melanomma pulvis-pyris (C.H.

344

Illinois Natural History Survey Bulletin

Persoon:E.M. Fries) L. Fuckel. Family:
Melanommataceae. Anamorphs: Apo-
sphaeria P.A. Saccardo and Pseudospiropes
M.B. Ellis. Remarks: Holm (1957)
maintained Melanomma for certain ligni-
colous species that appear to form a unit
based on the scleroplectenchymatic walls of
the ascocarps. The ascospores are 3-septate,
ellipsoidal or frequently cuneiform to nearly
clavate and uniformly brown. Melanomma
was further subdivided into four groups
based on variability and differentiation of the
ascocarp wall. A number of species on dead
herbaceous stems usually placed in Lepto-
sphaeria were transferred to Melanomma.
Holm (1957) considered Melanomma to be
closely related to Leptosphaeria, but Samuels
and Miiller (1978) believed that Melanomma
should be merged with Trematosphaeria
(Chesters 1938).

Montagnula A.N. Berlese, Icones Fungorum

Omnium Hucusque Cognitorum ad usum
Sylloges Saccardianae Adcommodatae 2:68.
1896. Type species: Montagnula infernalis
(G. Niessl v. Mayendorf) P.G. Crivelli.
Family: Phaeosphaeriaceae. Ascomata: With
or without a clypeus or surrounded by a
secondary stromatic development, globose.
Asci: Bitunicate, clavate, with filiform basal
stalk having a claw-like base, wall thickened.
Ascospores: Fusoid, mostly 3-, rarely 4-, 5-,
or 7-septate, dark red-brown to opaque, wall
often finely tuberculate, vertical walls in two
central cells, symmetric, constricted at the
central septum. Anamorph: Unknown.
Remarks: Considered a subgenus of
Pleospora by Wehmeyer (1961), Montagnula
was raised to generic rank by Crivelli (1983).
Characteristics distinguishing the group are
the tapered or elongate stipitate base of the
ascus, the usually very dark pigmentation of
the spore wall (which is often tuberculate-
roughened), the tardy insertion of vulgaris-
like septa in the central cells, the tendency to
form a thick ascostromatic wall, and a
surrounding clypeus or stroma.

Nodulosphaeria G.L. Rabenhorst, Klotzschii

Herbarium Vivum Mycologicum Sistens
Fungorum Per Totam Germaniam Cresen-
tium Collectionem Perfectam, Dresden,
Edition II, Century 8, No, 725. Anno. 1858.
Type species: Nodulosphaeria derasa (M.J.
Berkeley & C.E. Broome) L. Holm. Family:
Phaeosphaeriaceae. Ascomata: Clustered,
immersed-subepidermal, globose, setose,
wall pseudoparenchymatic, melanized
brown, of textura angularis, beak short, with
paraphyses. Asci: Many, bitunicate,
cylindro-clavate, short-stalked, containing
eight ascospores. Ascospores: Fusiform,

Vol. 34 Art. 3

constricted at mid-septum, 8- to 10-septate,
with one enlarged cell, with bipolar append-
ages, pale brownish yellow. Anamorph:
Unknown. Remarks: Once considered a
subgenus of Leptosphaeria, Nodulosphaeria
was reinstated by Holm (1957). This genus
can be distinguished from Leptosphaeria by
the setose ascomata composed of pseudopar-
enchyma.

Ophiobolus H. Riess, Hedwigia, Dresden 1:27—28.

1854. Type species: Ophiobolus disseminans
H. Riess. Family: Leptosphaeriaceae.
Ascomata: Scattered to clustered, immersed,
becoming erumpent, pyriform, black,
glabrous, outer wall of textura globulosa
melanized brown, inner wall of textura
prismatica, wall pseudoparenchymatic. Asci:
Bitunicate, numerous, long-cylindrical to
clavate, thick-walled at tips, short-stalked, 4-
to 8-spored. Ascospores: In a single fascicle,
yellow or brown, sclolecosporous, multisep-
tate, often with one or more enlarged cells,
sometimes with one or more gelatinous,
terminal, globose appendages. Anamorph:
Unknown. Remarks: Ophiobolus integrates
with Leptosphaeria subgenus or section
Nodulosphaeria (G.L. Rabenhorst sensu F. v.
Hohnel) E. Miiller, and authors do not agree
on the precise limit between Ophiobolus and
subgenus Nodulosphaeria. In general, the
ascospores of subgenus Nodulosphaeria are
only slightly curved or straight and shorter
than those in Ophiobolus, and dark periphy-
ses line the ostiole in subgenus Nodu-
losphaeria (Shoemaker 1976, Walker 1980).

Ophiosphaerella C.L. Spegazzini, Anales del

Museo Nacional de Historia Natural de
Buenos Aires 19(Series 3, 12):401. 1909.
Type species: Ophiosphaerella graminicola
C.L. Spegazzini. Family: Phaeosphaeriaceae.
Ascomata: Scattered to clustered, immersed,
subepidermal, globose, glabrous, wall of
textura angularis in face view, beak short,
papillate, bluntly conical with periphyses.
Asci: Bitunicate, long, cylindric, in a dense
basal layer. Ascospores: Filiform (scoleco-
sporous), phragmoseptate, brown, lacking
gelatinous sheaths. Anamorph: Unknown.
Remarks: A genus close to Ophiobolus H.
Riess sensu stricto and Phaeosphaeria 1.
Miyake, it is characterized by scoleco-
sporous, brown, septate ascospores and
occurs on Gramineae and Cyperaceae.

Paraphaeosphaeria O. Eriksson, Arkiv for Botanik,

Uppsala, Stockholm, Series 2, 6:405. 1967.
Type species: Paraphaeosphaeria michotii
(G.D. Westendorp) O. Eriksson. Family:
Phaeosphaeriaceae. Ascomata: Scattered,
intraepidermal, depressed globose to globose,
without prominent beak. Asci: Bitunicate,

—_

March 1991

numerous, cylindrical, short-stalked.
Ascospores: Clavate to cylindrical, 2- to 9-
septate, first-formed septum subtending an
enlarged cell, echinulate to rarely smooth,
with a thick, gelatinous sheath. Anamorph:
Coniothyrium A.C.J. Corda. Remarks:
Paraphaeosphaeria is a segregate of
Leptosphaeria resembling Phaeosphaeria,
but differs in having ascospores with more
septa and Coniothyrium anamorphs. In
Phaeosphaeria, the anamorphs belong to
Hendersonia P.A. Saccardo and Phaeosepto-
ria C.L. Spegazzini (Eriksson 1967).

Phaeosphaeria |. Miyake, Botanical Magazine,

Tokyo 23:93. 1909; Journal of the College
of Agriculture, Imperial University of Tokyo
2:246. 1910. Type species: Phaeosphaeria
oryzae |. Miyake. Family: Phaeosphaeri-
aceae. Ascomata: Superficial, globose,
glabrous, wall pseudoparenchymatic, thin.
Asci: Many, bitunicate, cylindro-clavate,
containing eight ascospores. Ascospores:
Fusiform to cylindrical, 3-septate, yellowish
brown, one cell clearly inflated. Anamorph:
Coelomycetous, Phyllosticta/Hendersonia.
Remarks: Ascospores of species in this genus
are very similar to those of Leptosphaeria
species. Phaeosphaeria species are distin-
guished by small thin-walled, pseudopar-
enchymatic ascomata, and by their occur-
rence mostly on monocots (Shoemaker and
Babcock 1989).

Platystoma V. Trevisan, Bulletin. Société R. de

Botanique de Belgique, Bruxelles 16:16.
1877. Type species: Platystoma compressum
(C.H. Persoon:E.M. Fries) V. Trevisan.
=Sphaeria compressa C.H. Persoon:E.M.
Fries. Family: Platystomaceae. Ascomata:
Clustered, immersed-subepidermal, globose,
glabrous, outer cells melanized brown, short
papillate, papilla conical with periphyses,
ostiole slit-like. Asci: Many, bitunicate,
cylindric-clavate, short-stalked. Ascospores:

Fusiform to subcylindric, constricted at septa,

5- to 6-septate, with one enlarged cell,
brown. Anamorph: Unknown. Remarks:
This genus differs from Leptosphaeria in
having trabeculate rather than cellular
pseudoparaphyses and a slit-like ostiole.

Pleospora G.L. Rabenhorst ex V. Cesati & G. de

Notaris, nom. cons., Commentario della
Societa Crittogamologica Italiana, Milan
1:217. 1863. Type species: Pleospora
herbarum (C.H. Persoon:E.M. Fries) G.L.
Rabenhorst type cons. Family: Pleospo-
raceae. Ascomata: Perithecioid, immersed-
erumpent, peridium relatively wide,
composed of pseudoparenchyma. Asci:
Bitunicate, clavate or oblong, short-stalked,
endotunica wide. Ascospores: Elliptical,

Nomenclator of Leptosphaeria 345

yellow-brown, with transverse and longitudi-
nal septa, with gelatinous sheath.
Anamorphs: Alternaria C.G. Nees v.
Esenbeck, Dendryphion K.F.W. Wallroth,
and Sremphylium K.F.W. Wallroth.
Remarks: Pleospora can be separated from
Leptosphaeria by ascospore septation and
anamorphs. Some species included in
Leptosphaeria have ascospores with
longitudinal septa, and these species must be
examined carefully for ascomatal features
and cultured for anamorphic states (Miiller
1951, Wehmeyer 1961).

Pyrenophora E.M. Fries, Summa Vegetabilium

Scandinaviae, Seu Enumeratio, Systematica
et Critica, Plantarum tum Cotyledonearum,
Quam Nemearum Inter Mare Occidentale Et
Album, Inter Eidoram et Nordkop, Hactenus
Lectorum, una Cum Singulae Distributione
Geographica, pp. 397-398. 1849. Type
species: Pyrenophora phaeocomes (G.L.
Rabenhorst:E.M. Fries) E.M. Fries. Family:
Pyrenophoraceae. Ascomata: Immersed to
erumpent, medium to large, usually setose or
bearing conidiophores. Asci: Bitunicate,
few, basal, clavate, oblong or cylindric.
Ascospores: Ellipsoidal, broader above the
middle, muriform, yellowish-brown,
sheathed. Anamorph: Drechslera R.A.
Shoemaker. Remarks: Pyrenophora is
closely related to Pleospora and differs from
Leptosphaeria in having large, muriform
ascospores with gelatinous sheaths. The
genus is nomenclaturally sound with the
conservation of Ceuthospora R.K. Greville,
1826, typified by Ceuthospora lauri (R.K.
Greville) R.K. Greville vs. Ceuthospora E.M.
Fries, 1825, typified by Ceuthospora
phaeocomes (J.F. Rebentish:E.M. Fries) E.M.
Fries (Shoemaker 1961, Ammon 1963, Barr
1972, Sutton 1972).

Rebentischia P.A. Karsten, Fungi Fenniae Exsiccati,

Century 9, No. 881. Anno. 1869; Mycologia
Fennica, p. 14. 1873. Type species:
Rebentischia pomiformis P.A. Karsten.
Family: Tubeufiaceae. Ascomata: Erumpent,
globose, roughened with protruding cells and
hyphae, pseudoparenchymatic, outer cells
melanized black, beak absent. Asci: Many,
bitunicate, cylindric-clavate. Ascospores:
Obovate, vinaceous brown 4- to 5-septate,
with basal appendage. Anamorph: Un-
known. Remarks: Miiller (1950) considered
the centrum type of this genus to be similar
to that of a typical Leptosphaeria. It differs
from Leptosphaeria in the ascospores, which
have a hyaline basal appendage (Dennis
1978, Barr 1980).

Sulcispora R.A. Shoemaker, Canadian Journal of

Botany, Ottawa 67:1594. 1989. Type

346

Illinois Natural History Survey Bulletin

species: Sulcispora pleurospora (G. Niessl v.
Mayendorf) R.A. Shoemaker. Family:
Phaeosphaeriaceae. Ascomata: Immersed
becoming erumpent, pyriform to globose,
smooth. Beak: Composed of brown
rectangular cells, the ostiole lacking
periphyses. Ascoma wall: Brown pseudopar-
enchymatic, rectangular cells. Asci: Few,
bitunicate, cylindric, 8-spored. Ascospores:
Tetraseriate, fusiform, straight, 5- to 6-
septate, constricted at first septum, reddish-
brown, longitudinally sulcate, sheathed.

Trematosphaeria L. Fuckel, Symbolae Mycologi-

cae, pp. 161-162. 1870. Type species:
Trematosphaeria pertusa (C.H. Persoon:
E.M. Fries) L. Fuckel. Family: Plat-
ystomaceae. Ascomata: Superficial to
partially immersed, globose to conic,
peridium pseudoparenchymatic, melanized
black, textura angularis in face view, beak
short, papillate. Asci: Bitunicate, cylindric-
clavate, short-stalked, containing eight
ascospores. Ascospores: Fusiform, con-
stricted at mid-septum, 3-septate, brown.
Anamorph: Unknown. Remarks: Petrak
(1923) considered the structural features of
Trematosphaeria to correspond to those of

Vol. 34 Art. 3

Leptosphaeria, and he included wood-
inhabiting Leptosphaeria-like species. Boise
(1984) redefined the genus by the presence of
trabeculate pseudoparaphyses and included it
in the Platystomaceae in the Melanom-
matales.

Trichometasphaeria A. Munk, Dansk Botanisk

Arkiv, Kjobenhavn 15(2):135. 1953. Type
species: Trichometasphaeria dianthi (E.
Rostrup) A. Munk. [=7richometasphaeria
gloeospora (M.J. Berkeley & F. Currey) L.
Holm]. Family: Lophiostomataceae.
Ascomata: Globose, subepidermal, with
ostiole bearing brown setae. Asci: Bituni-
cate, clavate to cylindrical. Ascospores:
Hyaline, fusiform to elliptical, 4- to 6-
septate. Anamorph: Unknown. Remarks:
This genus is similar to Keissleriella, but it
differs in having several septate spores versus
1-septate spores in Keissleriella. Bose
(1961) united the two genera under the
earlier name, Keissleriella, after observing
variation in ascospore septation in
Keissleriella aesculi. Barr (1987b) accepts
both genera and places each of them ina
different order.

Appendix 3. Synonyms of Leptosphaeria

Ampullina L. Quélet (1875). Type: Ampullina
acuta =Leptosphaeria acuta (v. Arx and
Miiller 1975).

Baumiella P.C. Hennings in H. Baum (1903). Type
species: Baumiella caespitosa P. Hennings in
H. Baum, Kunene-Sambesi Expedition,
Berlin, p. 165. 1903; =Leptosphaeria baumii
J.A. v. Arx & E. Miiller (1975) nom. nov.,
non Leptosphaeria caespitosa G. Niessl v.
Mayendorf.

Bilimbiospora B. Auerswald in G.L. Rabenhorst
(1860). Nomina generica rejicienda by
conservation of Leptosphaeria (Greuter et al.
1988).

Chaetoplea (P.A. Saccardo) F.E. Clements in F.E.
Clements & C.L. Shear (1931). =Lepto-
sphaeria (Eriksson & Hawksworth 1986).

Chitonospora E.C. Bommer, M.H. Rousseau, &
P.A. Saccardo in P.A. Saccardo (1891).
=Leptosphaeria (Miiller 1950).

Dendroleptosphaeria M. de Sousa da Camara
(1932). A possible synonym of Lepto-
sphaeria (Eriksson & Hawksworth 1986).

Dothideopsella F. v. H6hnel (1915). =Lepto-
sphaeria (v. Arx & Miiller 1975).

Exilispora L.R. Tehon & E.Y. Daniels (1927).
Type species: Exilispora plurisepta L.R.
Tehon & E.Y. Daniels, Mycologia, Lancas-
ter, Pennsylvania 19:112. 1927; =Lepto-
sphaeria plurisepta (L.R. Tehon & E.Y.
Daniels) J.A. v. Arx & E. Miiller (v. Arx &
Miiller 1975).

Humboldtina C.E. Chardon & R.A. Toro (1934).
=Leptosphaeria (Eriksson & Hawksworth
1986).

Leptosporopsis F. vy. Héhnel (1920). =Lepto-
sphaeria (Eriksson & Hawksworth 1986).

Macrobasis K. Starback (1893). =Leptosphaeria
(Petrak and Sydow 1923, Miiller 1950).

Metasphaeria P.A. Saccardo (1883). Nomen
ambiguum. =Leptosphaeria (Dothideales) (v.
Arx and Miiller 1975); a segregate of
Leptosphaeria comprising the hyaline-spored
species, otherwise as in Leptosphaeria
(Petrak 1923, Miiller 1950). However, the
genus contains both unitunicate and bituni-
cate Ascomycetes (Barr 1976).

Mycopyrenula E.A. Vainio (1921). =Leptosphaeria
(Miiller 1950), but considered a good genus
by Hawksworth et al. (1983).

Mycotodea W. Kirschstein (1936). According to
Petrak (1940), Mycotodea is a synonym of
Scleropleella and thus Leptosphaeria
according to E. Miiller (1950); =Lepto-
sphaeria (Dothideales) (v. Arx and Miiller
1975).

Myriocarpium H.F. Bonorden (1864). =Lepto-
sphaeria (Eriksson & Hawksworth 1986).

Nodulosphaeria G.L. Rabenhorst (1858). Nomina
generica rejicienda by the conservation of
Leptosphaeria (Grueter 1988).

Phaeoderris (P.A. Saccardo) F. v. Hohnel (1907b).
=Leptosphaeria (v. Arx and Miiller 1975).

Phyllophthalmaria (J. Muller Argov) A.
Zahlbruckner in A. Engler & K. Prantl
(1905). A questionable synonym of
Leptosphaeria (Eriksson and Hawksworth
1986).

Pocosphaeria (P.A. Saccardo) A.N. Berlese (1892).
=Leptosphaeria (Miiller 1950, v. Arx and
Miiller 1975). Represents Leptosphaeria
species with a bristly peridium.

Saccothecium E.M. Fries (1835). =Pringsheimia S.
Schulzer v. Miiggenburg in S. Schulzer v.
Miiggenburg, A. Kanitz, and J.A. Knapp
(1866). Placed in the Dothideales (Holm
1975); =Massaria G. de Notaris (1844)
(Pyrenulales) (Barr 1979).

Sclerodothis F. v. Hohnel (1918a). Based on
hyaline spores and therefore identical with
Metasphaeria, which is a synonym of
Leptosphaeria (v. Arx & Miiller 1975).

Scleropleella F. v. Hohnel (1918a). =Leptosphaer-
ulina D. McAlpine (1902) (Dothideales)
(Barr 1972); considered a good section or
subgenus of Leptosphaeria by Miiller (1950);
species in this group form transitional taxa
within Leptosphaeria.

Syncarpella F. v. Theissen & H. Sydow (1915).
=Leptosphaeria (v. Arx and Miiller 1975).
Accepted by Barr 1987b.

Appendix 4. Anamorphs of Leptosphaeria

Ascochyta M.A. Libert (Coelomycete). Connection:
Leptosphaeria pratensis P.A. Saccardo &
P.A. Briard [=Ascochyta meliloti (W.
Trelease) J.J. Davis]. Source: Kendrick and
DiCosmo 1979. Conidiogenesis: Phialidic
with periclinal thickenings of apex of
phialide. Conidia: Hyaline, 1- or, rarely, 3-
septate. Comments: Other anamorphs
reported for Leptosphaeria pratensis P.A.
Saccardo & P.A. Briard are Phoma meliloti
A. Allescher and Stagonospora meliloti
(W.G. Lasch) F. Petrak, both Coelomycetes
(Lucas and Webster 1967).

Ascochytula (A.A. Potebnia) H. Diedicke [=Pseu-
dodiplodia (P.A. Karsten) P.A. Saccardo]}
(Coelomycete). Connection: Leptosphaeria
obiones (H.M. Crouan & P.L. Crouan) P.A.
Saccardo (=Ascochytula obiones H.
Diedicke). Source: Grove 1935 (by
association). Conidiogenesis: Phialidic with
periclinal thickening of apex of phialide.
Conidia: Phaeodidymospores.

Asteromella G. Passerini & F. v. Thiimen (Coelo-
mycete). Connection: Leptosphaeria
artemisiae (L. Fuckel) B. Auerswald
(=Asteromella artemisiae E. Miiller).
Source: Miiller 1950, Lucas and Webster
1967. Conidiogenesis: Phialidic with
periclinal thickening of apex of phialide.
Conidia: Hyaloamerospores.

Camarosporium §. Schulzer vy. Miiggenburg
(Coelomycete). Connection: Leptosphaeria
maculans (J. Desmazieres) V. Cesati & G. de
Notaris [=Camarosporium affine (P.A.
Saccardo) E.C. Bommer & M.H. Rousseau]
(Coelomycete). Source: Miller and
Tomasevie 1957; Connection: Leptosphaeria
millefolii (L. Fuckel) G. Niessl v. Mayendorf
(=Camarosporium sp.). Source: Miiller and
Tomasevic 1957; Connection: Leptosphaeria
ogilviensis (M.J. Berkeley & C.E. Broome)
V. Cesati & G. de Notaris (=Camarosporium
sp.). Source: Miller and TomaSevié 1957;
Connection: Leptosphaeria orthosanthi E.
Miller (=Camarosporium sp.). Source:
Miller and Dennis 1965; Conidiogenesis:
Annellidic. Conidia: Phaeodictyospores.

Cladosporium J.H. Link (Hyphomycete).
Connection: Leptosphaeria ladina E. Miiller
(=Cladosporium ladium E. Miller). Source:
Miiller 1950. Conidiogenesis: Holoblastic,
sympodial. Conidia: Brown, o-pluriseptate.
Comments: Considered a culture contaminant
rather than an anamorph (Kendrick and
DiCosmo 1979).

Coniothyrium (A.C.J. Corda) (Coelomycete).
Connection: Leptosphaeria bondari A.A.
Bitancourt & A.E. Jenkins (=Coniothyrium
sp.). Source: Wehmeyer 1975; Connection:
Leptosphaeria coniothyrium (L. Fuckel) P.A.
Saccardo (=Coniothyrium fuckelii P.A.
Saccardo). Source: Zeller 1927, Punithal-
ingam 1980; Connection: Leptosphaeria
faullii G.D. Darker (=Coniothyrium faullii
G.D. Darker). Source: Darker 1964,
Wehmeyer 1975; Connection: Leptosphaeria
michotii (G.D. Westendorp) P.A. Saccardo
(=Paraphaeosphaeria O. Eriksson: =Conio-
thyrium scirpi J.W. Trail). Source:
Wehmeyer 1975; Sivanesan 1984;
Connection: Leptosphaeria obiones (H.M.
Crouan & P.L. Crouan) P.A. Saccardo
(=Coniothyrium obiones H. Diedicke).
Source: Grove 1935; Connection: Lepto-
sphaeria perichymeni C.A. Oudemans
(=Coniothyrium sp.). Source: Wehmeyer
1975; Connection: Leptosphaeria spartinae
J.B. Ellis & B.M. Everhart (=Coniothyrium
sp.). Source: Lucas and Webster 1967;
Conidiogenesis: Annellidic. Conidia:
Phaeoamerospores, phaeodidymospores.

Diplodina G.D. Westendorp (Coelomycete).
Connection: Leptosphaeria marcyensis (C.H.
Peck) P.A. Saccardo (=Diplodina sp.).
Source: Lucas and Webster 1967. Conidio-
genesis: Phialidic, periclinal thickening of
apex of phialide, eustomatic. Conidia:
Hyaline, 0- to 2-septate.

Hendersonia M.J. Berkeley. Nomina generica
rejicienda. [=Stagonospora (P.A. Saccardo)
P.A. Saccardo] (Coelomycete).

Leptophoma F. v. Héhnel (=Phoma P.A. Saccardo)
(Coelomycete).

Microdiplodia A. Allescher (Coelomycete).
Connection: Leptosphaeria obtusispora C.L.
Spegazzini [=Microdiplodia henriquesii (F.

March 1991 Nomenclator of Leptosphaeria 349

v. Thiimen) F. Petrak & H. Sydow]. Source:
Lucas 1963. Conidiogenesis: Unknown in
Microdiplodia henriquesii (F. v. Thiimen) F.
Petrak & H. Sydow. Conidia: Phaeodi-
dymospores.

Nakataea K. Hara (Hyphomycete). Connection:
Leptosphaeria salvinii A. Cattaneo
(=Nakataea sigmoidea K. Hara). Source:
Ellis 1971. Conidiogenesis: Holoblastic,
sympodial. Conidia: 3-septate, pale brown.

Pestalotia G. de Notaris (Coelomycete).
Connection: Leptosphaeria honiarensis T.
Matsushima (=Pestalotia sp.). Source:
Matsushima 1971. Conidiogenesis:
Annellidic. Conidia: Phaeophragmospores
with appendages. Comments: Leptosphaeria
honiaraensis may belong in Pestalosphaeria
M.E. Barr (Amphisphaeriaceae) (Kendrick
and DiCosmo 1979).

Pestalotiopsis R.L. Steyaert (Coelomycete).
Connection: Leptosphaeria elaeidis C. Booth
& J.S. Robertson. Source: Booth and
Robertson 1961. Conidiogenesis: Holoblas-
tic, annellidic. Conidia: Phaeo-
phragmospores with appendages.

Phaeoseptoria C.L. Spegazzini (Coelomycete).
Connection: Leptosphaeria culmifraga (E.M.
Fries:E.M. Fries) V. Cesati & G. de Notaris
(=Phaeoseptoria sp.). Source: Wehmeyer
1975; Connection: Leptosphaeria fuckelii G.
Niessl v. Mayendorf (=Phaeoseptoria sp.).
Source: Webster and Hudson 1957, Sivane-
san 1984; Connection: Leptosphaeria
lactuosa G. Niess| v. Mayendorf
(=Phaeoseptoria sp.). Source: Webster and
Hudson 1957, Sivanesan 1984; Connection:
Leptosphaeria macrospora (L. Fuckel) F. v.
Thiimen (=Phaeoseptoria sp.). Source:
Lucas and Webster 1967; Connection:
Leptosphaeria microscopica P.A. Karsten
[=Phaeoseptoria airae (W.B. Grove) R.
Sprague]. Source: Sivanesan 1984;
Connection: Leptosphaeria nigrans (M.R.
Roberge) V. Cesati & G. de Notaris

(=Phaeoseptoria sp.). Source: Hughes 1949,

Sivanesan 1984; Conidiogenesis: Holoblas-
tic. Conidia: Phaeophragmospores.

Phoma P.A. Saccardo (Coelomycete). Connection:
Leptosphaeria acuta L. Fuckel (=Phoma
acuta L. Fuckel). Source: Grove 1935;
Miiller and Tomasevic 1957; Connection:
Leptosphaeria agnita (J. Desmaziéres) V.
Cesati & G. de Notaris (=Phoma sp.).
Source: Lucas and Webster 1967, Sivanesan
1984; Connection: Leptosphaeria albo-
punctata (G.D. Westendorp) P.A. Saccardo
(=Phoma sp.). Source: Sivanesan 1984;
Connection: Leptosphaeria conferta G.
Niessl v. Mayendorf ex P.A. Saccardo
(=Phoma sp.). Source: Lucas 1963, Sivane-

san 1984; Connection: Leptosphaeria
congesta M.T. Lucas (=Phoma sp.). Source:
Lucas 1963; Connection: Leptosphaeria
cruenta P.A. Saccardo (=Phoma sanguino-
lenta E. Rostrup). Source: Grove 1935;
Connection: Leptosphaeria doliolum (C.H.
Persoon:E.M. Fries) subsp. doliolum var.
doliolum V. Cesati & G. de Notaris (=Phoma
hoehnelii H.A. Van Kerstern subsp. hoehnelii
var. urticae G.H. Boerema & H.A. Van
Kerstern). Source: Lucas and Webster 1967,
Boerema 1976; Connection: Leptosphaeria
dumetorum G. Niessl v. Mayendorf
(=Phoma sp.). Source: Lucas and Webster
1967; Connection: Leptosphaeria haematites
M.R. Roberge ex J. Desmaziéres) G. Niessl
v. Mayendorf (=Phoma sp.). Source: Lucas
and Webster 1967, Sivanesan 1984;
Connection: Leptosphaeria libanotis (L.
Fuckel) G. Niessl v. Mayendorf (=Phoma
sp.). Source: Lucas and Webster 1967,
Sivanesan 1984; Connection: Leptosphaeria
lindquistii M.J. Frezzi (=Phoma macdonaldii
G.H. Boerema). Source: Sivanesan 1984;
Connection: Leptosphaeria maculans (J.
Desmaziéres) V. Cesati & G. de Notaris
{=Phoma lingam (H.J. Tode:E.M. Fries)].
Source: Miller and TomaSevié 1957,
Sivanesan 1984; Connection: Leptosphaeria
millefolii (L. Fuckel) G. Niessl v. Mayendorf
(=Phoma sp.). Source: Miiller and
Tomasevic 1957; Connection: Leptosphaeria
ogilviensis (M.J. Berkeley & C.E. Broome)
V. Cesati & G. de Notaris (=Phoma sp.).
Source: Miiller 1971; Connection: Lepto-
sphaeria pontiformis (L. Fuckel) P.A.
Saccardo (=Phoma sp.). Source: Webster
and Hudson 1957, Wehmeyer 1975;
Connection: Leptosphaeria purpurea H.
Rehm (=Phoma sanguinolenta W.B. Grove).
Source: Lucas and Webster 1967, Sivanesan
1984; Connection: Leptosphaeria sacchari
H.J. Van Breda De Haan (=Phoma sp.; as
Phyllosticta sp.). Source: Hudson 1960;
Connection: Leptosphaeria solani L.G.
Romell (=Phoma sp.). Source: Lucas and
Webster 1967; Connection: Leptosphaeria
spartinae J.B. Ellis & B.M. Everhart
(=Phoma sp.). Source: Sivanesan 1984;
Connection: Leptosphaeria submaculans L.
Holm (=Phoma sp.). Source: Lucas and
Webster 1967; Connection: Leptosphaeria
thomasiana P.A. Karsten (=Phoma sp.).
Source: Zeller 1927 (by association), Lucas
and Webster 1967 (unnamed); Connection:
Leptosphaeria typhicola P.A. Karsten
(=Phoma sp.). Source: Lucas and Webster
1967 (unnamed), Sivanesan 1984; Conidio-
genesis: Phialidic with periclinal thickening
of apex of phialide. Conidia: Phaeoamero-
spores.

350

Illinois Natural History Survey Bulletin

Scolecosporiella F. Petrak (Coelomycete).

Connection: Leptosphaeria macrospora (L.
Fuckel) F. v. Thiimen [=Scolecosporiella
bernardiana (P.A. Saccardo) A. Sivanesan].
Source: Sivanesan 1984; Connection:
Leptosphaeria typharum (J. Desmazieéres)
P.A. Karsten [=Scolecosporiella typhae
(C.A. Oudemans) F. Petrak]. Source:
Sivanesan 1984; Conidiogenesis: Holoblas-
tic. Conidia: Pale brown, 3- to many-septate,
cylindrical.

Septoria P.A. Saccardo (Coelomycete).

Connection: Leptosphaeria avenaria G.F.
Weber (=Septoria avenae B. Frank). Source:
Grove 1935, Sivanesan 1984; Connection:
Leptosphaeria maydis G.L. Stout (=Septoria
zeae G.L. Stout). Source: Stout 1930 (by
association); Connection: Leptosphaeria
nodorum E. Miller [=Septoria nodorum
(M.J. Berkeley) M.J. Berkeley]. Source:
Lucas and Webster 1967, Sivanesan 1984;
Connection: Leptosphaeria phlogis C.A.
Oudemans (=Septoria phlogis P.A. Saccardo
& C.L. Spegazzini). Source: Grove 1935 (by
association); Connection: Leptosphaeria
sorbi A.L. Jaczewski (=Septoria sorbi W.G.
Lasch). Source: Grove 1935 (by associa-
tion); Conidiogenesis: Three types of
conidiogenesis have been found for the
pathogenic species of Septoria studied thus
far. They are (1) holoblastic, sympodial, (2)
simple holoblastic, and (3) phialidic with
periclinal thickening of the apex of the
phialide. Conidiogenesis has not been
determined for any of the Seproria ana-
morphs of Leptosphaeria. Conidia: Hyaline,
multiseptate, filiform.

Stagonospora (P.A. Saccardo) P.A. Saccardo

(Coelomycete). Connection: Leptosphaeria
anemones L. Hollés (=Stagonospora
anemones N.T. Patouillard). Source:
Sivanesan 1984; Connection: Leptosphaeria
arundinaceae P.A. Saccardo (=Stagonospora

Vol. 34 Art. 3

vexata P.A. Saccardo). Source: Grove 1935
(by association); Connection: Leptosphaeria
bicolor D.L. Hawksworth, W. Kaiser & B.N.
Ndimande (=Stagonospora sp.). Source:
Kaiser et al. 1979; Connection: Lepto-
sphaeria cistina M.J. de Urries y Azara
(=Stagonospora sp.; as Hendersonia cisti M.
de Sousa da Camara). Source: Lucas 1968:
Connection: Leptosphaeria dumetorum G.
Niessl v. Mayendorf (=Stagonospora sp.; as
Hendersonia sp.). Source: Lucas and
Webster 1967, Sivanesan 1984; Connection:
Leptosphaeria eustomoides P.A. Saccardo
(=Stagonospora sp.; as Hendersonia sp.).
Source: Webster and Hudson 1957;
Connection: Leptosphaeria gigaspora G.
Niessl v. Mayendorf (=Stagonospora gi-
gaspora P.A. Saccardo). Source: Grove
1935 (by association); Connection: Lepto-
sphaeria libanotis (L. Fuckel) G. Niess! v.
Mayendorf (=Stagonospora sp.: as
Hendersonia sp.). Source: Wehmeyer 1975;
Connection: Leptosphaeria polygonati E.
Miiller & M. TomaSevic¢ (=Stagonospora
sp.; as Hendersonia sp.). Source: Miiller and
TomaSevic 1957; Connection: Leptosphaeria
pontiformis (L. Fuckel) P.A. Saccardo
(=Stagonospora sp.; as Hendersonia sp.).
Source: Lucas and Webster 1967, Sivanesan
1984; Connection: Leptosphaeria pratensis
P.A. Saccardo & P.A. Briard [=Stagono-
spora melioti (W.G. Lasch) F. Petrak].
Source: Jones and Weimar 1938, Lucas and
Webster 1967; Connection: Leptosphaeria
taiwanensis W.Y. Yen & C.C. Chi (=Stag-
onospora taiwanensis W.H. Hsieh. Source:
Hsieh 1979, Sivanesan 1984; Connection:
Leptosphaeria viridella (C.H. Peck) P.A.
Saccardo (=Stagonospora sp.; as Hender-
sonia). Source: Lucas and Webster 1967:
Conidiogenesis: Holoblastic, sometimes
annellidic. Conidia: Hyaline, multiseptate,
cylindrical.

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Manuscripts of high quality dealing with any aspect of natural history will be considered for
publication in one of the Illinois Natural History Survey series: Bulletin, Biological Notes,
Circular, and Special Publications. Authors who are not employees of the Survey are required to
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full. The Survey expects to publish only one or two manuscripts by non-Survey authors each year.
Send three copies of manuscripts to be considered for publication to Office of the Chief, Illinois
Natural History Survey, 607 East Peabody Drive, Champaign, Illinois 61820. Before a manuscript
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Illinois Natural History Survey
607 East Peabody Drive
Champaign, Illinois 61820
217-333-6880

A Division of the Illinois Department of Energy and Natural Resources

hael R. Jeffords
iois Natural History Survey

ILLINOIS
NATURAL
HISTORY
SURVEY

NOV 20 1991

LIBRARY

ILLINOIS
NATURAL
HISTORY
SURVEY

Our Living Heritage:
The Biological Resources of Illinois

Edited by

Lawrence M. Page

Michael R. Jeffords

Illinois Natural History Survey

Proceedings of a symposium in celebration of Earth Day 1990
Illinois Department of Energy and Natural Resources

Illinois Natural History Survey

April 23 and 24, 1990

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

Printed by Authority of the State of Illinois
X12275-2M-4-91
US ISSN 0073-4918

Graphic Design: Gail Glende Rost
Computer Graphics: Molly Hardin Scott
Editor: Audrey S. Hodgins

A catalog of the publications of the Illinois Natural History Survey is available
without charge from the address below. A price list and order blank are
included with the catalog.

Illinois Natural History Survey

Distribution Center

607 East Peabody Drive

Champaign, Illinois 61820

Manuscripts of high quality dealing with any aspect of natural history will be
considered for publication in one of the Illinois Natural History Survey series:
Bulletin, Biological Notes, Circular, and Special Publication. Authors who are
not employees of the Survey are required to pay printing costs. Manuscripts
should follow the recommendations of the third edition of the Council of
Biological Editors Style Manual except that journal names in literature cited are
to be spelled in full. The Survey expects to publish only one or two manuscripts
by non-Survey authors yearly. Send three copies of manuscripts to be consid-
ered for publication to Office of the Chief, Illinois Natural History Survey,

607 East Peabody Drive, Champaign, Illinois 61820. Before a manuscript is
accepted for publication in the Bulletin or Biological Notes, it must be recom-
mended by two or more outside referees.

Citation: Page, L.M., and MLR. Jeffords, eds. 1991. Our living heritage: the
biological resources of Illinois. Illinois Natural History Survey Bulletin 34(4):
357-477.

Foreword

We live in a world of near continuous monitor-
ing. In our automobiles we monitor the status
of fuel, oil pressure, temperature, and seat belts
through gauges, lights, and electronic voices.
The consumption of electricity and fuel in our
homes is monitored as is the chlorine in our
drinking water and the alcohol in our beer.
Manufacturers retain quality assurance inspec-
tors and issue warrantees and guarantees to
convince us that all is well. We monitor our
schools and measure our own progress through
grades and proficiency scores. It seemed
appropriate, therefore, that the Illinois Natural
History Survey should take a measure of the
living natural resources of Illinois by bringing
together a knowledgeable group of persons to
summarize the state of the State. In order to
share this information and to provide an
opportunity for discussion, a symposium, “Our
Living Heritage: The Biological Resources of
Illinois,” was sponsored by the Illinois Depart-
ment of Energy and Natural Resources and
organized by the Survey. The event, timed to
coincide with Earth Day 1990 celebrations, was
held on April 23 and 24 on the campus of the
University of Illinois at Urbana-Champaign. It
was attended by nearly 250 professional
scientists from some 50 agencies and institu-
tions along with a number of interested and
dedicated citizens. To share the results of that
symposium with an even larger audience, we
have issued this publication of its proceedings.
To address the salient features of the
living resources of Illinois in an ordered
fashion, the symposium was presented in five
sessions: forests, prairies and barrens, wetlands,
streams and caves, and agro-urban ecology.
When we consider that only 0.5% of Illinois
remains in undisturbed natural areas, that
Illinois ranks 46th among states in publicly
owned open space per person, that forest
acreage has decreased by 73% in the past
century and tallgrass prairie by over 99%, that

85% of our wetlands have been lost, that soil
erosion proceeds at the rate of 200 million tons
per year, and that approximately 30,000 tons of
herbicide and 3,500 tons of insecticides are
used annually on agricultural crops in Illinois,
we can scarcely imagine the tone of the
symposium to have been anything but pessi-
mistic. In part, there was discouragement, but it
was tempered by positive developments,
including the designation of the Middle Fork of
the Vermilion River as a National Wild and
Scenic River, the acquisition of the Cache
River Basin, the initiation of a study to identify
high-quality Illinois streams based on biodiver-
sity, and the ever quickening actions of the
Nature Preserves Commission.
Preservation/conservation has been in
conflict with consumption/development since
the days of Theodore Roosevelt. At times one
side seems to prevail over the other, but the
balance has been clearly on the side of con-
sumption. Special interest groups have to a
considerable extent managed to give the word
environmentalist a pejorative cast and the word
development a positive ring. During the past
decade, the executive branch of the federal
government has determinedly downplayed
environmental concerns, and that stance has
been translated into inertia in a number of
federal agencies with responsibility for natural
resources. The focus of the United States
Environmental Protection Agency, for example,
has until very recently ignored the living
components of the environment. At the same
time, public sensitivity to environmental
concerns has dramatically increased, primarily
through public service television and other
media-generated presentations on tropical
deforestation, extinction of species, depletion
of the ozone layer, agro-chemical contamina-
tion of groundwater, and the effects of acid
rain. Some of this concern is now being
transformed into political action. Polls suggest

ill

that the public understanding of environmental
matters is quite high, and some believe that it
exceeds the perceptions of elected officials. A
Green Party has emerged in this country only
very recently, but Greens are a part of both
major political parties and the trend in federal
legislation may soon begin to sway in favor of
conservation/preservation and away from
consumption/development. The National
Institutes for the Environment may well
become a reality within the next several years.
Within this tentatively encouraging national
picture, the symposium was timely indeed.

One symposium event of special interest
cannot be documented in these proceedings—
the “citizens respond” program of Monday
evening, April 23—and I would like to note it
here. Michael Jeffords and Susan Post of the
Survey opened that session with a mulitmedia
presentation on the biodiversity of Illinois.
Their slides of representative plants and
animals and habitats of the natural divisions of
Illinois brought home to us the beauty and
fragility that can yet be discovered in the
landscape of our state. A panel presentation by
five environmental activists followed: Clark
Bullard, Office of Energy Research at the
University of Illinois at Urbana-Champaign;
Max Hutchison, Natural Land Institute of The
Nature Conservancy; Lawrence Page of the
Illinois Natural History Survey; Donna
Prevedell, farmwife and contributing editor to
the Progressive Farmer; and Michael Reuter,
Volunteer Stewardship Network of The Nature
Conservancy. They spoke briefly but openly on
preservation activities in which they had been
closely involved. The discussion was then
turned over to the audience, who asked ques-
tions and shared their experiences—successes
and failures—with preservation efforts.

I urge you to read on in order to under-
stand the status of the biological resources of
Illinois and to appreciate how much remains to
be accomplished to secure their future—and
ours. I would be remiss, however, if I did not
conclude by acknowledging the committee of
Survey staff who planned and conducted the
symposium: Lawrence Page, Michael Jeffords,
Joyce Hofmann, Susan Post, Louis Iverson, and
Audrey Hodgins. Their efforts included
developing the program, arranging for speakers
and facilities, producing and mailing promo-
tional materials, and welcoming the audience.

Without their enthusiasm and hard work, the
symposium would not have materialized and
our understanding of the biological resources of
Illinois would be much diminished.

Lorin I. Nevling, Chief
Illinois Natural History Survey

Contents

FOREWORD _ iii
INTRODUCTION 357

SESSION ONE: FORESTS = 359

Forest Resources of Illinois: What Do We Have and What Are They
Doing for Us? Louis R. Iverson 361

Forest Succession in the Prairie Peninsula of Illinois John E. Ebinger
and William E. McClain 375

Effects of Forest Fragmentation on Illinois Birds
Scott K. Robinson 382

SESSION TWO: PRAIRIES AND BARRENS = 383

Illinois Prairies: A Historical Perspective Roger C. Anderson 384
Prairie and Savanna-restricted Insects of the Chicago Region
Ron Panzer 392
Prairie Birds of Illinois: Population Response to Two Centuries of
Habitat Change James R. Herkert 393

SESSION THREE: WETLANDS 400

Aquatic and Wetland Plants of Illinois John E.Schwegman 401
Breeding Biology and Larval Life History of Four Species of
Ambystoma (Amphibia: Caudata) in East-central Illinois
Michael A. Morris 402
Ecological Integrity of Two Southern Illinois Wetlands
M. Ann Phillippi 403
Status and Distribution of Wetland Mammals in Illinois
Joyce E. Hofmann 409

SESSION FOUR: STREAMS AND CAVES 416

The Fishes of Illinois: An Overview of a Dynamic Fauna
Brooks M. Burr 417

The Aquatic Mollusca of Illinois Kevin S.Cummings 428

Streams of Illinois Lawrence M. Page 439

Illinois Caves: A Unique Resource James E. Gardner 447

SESSION FIVE: AGRO-URBAN ECOLOGY 453

The Land Use Controversy: Maintaining and Increasing Biotic
Diversity in the Agricultural Landscape of Illinois
Michael E. Irwin 454

Farm Programs, Agricultural Technologies, and Upland Wildlife
Habitat Richard E. Warner 457

Evaluating Alternatives for Urban Deer Management
James H. Witham 458

Illinois Railbanking Study Richard Pietruszka 459

Closing Remarks Brian D. Anderson 460

APPENDICES

Appendix One: Native Illinois Species and Related
Bibliography Susan L. Post 463
Appendix Two: County Reference Map 476

Introduction

The term biodiversity has not yet made its way
into most dictionaries, but the word is generally
accepted to mean the organisms that inhabit the
Earth and the ecosystems in which they live.
Lying at the junction of the eastern forest,
western great plain, southern coastal plain,
Ozark uplift, and northern forest biomes, Illinois
provides habitat for an extremely varied native
flora and fauna. Scientists at the Illinois Natural
History Survey recently compiled data on the
biodiversity of Illinois and conservatively
estimated that more than 53,000 species are
native to the state (Appendix I). The largest
groups are insects with about 17,000 species and
fungi with about 20,000 species. In addition,
Illinois is home to 2,068 species of vascular
plants and 649 species of vertebrates (mammals,
birds, reptiles, amphibians, and fishes).

The biodiversity of Illinois is more readily
appreciated when it is compared to that of other
regions. Consider, for example, that the Pine
Hills—LaRue Swamp region of southwestern
Illinois contains about 1,000 native species of
plants. The Great Smoky Mountains National
Park, an area of wilderness about 260 times
larger, contains only 1,200 native plant species.
That same region of southwestern Illinois also
has more amphibian and reptile species (61)
than are found in any region of comparable size
in the United States. Perhaps equally surprising,
one-fourth of all the freshwater fishes and
mussels of North America north of Mexico are
found in Illinois.

The destruction of tropical rainforests,
which are thought to contain over half the total
species of organisms, has been widely publi-
cized, but all ecosystems are threatened as
human populations and their support systems
expand. Illinois, one of the most altered regions
on Earth, is experiencing an ongoing and
accelerating loss in variety as well as absolute
numbers of organisms. At least 115 species are
known to have been extirpated in recent decades

(Appendix I), and another 497 are officially
listed in Illinois as threatened or endangered.
Unless circumstances change dramatically,
Illinois will soon have lost 1 in 5 of its native
species of fishes, 1 in 5 of its native flowering
plants, | in 5 of its native birds, | in 4 of its
native mammals, and a startling one-half of its
native freshwater mussels!

Historical accounts of Illinois noted
huge trees, vast grasslands, and extensive
wetlands. Illinois was chiefly a combination of
flat, mesic, “marshy” prairies and forested
hilly country. Interspersed in these habitats
were sand dunes, bogs, fens, sedge meadows,
savannas, and swamps. Unfortunately, little of
that original landscape remains. In fact, [llinois
ranks an unenviable 49th among states in the
percentage of natural areas surviving. Of the
original 22 million acres of prairie, only 2,300
acres (0.01%) remain. Of the 14 million acres
of forest present in Illinois in 1820, only
13,500 acres of primary (undisturbed) forest
survive (0.10%). Many of our wetlands have
been, and continue to be, drained before they
can be biologically inventoried and their value
determined. Our streams are polluted and
increasingly degraded by the influx of soil
from surrounding farmland. A significant
portion of the biodiversity of Illinois will soon
disappear unless the remaining species-rich
areas are protected.

Several factors contribute to the global
loss of biodiversity: the explosive growth of
the human population, widespread and
extreme poverty and malnutrition, and a
notable lack of sustainable, productive agricul-
tural and forest systems in many regions of the
world. This loss is of paramount importance
because human existence depends on the
biological resources of the planet. Our
prosperity and well-being are based largely on
our ability to take advantage of the properties
of plants, animals, and microorganisms for

357

358 Illinois Natural History Survey Bulletin

food, clothing, medicine, and shelter. As
species are lost, we reduce our options for
future development of vital commodities. As
habitats and ecosystems are lost, we lose the
recreational potential of wild places, and we
disturb the balance of atmospheric gases,
including oxygen, carbon dioxide, and ozone.
Although the link between biodiversity and
human survival is clear, we must also learn to
value the biodiversity of our planet and state
for its own sake, quite apart from direct
benefits to us.

The loss of biodiversity is a global
problem, but the loss of Illinois biodiversity is
of special concern to Illinoisans. In our state,
the major cause of the loss of species is the
destruction and degradation of habitat. The
anthropogenic changes associated with agricul-
ture and urbanization cause environmental
degradation and lead to the extincfion of
species. If the loss of its native biodiversity is
not halted, Illinois could become a biological
desert unable to respond to the need for new
products and incapable of developing resource-
based solutions to human problems. At issue is
how we will protect the natural habitats that
remain, restore some of the natural areas that
have been lost, and balance the protection of
biodiversity against conflicting social and
economic interests. If we are to make informed
decisions, we must first complete the following
tasks.

Inventory the biological resources of
Illinois. Our knowledge about the biodiversity
of Illinois is incomplete. This lack of informa-
tion hampers our ability to estimate the size and
nature of the problem and to recommend
remedial measures. We are unable to identify
all the biological resources at risk because no
inventory of all life forms exists. Although our
knowledge of some taxa is extensive, other
groups are largely unknown. Species are lost
before they are discovered and studied. Even in
groups that are well studied (e.g., birds and
fishes), changes are occurring so rapidly that
additional data are needed if wise decisions
relative to development and management are to
be made.

Develop the scientific base on which
the emerging fields of conservation biology,
restoration ecology, and environmental
management can be built. Recent global and
regional environmental changes and the

Vol. 34 An. 4

inevitability of future modifications underscore
the need for prudent decisions regarding the
protection and use of natural resources. Indices
are needed that will enable us to compare
habitats and select outstanding natural areas for
management and protection.

Educate Illinoisans regarding the im-
portance of biological diversity. Biodiversity
is of particular interest to biologists and
ecologists, but all citizens must be informed
about the global biodiversity crisis if protective
legislation is to be enacted and funding
ensured.

Encourage socio-economic research
related to the wise use of biodiversity. We
need theoretical and empirical studies on the
economic and social causes of the biodiversity
crisis, its consequences, and its remedies.

Sponsored by the Department of Energy
and Natural Resources and the Illinois Natural
History Survey, the symposium “Our Living
Heritage: The Biological Resources of Illinois”
was held in celebration of Earth Day 1990 on
the Urbana-Champaign Campus of the Univer-
sity of Illinois. Two days, April 23 and 24,
were spent reviewing present information about
the biodiversity of Illinois and identifying
actions necessary to understand and conserve
the remaining resources of our state. Sessions
were arranged by ecosystem (forests, prairies
and barrens, wetlands, streams, caves, and
agro-urban habitat), and contributors discussed
what is known about how these ecosystems
function, how they have been modified, and
how various decisions are likely to affect their
survival. The proceedings that follow summa-
rize information on the biodiversity of Illinois
and suggest where additional research is
needed. Nineteen of the twenty-two presenta-
tions delivered at the symposium are included
here, either as abstracts or papers.

Although the audience agreed that more
information on certain subjects and groups of
organisms is needed, they also acknowledged
that we know enough to conclude that we have
already drastically altered most of our native
landscape and that we are rapidly losing native
species. Without greater protection and more
extensive management of natural areas, the loss
of habitats and species can only accelerate.

Session One: Forests

Like the first farmsteads, towns of the frontier were built in stumpland meadows. The trees were gone. The
civic landscapes sweltered in the sun. Never so quick an afterthought: fast-growing black locust trees were
imported and planted everywhere, from college campuses to courthouse squares, to provide a promise of
shade. What irony—the sons of the world’s most incredible axemen planting seedlings in the shadow of

stumps five feet across.—Robert O. Petty

In 1820, approximately 13.8 million acres of
Illinois were forested. The midcontinental
location of the state and its north to south
distance of nearly 400 miles allowed an
unusual variety of forest types to exist. The pre-
settlement forests of Jo Daviess County
covered nearly 80% of the land surface and
were noted for their rugged topography and the
presence of Pleistocene relic species. In 1830, a
U.S. Government geologist surveying the
Grand Prairie Division in central Illinois
observed, “Sometimes the woodland extends
along this river for miles continuously, again it
stretches in a wide belt off into the country,
marking the course of some tributary streams,
and sometimes in vast groves of several miles
in extent, standing alone, like islands in the
wilderness of grass and flowers.” Robert
Ridgway, a Smithsonian naturalist, noted the
immense size and diversity of the trees along
the lower Wabash Valley in the 1870s. With
photographs and measurements, he documented
the extraordinary nature of the bottomlands. In
the Shawnee Hills the relatively broad, flat-
bottomed ravines, originally cut by the melt-
waters of the Illinoian glacier, were verdant,
damp jungles filled with trees—beech, sugar
maple, and tulip—that reached and overtopped
the sandstone bluffs. South of the Shawnee
Hills the terrain flattened and a distinctly
southern forest grew in the past and present
Ohio River valleys. Great expanses of bald
cypress—water tupelo swamps filled the
lowlands along the Cache and Ohio rivers. Rare
species like willow oak, silverbell, water
hickory, and American chestnut occupied river
terraces, flatwoods, and ravines.

We know of these magnificent forests for
several reasons. Early settlers to Illinois, while
greatly impressed with the vast expanse of
prairie, chose to live in the woodlands, a
landscape with which Europeans felt more

familiar. Thus the nature of these forests came
to be better documented than that of other
landscape types. In addition, early biologists
like Ridgway and the St. Louis physician
George Engelmann described the presettlement
condition of Illinois forests in considerable
detail.

To begin to understand the current
condition of Illinois forests we must reflect
upon their past and on what has been lost.
Robert Ridgway, writing in the American
Naturalist in the 1870s, described the forests
along the Wabash River. “If the forest is
viewed from a high bluff, it presents the ap-
pearance of a compact, level sea of green,
apparently endless . . . the tree-tops swaying
with the passing breeze, and the general level
broken by occasional giant trees which rear
their massive heads so as to overlook the
surrounding miles of forest . . . while the
occasional, and by no means infrequent,
‘monarchs’ which often tower apparently for
one-third their height above the tree-top line,
attain an altitude of more than one hundred and
eighty feet, or approach two hundred feet.” In
the visitor center of Beall Woods, an Illinois
Nature Preserve in Wabash County, an im-
mense yellow outline painted on the floor
represents one of these last great trees. The
circle is seventeen feet in diameter.

Today nearly 4.3 million acres of trees
can be found in Illinois, not too startling a
decline in acreage from 1820 if we consider the
agricultural and urban development that now
blankets the state. Lest we are too complacent,
however, we should recall that much of the
forest acreage of today is second- or third-
growth timber or pine plantations; only 13,500
acres of relatively undisturbed forests remain—
a shockingly small percentage of our rich,
forested heritage. Fortunately, fragments
remain of nearly all forest types found in

Se})

360 Illinois Natural History Survey Bulletin

presettlement times and these, in conjunction
with land survey records, early written ac-
counts, and good biological detective work,
allow us to mentally reconstruct, and some-
times physically restore, the various forest
habitats. These efforts, to some extent, provide
a glimpse of what was once Illinois.

The three papers given at this session
help us to conceptualize the forests that were
once so integral to the Illinois landscape and to
understand how the forests that exist today
came to be. In addition, they enable us to
appreciate the role that forests play in the
economy of the state, in preserving biodiversity
and habitat for wildlife, in controlling erosion
and improving the quality of surface water, and
in conserving energy and slowing global
warming.

Vol. 34 An. 4

Forest Resources of Illinois:

What Do We Have and What Are They Doing for Us?

Louis R. Iverson, Illinois Natural History Survey

Forests occupy only a relatively small propor-
tion (12%) of the land area of Illinois (Figure
1), yet they provide tremendous benefits to the
citizens of the state. We need only walk
through the woods to be aware of some of these
benefits: aesthetic beauty, habitat for special-
ized plants and for birds and other wildlife,
recreational opportunities, and high-quality
hardwood. The more subtle but equally impor-
tant benefits that forest ecosystems provide,
however, are not so readily perceived. Forested
acres, for example, dramatically inhibit soil
erosion, thereby reducing the sediment load
that eventually finds its way into our water
courses; no forest benefit is more important
when we consider that 3.3 pounds of soil are
lost for each pound of grain produced in Illinois
(Iverson et al. 1989). Global warming, due
largely to the excessive buildup of carbon
dioxide in the atmosphere, is also counteracted
to some degree by our forests because plants
convert tremendous quantities of carbon
dioxide into plant tissue and oxygen each day.
Then too, our forests contribute greatly to the
maintenance of biological diversity, a benefit of
crucial importance in Illinois where the land-
scape is dominated by a row-crop monoculture.
The purpose of this paper is to review the
historic trends that shaped the Illinois forest, to
document its present status, and to summarize

Cropland 24.7

Forestland 4.3

L— | Urban 2.6
yy Pasture 2.4
Other 1.2

Nonforest with trees 0.9

Figure |. Major land use in Illinois in millions of
acres, 1985. Total acres in Illinois = 36,061,000.
Source: Hahn 1987.

the benefits it currently provides. The material
is largely condensed from a more detailed and
complete document, Forest Resources of
Illinois: An Atlas and Analysis of Spatial and
Temporal Trends (Iverson et al. 1989). Readers
are encouraged to consult that book and the
map (Iverson and Joselyn 1990) that accompa-
nies it for a great deal more information
regarding the forests of Illinois, including data
specific to the counties in which they may be
particularly interested. Both the book and map
are available as Special Publication 11 from the
Illinois Natural History Survey.

Much of the story of the Illinois forests
can be understood by comparing the earliest
systematic vegetation data available for the
state, data recovered from the original land
surveys made during the first half of the
nineteenth century, with recent land-use infor-
mation taken via remote sensing from airplanes
and satellites.

FORESTS OF 1820

Illinois was surveyed by the United States
General Land Office between 1807 and 1844.
Starting from southern Illinois and working
northward, surveyors divided the land into
townships and sections, prepared plat maps,
and made notes on the vegetation they encoun-
tered. These records provide a fairly complete
picture of the landscape prior to the massive
disturbance caused by European settlement.
Anderson (1970) published a map showing the
statewide distribution of forest and prairie as
deduced from these data (Figure 2). Large
expanses of forest existed, primarily in the
south and west. Approximately 38.2% of the
state (13.8 million acres) was forested at the
time of the European settlement, 61.2% was
prairie, and 0.6% was water. Fifteen counties
were at least 80% forested, and only 21
counties had less than 20% forest cover.

361

362 Illinois Natural History Survey Bulletin

FOREST TRENDS 1820-1980

Illinois forests have undergone drastic changes
in the decades since European settlement. Only
31% of the forest area present in 1820 exists
today (Figure 3). The lowest percentage of
forest occurred about 1920 when only 22% of
the land forested in 1820 remained in forest
(Telford 1926; U.S. Forest Service 1949; Essex
and Gansner 1965; Hahn 1987). Although
forest area has increased in recent decades,
most of today’s forest is secondary forest, and
only about 11,600 acres exist in a relatively
undisturbed condition (Illinois Natural Areas
Inventory as reported in Iverson et al. 1989).
Illinois ranks 49th, next to Iowa, in percent of
the state converted from its “potential” vegeta-
tion type (Kiichler 1964); only 11 percent of
the state remains in its “potential” vegetation
type and essentially all of that is forest
(Klopatek et al. 1979).

Figure 2. Forests in Illinois about 1820. Source:
Anderson 1970.

Vol. 34 Ant. 4

The pattern of deforestation of the
primary (i.e., “virgin’’) forests of Illinois can be
deduced to some degree by relying on estimates
of forestland in 1820 and 1924 and on other
written accounts (especially Telford 1926).
From initial settlement in the early 1800s to
1860, agriculture was the only important
industry associated with wooded lands. Until
1830, forests were the sole source of potential
agricultural land; however, when settlers
realized that the prairies made good cropland
and after the invention of the moldboard plow,
the prairies were converted to cropland at an
astonishing rate of approximately 3.3% per
year (Table 1). Over 300,000 people settled the
prairies during the decade of the 1830s, and this
burgeoning population created an enormous
demand for housing material, fuel, and fence
posts. Railways were not yet in place to import
lumber, and most of the timber in the prairie
counties rapidly disappeared.

Figure 3. Forests in Illinois about 1980. Source:
U.S. Geological Survey land-use data, 1973-1981.

April 1991

By 1860, a timber industry had begun to
flourish in Illinois. Ninety-two of the 102
counties had industries based on wood products
by 1870, and forestland had dwindled to 6.02
million acres (Telford 1926). During the 1880s,
annual lumber production exceeded 350 million
board feet, 2.2 times the present production,
and continued to increase until 1900, when it
began to decrease as the resource itself de-
clined. By 1923, only 22,000 acres of the
original 13.8 million acres of primary forest
remained.

A useful comparison can be made
between deforestation in Illinois in the nine-
teenth century and the deforestation presently
under way in the tropics. The primary forests of
Illinois went from 13.8 million acres in about
1820 to 6 million acres in about 1870, to
22,000 acres in about 1920 (Figure 4), an
overall deforestation rate of 1% per year
(1.13% of the original primary forest lost
during the first half of the century, 0.87%
during the second half). Deforestation rates,
however, were not a constant during the period
and probably followed a curve such as that
shown in Figure 5, with maximum deforesta-
tion in the late 1800s. Rates of deforestation
have also been compiled for Rond6nia in Brazil
(Malingreau and Tucker 1988), for Costa Rica
(Sader and Joyce 1988), and for Malaysia
(Iverson et al. 1990) and are shown in Table 1.
The fastest rate, 2.47% annually, was found
from 1972 to 1982 in peninsular Malaysia,
even though more forestland was being
removed in Rondonia. This rate was probably
equaled in Illinois in the late 1800s (Figure 5).
A similar curve is currently found in the other
countries, with Malaysia at the apex of the

Symposium Proceedings: Our Living Heritage

363

curve, ROndonia on the upward slope with
increasing rates, and Costa Rica on the down-
ward slope with a declining resource and a
dropping rate. History does indeed repeat itself,
and we Americans should acknowledge our
own history of deforestation as we now attempt
to curb the destruction of tropical forests.

FOREST TRENDS 1962-1985

Forest area increased by 10% from 1962
through 1985, from 3.87 to 4.26 million acres.
This increase is partially explained by the
reduced number of cattle raised in Illinois and
the conversion of pastures and hayland to
secondary forest. Total net volume of growing
stock has also increased 40% since 1962 (Table
2). Pine plantations have shown the highest
percentage of increase in volume (up to 375%),
but the largest absolute increase in volume was
shown by oaks (an increase of 0.64 million
cubic feet).

Acres

10,000,000 |-

1,000,000 |;

100,000 };

10,000
1820 1870 1924 1948 1985
Figure 4. Extent of Illinois primary forests, 1820—
1985. Interpreted from Telford 1926; U.S. Forest
Service 1949; and Anderson 1970.

1962

Table 1. Recent rates of land clearing in three tropical countries compared with rates of land clearing in

Illinois from 1820 to 1923.

Sq km of land Percent cleared per year

Location Land use Year
Rondonia, Brazil Forest 1978
1987
Malaysia Forest 1972
1982
Costa Rica Forest 1940
1983
Illinois Forest 1820
1870
1923
Illinois Prairie 1830

1860

239,800
208,000 1.47

48,970
36,870 2.47

34,210
8,710 ns

55,870
24,290 1.13
90 0.87

87,550
10 3.33

364

Compositional changes during
1962-1985 were especially profound, with vast
percentage increases in commercial acreage of
white, red, and jack pines, oak—gum-—cypress,
and especially maple—beech forest types
(Figure 6). Maples increased 41-fold in the past
25 years—from 0.025 million acres to 1.046
million acres! Concomitantly, oak—hickory
decreased by 337,000 acres (14%), and over
half of the state’s elm—ash—soft maple dis-
appeared. The loss of oak—hickory is largely
from maple “take-over” as shade-tolerant
maples replace oak—hickory stands following
mortality or harvest. A documented case of the
maple take-over of a forest in east-central
Illinois is presented later in these proceedings
(Ebinger and McClain, page 375) and else-
where (Ebinger 1986). The reduction of
elm—ash-soft maple is due to mortality from
Dutch elm disease and the conversion to
cropland of bottomland forests that once
supported this forest type. These data make
clear that although forest acreage and volume
have increased since 1962, the quality and
value of the timber resource has diminished, at
least by today’s standards. Maple-dominated
forests also support a somewhat different array
of wildlife than that supported by oak-domi-
nated forests, and such “hard mast” (acorns and
hickory nuts) feeders as squirrels and wood-
peckers are less abundant in maple-dominated
forests.

ILLINOIS FORESTS TODAY

A closer look at the current status of the Illinois
forests reveals some interesting and on occa-
sion surprising information.

Area

Estimates of current forestland compiled from
the 1985 U.S. Forest Service inventory indicate
that about 12% (4.27 million acres) of the land
area of Illinois is forested (Hahn 1987). The
extent of this forestland can be seen in Figure 3
(as well as in several forms on the 1:500,000
scale map of Iverson and Joselyn 1990). The
importance of the southern and western
counties is clear. At one extreme is Ford
County with only 3,000 acres of forestland; at
the other is Pope County with 149,200 acres,
Jackson with 134,500, and Pike with 122,500.
Included in this 4.27 million acres are
4,029,900 acres of commercial (capable of and
potentially available to produce commercially

Illinois Natural History Survey Bulletin

Vol. 34 Art. 4

valuable trees) forestland and 235,600 acres of
reserved or protected timberland.

Wooded strips less than 120 feet wide
and land on which at least one tree (5 inches in
diameter at breast height) occurs per acre make
up a category that has been designated “non-
forestland with trees.” Included in this category
are wooded strips (178,500 acres), wooded
pastures (162,400), urban and other built-up
land (139,500), windbreaks (133,100), im-
proved pastureland with trees (103,600), urban
forest (102,800), and several miscellaneous
classes. Taken together, 900,800 acres of
nonforestland with trees are found in Illinois.

Composition

The composition of many Illinois forests has
changed over the past several decades. Today,
about one-half of the commercial forest acreage

Percent cleared per year
2S

— rate of forest loss
— average annual rate

i)

1820 1840 1860 1880 1900

Figure 5. Rate of forest clearing in Illinois, 1840—
1920. Interpreted from Telford 1926; U.S. Forest
Service 1949; and Anderson 1970.

1920

Acres (x 1000)
2,500

2,000

1,500

1,000

P WRIJP SP
+3900% +23% +4% -14% +674% -53%+4120%
Figure 6. Composition of Illinois commercial
forests, 1962-1985. Percent change is given below
each pair of bars. Abbreviations are decoded as
follows: WRJP = white-red—jack pine, SP =
shortleaf pine, OP = oak—pine, OH = oak-hickory,
OGC = oak—gum-cypress, EASM = elm—ash-soft
maple, MB = maple—beech. Source: Hahn 1987.

OP OH OGC EASM MB

April 1991

Symposium Proceedings: Our Living Heritage

Table 2. Net volume of growing stock on commercial forestland in Illinois by species group for 1962 and
1985, percent change between those dates, and net annual growth estimated from 1985 data.

1962 1985 Net annual growth
Species group (thousand cubic feet) Percent change —_ (thousand cubic feet)
Softwoods
Loblolly—shortleaf pine 15,200 64,700 +327 1,891
White pine! _ 16,800 _ 393
Red pine! -- 12,000 = 310
Eastern red cedar 2,400 11,400 +375 445
Bald cypress 6,800 8,900 +31 13
Jack pine! —_ 700 36
Other softwoods 700 3,000 +329 110
Total 25,100 117,500 +368 3,224
Hardwoods
Red oak 701,800 1,062,400 +51 18,352
White oak 739,700 1,017,600 +38 15,075
Hickory 343,900 522,500 +52 7,443
Soft maple 259,200 341,600 +32 14,144
Elm 367,700 267,400 -27 -5,106
Green—white—black ash 218,200 261,000 +20 6,932
Hard maple 99,800 163,100 +63 3,717
Cottonwood 114,100 157,800 +38 1,976
Sycamore 123,300 134,600 +9 2,412
Black walnut 77,500 119,100 +54 2,279
Hackberry — 93,500 — 5,683
Black cherry? —_ 87,700 3,663
Basswood 25,800 54,100 +110 1,215
Yellow poplar 26,400 51,800 +96 1,609
Willow? o= 50,300 a 1,427
Sweetgum 58,600 45,100 -23 1,163
River birch? a 36,800 — 1,257
Tupelo 13,900 28,000 +101 209
Beech 14,500 12,100 -17 242
Butternut? 5,700 = 105
Aspen 9,100 1,900 -79 28
Other hardwoods 223,100 203,500 9 8,966
Total 3,416,600 4,717,600 +38 92,791
Total all species 3,441,700 4,835,100 +40 96,015

365

'Tabulated only in 1985 survey, included with other softwoods in 1962.
> Tabulated only in 1985 survey, included with other hardwoods in 1962.

Source: Hahn 1987; reprinted from Iverson et al. 1989.

(2.03 million acres) is oak—hickory, one-fourth
is maple—beech (1.05 million acres, almost
exclusively sugar maple), and one-sixth is
elm—ash-soft maple (0.72 million acres)
(Figure 6). Together, the remaining forest types
(white—red—jack pine, loblolly—shortleaf pine,
oak—pine, and oak—gum-—cypress) account for
an additional 216,800 acres of commercial
forestland.

The location of these various forest types
has been mapped (Iverson et al. 1989; Iverson
and Joselyn 1990). Oak—hickory is found
throughout the state with maximum levels in
the western and southern counties. Maple—

beech, a forest type also found throughout
Illinois, has the highest average number of
acres per county in western Illinois but is
proportionally most prominent in the central
Grand Prairie counties. Elm—ash-—soft maple is
found in bottomland forests, and these forests
are more frequently located in the southern
counties. Oak—pine, oak—gum-cypress, and
shortleaf pine types are confined to the south-
ern counties, but the white pine type is most
common in the western part of the state.
According to the Illinois Plant Informa-
tion Network (Iverson and Ketzner 1988), 508
woody taxa have been recorded in Illinois, a

366 Illinois Natural History Survey Bulletin

high diversity of woody plant species consider-
ing the extensive agricultural acreage. Trees
account for 261 taxa, shrubs 284, and lianas 47
(some taxa include more than one type). These
woody plants account for a diversity of cover
types and occupy a variety of habitats. On
average, 70 tree taxa and 54 shrub taxa have
been recorded from each county (Iverson et al.
1989). Southern counties have the largest
number of tree taxa (Jackson has 145 taxa,
Pope 129, and Union 128), and northeastern
counties have the most shrub taxa (Cook has
153 and Lake 136).

Volume, Annual Growth, and Number
Net volume estimates for 1985 showed the
prominence of oak and hickory in commercial
forests, with considerable amounts of ash,
black walnut, cottonwood, elm, maple, and
sycamore as well (Figure 7). The data shown in
Figure 7 may have greater immediacy if we
consider that 1 million board feet provide
enough lumber to build an estimated 73 wood
houses. The total net volume of Illinois timber
in 1985—17.5 billion board feet—would
theoretically build 1.3 million wood houses!

Total net volume estimates of growing
stock were 4.8 billion cubic feet, an average of
47.4 million cubic feet per county or 1,200
cubic feet per acre of commercial forestland in
the state. Hard hardwoods (predominately oak,
hickory, and ash) accounted for 68% of total
volume; soft hardwoods (e.g., elm and soft
maple) accounted for 30% and softwoods (e.g.,
pine) made up 2%.

According to annual growth estimates for
1985 (Hahn 1987), growing stock showed 96
million cubic feet of growth, or 437 million

Other Hardwoods 1,815
Maple 1,766

L— ie 959
WG Ash 783
Cottonwood 710
P Sycamore 605

\ \ Elm 483

\ Walnut 368
Softwoods 338

Figure 7. Total volume of Illinois commercial

forestland in 1985 in million board feet. Total net

volume of sawtimber was 17.5 billion board feet.

Source: Hahn 1987.

Oak 8,833

Vol. 34 Art. 4

board feet of sawtimber growth. Over 42% of
net annual sawtimber growth was accounted for
by oaks, with another 10% from soft maple,
6.3% from ashes, 3.7% from black cherry,
3.3% from hard maple, and 3.2% from black
walnut. Only elm and black ash showed
negative growth rates between 1962 and 1985,
and these are attributed to Dutch elm disease
and the clearing of bottomlands.

The estimated number of trees in Illinois
commercial forests revealed a somewhat
surprising statistic: the elms, with 344 million
trees, were the most common group. Most of
these, however, are small slippery (or red) elms
with little commercial value (Figure 8).
Overall, white oaks (99 million), red oaks (136
million), hickories (185 million), hard maples
(117 million), and soft maples (91 million)
were very abundant.

Age

Illinois forests are reasonably well distributed
among age classes, with 61-year to 80-year
classes most prevalent; however, certain trends
appear when the ages of major forest types are
considered (Figure 9). Oak—hickory forests
show a very uneven age distribution, with the
majority older than 60 years. A predominance
of maple—beech is found in younger age classes
(<30 years) relative to oak—hickory and
elm—ash-soft maple. This pattern again
illustrates, as it did in the data on acreage
trends (Figure 6), two important aspects of
Illinois forests today: maples are rapidly
increasing in younger age Classes and forest
types dominated by oaks and elms are declining
and have relatively fewer trees in younger age
classes. Among the other forest types, white

Oak 236
Noncommercial 217

Elm 344 Maple 208

\ee 185
Ash 114

Other Hardwoods \ Walnut 66

508 Softwoods 49

Figure 8. Number of live trees in 1985 in Illinois
commercial forestland in millions of trees. Total
number of trees was 1.93 billion. Source: Hahn 1987.

April 1991

and shortleaf—loblolly pine peak in the 21- to
30-year class with very little stand acreage
under 10 years of age. Pine plantations are no
longer being planted to the extent they were
from 1930 to 1960, primarily because of
changes in the management of the Shawnee
National Forest (U.S. Forest Service 1986).

Site

Forest stands can also be classified according to
an index that measures the quality of a site
based on the height its trees attain after 5O
years of growth. The soils of Illinois are
superior for forest growth compared to the
relatively shallow or infertile soils of neighbor-
ing states like Missouri or Kentucky. According
to this index, fully 84% of the trees in the
commercial forestlands of Illinois are capable
of supporting growth of 61 to more than 100
feet during a 50-year interval.

Mortality

In 1985, the forests of Illinois experienced an
annual mortality of over 200 million board feet
of sawtimber (67 million cubic feet of growing
stock) (Hahn 1987). In contrast, 161 million
board feet of timber were cut in 1983 (Blyth et
al. 1987); at that time, therefore, more timber

Acres (xX 1000)

Symposium Proceedings: Our Living Heritage 367

was dying than was being cut. These mortality
data represent an annual death rate of 1.36% of
the total inventory and 69% of the annual
growth of growing stock. These rates are quite
high in comparison to the mortality rate (0.9%)
in Illinois in 1962 and to rates in neighboring
states—central Wisconsin, for example, had an
average mortality rate of only 0.8% of its total
inventory in 1983 (Raile and Leatherberry
1988). The Illinois secondary forests are aging,
with concomitant increasing mortality. Disease
accounted for 38% of the mortality, but
weather, suppression, and unknown causes
were also important (Hahn 1987). Elms
suffered the greatest mortality and accounted
for 26% of total mortality; 56% of the elm
mortality was due to disease.

Ownership

Over 90% (3.64 million acres) of the commer-
cial forests in Illinois are privately owned,
mostly by farmers (45.3%) and other individu-
als (38.1%) (Figure 10). The remaining 10% is
publicly owned, primarily by the federal
government (7.2%) in the form of the Shawnee
National Forest. The Cooperative Extension
Service of the U.S. Department of Agriculture
estimated that Illinois had 169,073 private

600

(_] Other
C) Elm-Ash-Soft Maple _
fa Maple—Beech

a Oak—Hickory

500-7 <->:

400|------

300

200

100

0

5 15 25 35 45 35)

65 13 85 95 110 130 150

Figure 9. Acreage by age classes (in years) of the three major forest types in Illinois in 1985. Source: Hahn

1987.

forestland owners, each of whom owned an
average of 21.5 acres of forest. The primary
reasons for forest ownership given by the
holders of small parcels were wildlife habitat
and aesthetic value (Young et al. 1984); income
was of greater importance for those who owned
large forest parcels (McCurdy and Mercker
1986).

BENEFITS OF ILLINOIS FORESTS

Although Illinoisans would undoubtedly
respond in different ways if queried on the
benefits of the forests of our state, probably
none of them would be in error. The forests of
Illinois truly offer multiple benefits and
perhaps one of the most encouraging aspects of
management is that plans can be designed to
accommodate and enhance these varied
benefits.

Natural Communities

In the late 1970s, a search for natural communi-
ties relatively undisturbed by human activity
was undertaken throughout the state (White
1978). Of the 1,089 natural areas selected for
inclusion in the Natural Areas Inventory, 392
(36%) contained forestland; however, only 149
natural areas, a mere 11,593 acres of forestland,
were classified as Grade A (relatively undis-
turbed) or Grade B (some disturbance). Of that
total, about a third was classified as Grade A.
Since that inventory, a few additional high-
quality sites have been added, for a total of 157
areas from 62 counties. Lake and St. Clair
counties contain the largest number of forested
natural areas (12 and 11, respectively); Peoria
has 7, Washington and Mason 6 each, and
Massac 5. Adams County has the most exten-
sive acreage of high-quality forestland, 1,950
acres, followed by St. Clair (963 acres), Lake

Government 9.6%
(federal 7.2%,

state 1.4%, local 1.0%)
Corporate
ownership

6.8%

Individuals
83.4%
(farmers 45.3%, nonfarmers 38.1%)

Figure 10. Ownership of Illinois commercial forests,
1985. Source: Hahn 1987.

Illinois Natural History Survey Bulletin

Vol. 34 Ar. 4

(635 acres), Johnson (622 acres), McLean (450
acres), Saline (447 acres), Cook (444 acres),
and Pike (431 acres).

Many high-quality forests in Illinois are
undergoing degradation because of the invasion
of exotic plants. Over much of the state, forests
are threatened by garlic mustard (Alliaria
petiolata), Amur honeysuckle (Lonicera
maackii), tatarian honeysuckle (L. tatarica),
Japanese honeysuckle (L. japonicus), multiflora
rose (Rosa multiflora), autumn olive (Elaeag-
nus umbellata), and other introduced species.
These exotics reduce the diversity of forest
communities by eliminating native understory
species. Management strategies must be
adopted within the few remaining high-quality
forests if they are to be protected from aggres-
sive species. Control measures include recruit-
ing volunteers for hand weeding, the cautious
application of pesticides, and the implementa-
tion of biological controls. Perhaps most
important is an educational program to teach
the public how to identify and control these
dangerous invaders.

Botanical Diversity

Illinois forests provide habitat for an excep-
tional diversity of plant species and are the
natural home for most trees and other woody
species. The 508 taxa of trees, shrubs, and
lianas found in Illinois represent 15.9% of the
state’s reported flora, and 346 (69%) of them
are associated with forest habitats (ILPIN data:
Iverson and Ketzner 1988) (Figure 11). Most of
the remaining taxa are cultural (escaped from
cultivation). Of the 508 taxa, 370 (73%) are
native to Illinois; the remaining are introduced.
A relatively high proportion of the state's
woody taxa are listed as rare in Illinois (40%):
15% occur commonly, 33% occur occasionally
(common in localized patches), and 12% are

Forest, nonwoody taxa

1,235 (38%)

Forest, woody taxa

Nonforest. 346 (11%)

nonwoody taxa

1.461 (46%) Nonforest. woody taxa

162 (5%)
Figure 11. Number of plant taxa by habitat and habit
(woody and nonwoody). Total taxa in Illinois =

3,204. Source: Iverson and Ketzner 1988.

April 1991

uncommon (localized distribution or sparse
throughout).

Illinois forests also provide habitat for an
amazing number of nonwoody taxa. Including
the woody taxa, fully 1,414 native taxa (61% of
the native Illinois flora) are associated with
forest habitats (Figure 11). Thus Illinois forests,
which occupy only 12% of the area of the state,
provide habitat for over half of its native flora.
If we are to protect this irreplaceable biological
diversity, we must maintain and restore forest
communities. Beyond the importance of
forestland as habitat for total plant diversity,
rare plant species are frequently found in forest
habitat, for example, 166 taxa (47%) of the 356
plants listed as threatened or endangered in
Illinois are forest inhabitants. The importance
of high-quality forests as refuges for these taxa
cannot be overemphasized, especially in the
face of extreme pressures from urban and
agricultural growth.

Wildlife Habitat

Illinois forests provide the major habitat for
numerous wildlife species, and losses in the
quality and quantity of that habitat severely
affect wildlife populations (Illinois Wildlife
Habitat Commission 1985). Game species—
gray squirrel, eastern wild turkey, quail, and
white-tailed deer—depend on woodlands as do
many more nongame animals—thrushes,
warblers, woodpeckers, nuthatches, kinglets,
and whippoorwills—to mention only a few
bird species. But some relationships between
wildlife and forests are more subtle. Most of us
recognize the dependence of wood ducks on
natural cavities in the trees of bottomland
forests, but bottomland forests also provide
food and habitat for fish, mitigate the effects of
floods, restrain the movement of harmful
chemicals into lakes and streams, and provide
shade, thereby lowering water temperatures
during stressful summer months.

One method of summarizing the value of
Illinois wildlife habitat is based on land use.
Complete details are presented in Graber and
Graber (1976), and revised calculations based
on current data are given in Iverson et al.
(1989). The habitat evaluation index devised by
Graber and Graber is based on the relative
amount of a particular habitat type within a
given area, the availability of that habitat type
within the state or region, the changing
availability of that habitat (Is it increasing or

Symposium Proceedings: Our Living Heritage

369

decreasing over time?), and the “cost” of a
given habitat measured in years required to
replace the ecosystem. A summary of habitat
factors for Illinois as a whole is presented in
Table 3. By this calculation, over three-quarters
of the wildlife habitat (88 of 115.7 habitat
factor points) is derived from forests. Elm—
ash—cottonwood rates highest because this
forest type has been disappearing so quickly
over the past two decades (Figure 6). Oak—
hickory values would be higher except that
numbers in older age classes are increasing as
secondary forests mature, even though numbers
in younger age classes are decreasing (Figure
9). A very minor rating was earned by
maple—beech because this forest type has
increased so dramatically in recent years
(Figure 6).

This method can be used to evaluate
wildlife habitat on parcels of various size (see
examples in Iverson et al. 1989). In the final
calculation, the habitat factor for a given site or
region is divided by a regional or statewide
habitat factor (115.7 for the state). An index of
1.0, therefore, means that the value of the
habitat under consideration is about average for
the state or region as a whole. Thus, a habitat
evaluation index of 1.5, the value calculated for
the 16 southern counties, indicates a much
higher wildlife value than the value of the state
overall. Similarly, the value of 0.66 for the 60
northern counties indicates a relatively poor

Table 3. Habitat factors for Illinois, 1985, calculated
according to Graber and Graber (1976).

Percent of

Land type Habitat factor habitat factor
Forest
Pine 5.70 4.9
Oak-hickory 30.07 26.0
Oak—gum-cypress 11.97 10.3
Elm—ash—cottonwood 40.19 34.7
Maple—beech 0.14 BeOS
Subtotal 76.0
Nonforest
Cropland 0.29 0.3
Pasture/hayland 10.01 8.7
Prairie 1.46 1.3
Marsh 15.28 13.2
Water 0.38 0.3
Urban, residential 0.03 0.0
Fallow 0.19 0.2
Subtotal 24.0
Total 115.73 100.0

370 Illinois Natural History Survey Bulletin

habitat for wildlife, and the value of 1.09 for
the 26 south-central counties indicates wildlife
habitat somewhat above that of the state as a
whole.

Fragmentation of forest habitat has
negative implications for wildlife, especially
for neotropical migrant birds that need large
blocks of uninterrupted forest for successful
nesting (Harris 1984; Blake and Karr 1987;
Robinson 1988). As large tracts of forest are
broken into small, isolated woodlots, more
forest edge is created and more opportunities
exist for edge-adapted species, most impor-
tantly the cowbird, to invade the area and
parasitize the nests of many forest songbirds.

The extent of fragmentation in Illinois
forests was made clear in a recent examination
of forest parcels by size. Relying on the Illinois
Geographic Information System and data from
the U.S. Geological Survey, researchers
determined that 10,121 forested parcels exist in
the state and that the average size per parcel is
358 acres (Iverson et al. 1989). About 44% of
the parcels are less than 100 acres in size and
about 10% are larger than 600 acres (Figure
12). Perhaps the density of forest parcels can be
pictured more clearly if we envision an area the
size of a township—36 square miles. On
average, 6.1 parcels exist per township-sized
area, with 69% of them roughly 40 (limit of
resolution of the data) to 200 acres in size. This
perspective makes clear that Illinois forests are
extremely fragmented and that a concentrated
effort must be made to protect larger forest
patches and to aggregate smaller ones.

100—200 acres:
2,476 (1.5)

201—600 acres:
2.099 (1.3)

< 100 acres:

4,479 (2.7) 601—1,100 acres:

525 (0.3)
> 1,100 acres: 542 (0.3)

Figure 12. Number of forested parcels in Illinois by
size and average number of parcels per township
equivalent (36 square miles). Total number of
parcels in Illinois of a given size is the number
immediately following the size (e.g., <100-acre
parcel: 4,479). Average number of parcels of a given
size per township equivalent is given in parentheses.
Source: Iverson et al. 1989.

Vol. 34 Art. 4

Soil and Water Quality Protection

Soil erosion with its accompanying degradation
of surface water is indeed a serious threat to the
future of an agricultural state: for every pound
of corn, soybeans, wheat, or oats grown in
Illinois, 3.3 pounds of soil are lost (Iverson et
al. 1989). In contrast to cropland, forest
vegetation protects against excessive soil loss.
Average erosion of cropland proceeds at about
four times the annual rate of nongrazed
forestland—7 tons per acre compared to 1.6
tons, respectively. The difference in soil loss is
even greater on sloping, highly erodible soils.
Soils with land capability ratings of [Ve to VIle
lose 24.2 to 39.4 more tons per acre each year
they are under cultivation than they would lose
if they were forested. In 1982, 1.75 million
acres of cropland had these capability ratings.
Had those acres been converted to nongrazed
forestland, 36.5 million of the 157.8 million
tons of soil lost annually from cropland would
have been saved. Figure 13 shows that the soil
savings that would result from converting
cropland with higher capability ratings to
nongrazed forest would be disproportionately
higher than conversions from cropland with
lower ratings.

The Conservation Reserve Program is
designed to remove marginal cropland from
cultivation, and it is helping; however, over
96% of the cropland currently being removed
from production in Illinois is going into grass
rather than trees. The U.S. Department of
Agriculture and the Illinois Council on Forestry
Development are working together to alter this
percentage in favor of trees.

Acres (x 1000) Tons lost (x 1000)

16° r 70
147 Sr = : - = 5 60
12° G Cropland —_

= 7 50

— Soil loss

Il Le ey’
Capability Class

Figure 13. Cropland acreage and annual soil loss by
capability class. Class I soils are most productive:
Class VII soils are least productive. Source: U.S.
Soil Conservation Service data base 1982.

April 1991

Heavy grazing, and especially feedlot
operations, in forestlands largely negates the
benefits of soil protection. Average soil loss
from forestland that is heavily grazed or under
feedlot operations is 13.1 tons per acre per year
in contrast to only 1.6 tons per acre per year on
nongrazed forest. Thus, 66% of the 12.6
million tons of soil lost annually from forest-
land is lost from these areas, even though only
19% of Illinois forests are categorized as
grazed. Light grazing of forestland generally
does not increase soil loss significantly and is
certainly to be preferred over cultivation of
marginal lands.

According to estimates by the U.S. Forest
Service, 133,100 acres of windbreaks existed in
Illinois in 1985 (Hahn 1987). Windbreaks
retard soil loss due to wind erosion, but they
also provide shade for livestock and shelter for
wildlife. Their aesthetic qualities are not to be

overlooked, but their role in the conservation of

energy is growing in importance. Back in 1981,
the Soil Conservation Service estimated that
124,000 buildings in rural Illinois needed
windbreaks. Had they been planted, energy
equivalent to 941 million kilowatt-hours of
electricity could have been saved (USDA Soil
Conservation Service 1982).

Recreation and Scenic Values
In 1987, surveys by the Illinois Department of
Conservation indicated that Illinoisans spent
about 240 million days or portions of days
pursuing recreation on or near forestlands; in
the process they spent approximately $6.3
billion (Illinois Department of Conservation
1989). Activities closely aligned with forest
recreation (picnicking, observing nature, cross-
country skiing, backpacking, hiking, camping,
canoeing, horseback riding, snowmobiling,
riding off-road vehicles, trapping, and hunting)
accounted for 206 million of those days, an
average of 18.7 days per resident (Figure 14).
The majority (93%) of the 4,528 areas
developed for recreation in Illinois (almost
900,000 acres) are publicly owned and oper-
ated. Total land available for recreation totals
roughly 2.7% of the state’s land and water area,
a per capita outdoor recreation acreage of less
than 0.1 acre. Among states, Illinois rates 46th
in total public open space per capita. In
addition, most of the publicly owned land
available for recreation is located in the south-
ern part of the state; the majority of IIlinoisans,
however, live in the north.

Symposium Proceedings: Our Living Heritage 371

Urban Forests

Most Illinoisans (83%) live in urban centers,
and urban forests are often their only exposure
to a natural environment. Urban forests provide
many benefits beyond those normally associ-
ated with rural forests, including temperature
modification and energy conservation; the
abatement of air, water, and noise pollution; the
masking of unpleasing urban views; and
physical and psychological benefits to city
dwellers. Because the urban forest exists in
such a heterogeneous environment, an accurate
assessment of its extent and function is
difficult. The U.S. Forest Service, however,
has estimated that 102,800 acres of urban forest
and 139,500 acres of urban areas with trees
existed in Illinois in 1985 (Hahn 1987). Cook
County alone has over 67,000 acres of forest
preserves, and much of this land is available for
recreation. A recent remote-sensing study
revealed that 21.3% of the land area in the six-
county Chicago area had tree cover in 1988
(Cook and Iverson 1991). Yet less than 0.01
acre per capita of publicly owned forestland
exists in that six-county area, and Chicago
ranks last among the nation’s ten largest urban
centers in this regard.

Urban forests face three problems. First,
maintenance and management are inadequate.
A recent survey by the Illinois Council on
Forestry Development (1988) estimated that
6.5 million municipal street trees exist in
I]linois with an estimated value of $3 billion.
These trees are generally not adequately
maintained because of inadequate budgets and
the lack of trained foresters. In addition, less
than half the potential number of street trees are
presently in place, and removals outstrip
plantings (American Forestry Association
1988). Second, forestlands are jeopardized by

ORV: 29,437

Picnicking: 28,307
[— ae 16,552

a riding:
Ny: 3,388

Camping: 12,961
Observing nature: Hunting: 10,166
87,449 Hiking: 7,899

Figure 14. Days (in thousands) spent in recreational
pursuits on or near forestlands in Illinois, 1987.
Source: Illinois Department of Conservation 1989.

372 Illinois Natural History Survey Bulletin

development and population pressures.
Tremendous growth is now occurring in the six
collar counties around Chicago. Information
from the Northeastern Illinois Planning
Commission (1987) shows that 867 quarter
sections (about 5.6% of the area) were urban-
ized (population density exceeding 1,000 per
square mile) between 1970 and 1980. Much of
this growth was at the expense of forestland.

A third problem is the absence of a policy for
using wood waste. Until recently, much of the
debris from tree removals and large amounts of
other wood wastes were deposited in landfills,
an enormous waste of wood and leaf mulch and
the needless use of costly landfill space. Better
uses for this material must be developed and
marketed.

Timber Products

Illinois ranks fifth in the nation in demand for
wood but 32nd in production. As a result,
Illinois imports much of the wood it uses from
neighboring states. In addition, 14.2% of the
wood harvested in Illinois is processed in
neighboring states and then often imported
back into the state. Currently, the annual
growth of timber (96 million cubic feet)
exceeds timber removals (68.6 million cubic
feet removed for timber products, logging
residues, and changing land uses), and a higher
proportion of the state’s demand for wood
could be met within its own boundaries if the
processing facilities were at hand. With
judicious management, harvesting could be
increased, negative effects on the environment
minimized, and multiple benefits achieved.

In 1983, 161 million board feet of timber
(mbf) were harvested in Illinois (Blyth et al.
1987); 146 mbf were processed in 178 Illinois
sawmills. Red oak (29%), pin oak (19%), white
oak (16%), and cottonwood (10%) accounted
for the majority of sawlogs processed in the
state. Of the 4 mbf of veneer and other high-
quality logs (mostly white oak, walnut, and red
oak) cut in Illinois during 1983, only 0.3%
remained in the state. Additionally, all pulp-
wood (7.2 million cubic feet) produced in the
state were processed elsewhere. The veneer and
pulpwood statistics are not surprising because
virtually no plants for either veneer or pulp-
wood are found in Illinois.

Vol. 34 Art. 4

An enormous quantity of fuelwood is
harvested from Illinois woodlands. In 1982,
nearly 2 million cords of firewood were cut or
gathered, a figure that represents 43% of the
total trees utilized that year! The major harvest
of fuelwood takes place in the heavily popu-
lated northeastern counties. Cook, McHenry,
and Will counties, for example, each harvested
over 150,000 cords of fuelwood in 1983 (Blyth
et al. 1985). The majority of firewood (97%)
was Cut from private lands, and 75% was
gleaned from dead trees.

According to U.S. Department of
Commerce figures, forest-related industries in
Illinois employ 55,000 people with an average
payroll of $965 million. These firms contribute
more than $2 billion annually to the state’s
economy through value added by manufacture;
in addition, they invest more than $144 million
in capital improvements annually (U.S.
Department of Commerce 1982-1985).

According to 1984 data from Dun &
Bradstreet, 166,900 employees work for 957
Illinois firms that are primarily involved in the
manufacture of wood products. If the paper
industry is included, an additional 576 firms
and 367,450 persons are involved (Figure 15).
The Dun & Bradstreet numbers are much
higher than those released by the U.S. Depart-
ment of Commerce because Dun & Bradstreet
include the total number of employees, even
those not directly associated with the wood-
manufacturing component. Nonetheless, a large
number of employees work in forest-related
industries, most of which are located in the
Chicago region.

Millwork/Plywood 376

Miscellaneous 199

Furniture 161

Paper 576

Containers 101

Sawmills 89
Buildings 31

Figure 15. Forest-related industries in Illinois, 1984.

These 1,533 sites employed 534,342 workers.

Source: Dun & Bradstreet data base 1984.

April 1991

CONCLUSIONS

A great deal of information has been presented
to establish the initial contention of this paper:
the Illinois forests provide numerous important
benefits to the citizens of the state. Neverthe-
less, considerable improvement in the quantity
and quality of these benefits could be achieved
if forestlands were better managed. Over most
of the state, little forest management is under-
way, and the potential of our forests to provide
wildlife habitat, preserve biodiversity, and
extend wood production has not been tapped.
Even in “wilderness” areas, management is
often necessary to maintain the status quo (e.g.,
remove exotic invaders). Ecosystems are not
static entities; change is inevitable, but only
with management can change benefit the
resource as well at its human guardians.

We need to manage the forest resources
we currently possess, but we also need to plant
more forests if we are to assure continuing
benefits from our forests. Recent political
developments have and may continue to
support tree planting programs; however,
caution is in order. Planting trees requires more
than seedlings and a spade. Species most
appropriate to a given site must be selected,
follow-up care must be available, and long-
term management must be provided if the
success of these programs is to be ensured.

The environmental problems facing
Illinois, the nation, and the planet are grave
indeed. Yet we are learning the important role
that forests can play in mitigating some of these
problems. We have, however, only begun to
realize the enormity of the task. We have only
begun to take the actions needed to create a
sustainable world.

LITERATURE CITED

AMERICAN FORESTRY ASSOCIATION. 1988. Plant a
tree—cool the globe. Urban Forest Forum 8:1, 11.

ANDERSON, R.C. 1970. Prairies in the prairie state.
Transactions of the Illinois State Academy of
Science 63(2):214-221.

Biake, J.G., AND J.R. Karr. 1987. Breeding birds
of isolated woodlots: area and habitat relationships.
Ecology 68:1724-1734.

Symposium Proceedings: Our Living Heritage 373

Biyth, J.E., D.R. McCurpy, J.H. BURDE, AND
W.B. Situ. 1985. Fuelwood production and
sources from roundwood in Illinois, 1983. U.S.
Department of Agriculture, Forest Service, Resource
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BiyTH, J.E., J.A. SEsTER, AND G.K. RaILe. 1987.
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change detection of urban land cover in Illinois using
Landsat thematic mapper data. Proceedings of the
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MD. (in press)

EBINGER, J.E. 1986. Sugar maple, a management
problem in Illinois forests? Transactions of the
Illinois State Academy of Science 79(1 & 2):25-30.

Essex, B.L., AND D.A. GANSNER. 1965. Illinois’
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Forest Service, Resource Bulletin LS-3. St. Paul,
MN. 56 p.

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mental evaluations using birds and their habitats.
Illinois Natural History Survey Biological Notes 97.
39 p.

Haun, J.T. 1987. Illinois forest statistics, 1985. U.S.
Department of Agriculture, Forest Service, Resource
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Harris, L.D. 1984. The fragmented forest. Univer-
sity of Chicago Press, Chicago, IL. 211 p.

ILLINOIS COUNCIL ON FORESTRY DEVELOPMENT.
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1990. Forest fragmentation and biomass degradation
in West Malaysia during 1972-1982. Bulletin of the
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374 Illinois Natural History Survey Bulletin

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resources of Illinois, 1820-1980, with accompanying
maps on forest volume, diversity, and composition.
Map. Illinois Natural History Survey, Champaign.

Iverson, L.R., AND D.M. KETZNER. 1988. The
Illinois Plant Information Network user’s guide.
Illinois Natural History Survey Internal Document.
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Risser, C.D. BURNETT, AND R.G. RAYBURN. 1989.
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spatial and temporal trends. Illinois Natural History
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J.L. Jones. 1979. Land-use conflicts with natural
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the conterminous United States. American Geo-
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Illinois’ forest resources. U.S. Department of Agri-
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St. Paul, MN. 113 p.

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Wildlife and Natural Resources Conference
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Vol. 34 Art. 4

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Illinois Forestry Research Report 84-2. 5 p.

Forest Succession in the Prairie Peninsula of Illinois

John E. Ebinger, Botany Department, Eastern Illinois University, and William E.
McClain, Division of Natural Heritage, Illinois Department of Conservation

Presently most of central Illinois is in the
Grand Prairie Natural Division (Schwegman
1973), classified as a part of the prairie
peninsula of the oak—hickory forest region by
Braun (1950), as a mosaic of bluestem prairie
and oak—hickory forest by Kiichler (1964), and
as a part of the prairie—deciduous forest ecotone
by Davis (1977). At the time of settlement by
Europeans, prairie dominated most of Illinois.
Forests were common, however, occurring on
rough terrain such as moraines and dissected
valleys of streams and rivers and as isolated
groves on the flat to gently rolling prairie.

During postglacial times, the vegetation
of Illinois changed extensively (King 1981).
Pollen diagrams from the prairie peninsula in
Illinois record the climatically related vegeta-
tion shifts that have occurred since the late
Pleistocene. The pollen record for Chatsworth
Bog, Livingston County, in the center of the
prairie peninsula, suggests that a mosaic of
open spruce woodlands and tundra existed
there from 14700 to 13800 BP. This cover type
in turn was replaced by an ash/tundra assem-
blage that reflected the slowly increasing
temperatures of the late-glacial from 13800 to
11600 BP. After 11600 BP, pollen from
deciduous trees and shrubs increased dramati-
cally, starting with cool-climate species (birch,
hazel, black ash) and followed by such warm-
tolerant taxa as elms, oaks, and hickories. By
8300 BP, prairie dominated the area as indi-
cated by a dramatic decrease in tree pollen and
a corresponding increase in the amount of
pollen from herbaceous plants. Oak pollen was
still present, however, suggesting that prairie
vegetation was probably common on the drier
flat uplands while the lowlands and river
valleys retained their,forest cover. These open
expanses of prairie with savanna and forest
communities restricted to the more dissected
lands were what the early European settlers
found when they entered the prairie peninsula
of Illinois in the early 1800s.

The presettlement distribution of the
major vegetation types in Illinois (prairie,
savanna, and forest) was determined largely by
firebreaks such as lakes and rivers and by
topographic relief that controlled the frequency
and intensity of fire (Gleason 1913; Wells
1970; Grimm 1984). Gleason (1913) found that
forests were more extensive on the east side of
firebreaks, while prairie tended to be more
extensive on the west side. This distribution
pattern was the result of prevailing westerly
winds that carried fires to the western sides of
firebreaks, thus encouraging the development
of prairies. In contrast, the eastern sides were
protected from fires, and forest developed at
these locations.

PRESETTLEMENT FORESTS

In presettlement times, according to survey
records of the General Land Office, prairie
occupied 61.2% of Illinois and forest and
savanna accounted for 38.2% (Iverson et al.
1989). In general, prairie vegetation was most
common on flat to gently sloping ground;
savanna and forest were most common in
dissected areas. The segregation of forest,
savanna, and prairie on the basis of topography
apparently occurred because dissected land-
scapes do not readily carry fire. For the most
part, these dissected landscapes have well-
developed drainage systems that support
permanent or temporary streams, which serve
as firebreaks. In addition, fires in hilly areas
tend to move up slope relatively rapidly due to
rising convection air currents, but convection
currents work against fires when they move
down hill, not uncommonly causing them to
burn themselves out.

A great deal of vegetation information
can be obtained from survey records of the
General Land Office (Bourdo 1956). The job of
the surveyors was to establish a grid system of
township, range, and section lines by the

375

376 Illinois Natural History Survey Bulletin

placement of section and quarter section corner
posts. In prairie and marsh areas, only posts
were used. In timbered areas, however, two (or
four) witness trees were blazed, and the
distance and direction of these trees from the
corner posts were recorded along with their
species and estimated diameter at breast height
(dbh). Because the placement of the corner
posts and the selection of witness trees were
essentially random, the principles of the
distance method (Cottam and Curtis 1956) can
be applied to the witness tree data and the
composition and tree density of the presettle-
ment savannas and forests determined.

In Illinois, several researchers have used
survey records of the General Land Office to
determine the extent, composition and densities
of tree species for various counties. Some of
their studies are summarized here and indicate
the extent and composition of the presettlement
vegetation of the prairie peninsula.

Kilburn (1959) found that the original
forest in Kane County consisted largely of oak
openings composed of pure bur oak or bur/
white oak stands. Lowlands and swamp forests
were found along rivers and streams, but a
more mesic forest occurred on the heavier soils
of the Big Woods area. Overall, three-fifths of
the county was prairie. Topography accounted
for most of the vegetation pattern: level areas
were in prairie vegetation; protected ravines,
valleys, steep bluffs, and hills were largely
forested. Overall, 87% of the witness trees
recorded by the surveyors were oaks and
hickories.

In Lake County, the situation was similar.
Oak and hickory species accounted for 95% of
the trees recorded (Moran 1976). In this county,
however, savanna was the dominant vegetation
type, occupying 51% of the area. It was found
mostly on rolling uplands that were frequently
broken by small wetlands or streams; bur oak
was by far the most common species with black
and white oaks in lesser numbers. Prairie, wet
prairie, and marsh occupied 33% of the county
while forests occurred in the remaining 16%.
For the most part, prairies were situated on flat
terrain and forests were restricted to areas of
rough topography or where natural firebreaks
afforded some protection.

In McLean County, located in west-
central Illinois, the presettlement vegetation
was 89.5% prairie, 5.4% savanna, 1.8% open
forest, and 3.3% closed forest (Rodgers and

Vol. 34 Art. 4

Anderson 1979). The forested areas occurred
on the more rugged topography associated with
rivers, streams, and glacial moraines. White
and black oaks were the most numerous species
recorded, but in the closed forests (273 trees/
ha) the more mesic species (i.e., sugar maple,
elm, red oak, buckeye) accounted for about
one-third of the trees present. These more
shade-tolerant, mesic species, which for the
most part are fire-sensitive, occupied sheltered
ravines and areas adjacent to streams where
fires occurred infrequently. In contrast, the
relatively shade-intolerant oaks, which depend
on periodic fires to maintain their dominance,
were more common on less dissected uplands.

In adjacent Mason County, similar results
were obtained (Rodgers and Anderson 1979).
Located in the Illinois River Sand Area Section
(Schwegman 1973), on soils developed from
deep sand deposits laid down by glacial
meltwater during the Pleistocene (Willman and
Frye 1970), prairie was the dominant vegeta-
tion type, occupying 67.7% of the county.
Savanna (14.4%) and forest (13.3%) occurred
on most of the remaining land and 4.6% was
covered by lakes and swamps. The dominant
tree species in the presettlement forests and
savannas were shade-intolerant, fire-tolerant
black and blackjack oaks. In the closed forests
(263 trees/ha), the oaks and hickories were still
the most numerous species. The more mesic,
shade-tolerant, fire-sensitive tree species (1.e..
sugar maple, elm, walnut) were also found in
the closed forests, particularly in areas of rough
topography.

In Douglas County, near the southern
edge of the Grand Prairie Natural Division
(Schwegman 1973), prairie was the most
widespread plant community (85%). Closed
forest, which was generally restricted to the
major river systems, accounted for the remain-
ing 15%. These forests were dominated by
white and black oaks and hickories, species that
accounted for 70% of the witness trees re-
corded by the surveyors. Mesic, shade-tolerant,
fire-sensitive species were present but restricted
to areas of rough topography and river valleys
(Ebinger 1986a).

Prairie was the most widespread vegeta-
tion type (60%) in Coles County, the southern
half of which is located on the Shelbyville
Moraine, the terminal moraine of Wisconsin
glaciation. Prairie was most common on the flat
to gently rolling uplands in the northern and

April 1991

central parts of the county. Forests, which
accounted for most of the remaining 40%, were
restricted to the rough topography of the
terminal moraine and to the valleys of the
Kaskaskia and Embarras rivers. More than 80%
of the witness trees recorded were oaks and
hickories, with white, black, and red oaks most
numerous. Again, more mesic species were
restricted to rough topography (Ebinger 1987).
Information extrapolated from the
records of early surveyors indicates that prairie
vegetation dominated most of Illinois in
presettlement times and was found on the flat
to gently rolling uplands throughout most of the
state. Savannas and forests, in contrast, were
more common in rough topography, especially
in the driftless areas, along major waterways,
and where morainal systems provided topo-
graphic relief. For the most part, savannas
developed on sites where the frequency of fire
was reduced, thereby permitting the establish-
ment of fire-tolerant tree species (Anderson
1970; Anderson and Anderson 1975; Grimm
1984; Anderson and Brown 1986). Forests,
particularly closed forests, developed in places
of rough relief, in river valleys, and in other
protected areas where fires were less likely to
occur. Oaks and occasionally hickories
dominated the open savannas. In the forests,
oaks and hickories were also the dominant
species, but more mesic, shade-tolerant, fire-
sensitive tree species were common forest
components. Furthermore, the transition from
forest to prairie varied from being rather abrupt
in some locations in the prairie peninsula to
others where savannas formed a broad transi-
tion between forest and prairie (Nuzzo 1986).
This transition was probably determined by
topographic relief, firebreaks, fuel loads, and
other edaphic and climatic factors that con-
trolled the frequency and intensity of fires.

PRESENT SUCCESSION TRENDS

During the past century and a half of agricul-
tural development, periodic fires have ceased in
the prairie peninsula, and the oak savannas and
open oak forests on the uplands have become
closed-canopy forests. As a result, these
woodlots have been changing to forests domi-
nated by such mesic, shade-tolerant, fire-
sensitive species as sugar maple, American and
red elms, white and green ashes, and ironwood
(Anderson and Adams 1978; Adams and
Anderson 1980; Ebinger 1986b).

Symposium Proceedings: Our Living Heritage 377

In particular, sugar maple has increased
in importance in most Illinois forests (Iverson
et al. 1989). If this trend continues, many of the
oak—hickory forests, their understories, and the
wildlife that depends upon them will be in
serious trouble in the near future. Even the best
quality oak—hickory communities are appar-
ently undergoing an irreversible change as
sugar maple and other mesic, shade-tolerant
species replace many of the original forest
components. Almost no work has been done
concerning methods to reverse this trend, and
the problem now concerns many ecologists and
managers of natural areas.

Many of the better quality forests that
presently exist in the prairie peninsula have
been surveyed during the past thirty years. In a
few of them, sugar maple is not an important
component, though other mesic species are
sometimes common. At Walnut Point State
Park in Douglas County (Ebinger et al. 1977),
sugar maple is rarely encountered, and oaks
and hickories are by far the most numerous
species. In the forests and savannas of the
Kankakee Sand Area Section (McDowell et al.
1983) and the Illinois River Sand Area Section
(Rodgers and Anderson 1979) oaks dominate
and mesic species are rarely encountered. In
most of the stands studied, however, mesic
species, particularly sugar maple, are relatively
important components. These mesic species are
also well represented in the seedling and
sapling categories and in the smaller diameter
classes. Oaks and hickories, in contrast, are
poorly represented in these categories.

Mesic, shade-tolerant, fire-sensitive
species are common components of many
recently surveyed forests in the prairie penin-
sula. Two “prairie grove forests” in Champaign
County have been surveyed at various times in
the past, and sugar maple is an important
component in both. In Trelease Woods
(Boggess 1964; Pelz and Rolfe 1977), sugar
maple dominates the seedling and sapling
categories as well as most of the diameter
classes. Similar results were obtained for
Brownfield Woods by Boggess and Bailey
(1964) and Miceli et al. (1977).

An inventory of the woody vegetation of
Funks Forest Natural Area in McLean County
was conducted by Boggess and Geis (1966).
This forest is an example of a mesophytic
forest that is transitional between the upland
oak—hickory cover type and the “prairie grove

378

forest.” Sugar maple, the dominant species in
Funks Forest, is followed closely by white oak
and elm. Sugar maple and white oak, however,
represent two distinct age classes. White oak,
which predominates in the 30-inch-diameter
class, is a “pioneer” species; and sugar maple,
which predominates in the 16-inch-diameter
class, has perhaps been increasing steadily in
importance during the past century.

One recently documented example of the
increase in importance of sugar maple is at
Baber Woods Nature Preserve in Edgar County.
This 16-ha forest is located on the flat to gently
rolling ground just north of the Shelbyville
Moraine, the terminal moraine of Wisconsin
glaciation. Two decades ago, McClain and
Ebinger (1968) reported that sugar maple
ranked second in importance in the woods and
dominated the seedling, sapling, and smaller
diameter classes. In a more recent survey of the
same area, Newman and Ebinger (1985) found
that this trend had continued. Sugar maple was
now first in importance, and the number per
acre had almost doubled. Further, sugar maple
continued to dominate the seedling and sapling
categories and accounted for nearly half of the
individuals in smaller diameter classes. Sugar
maple and oaks represent two distinct age
classes in Baber Woods, as shown in Figure 1.
These curves show that oaks predominate the
larger diameter classes and suggest that these
species have been an important forest compo-
nent for an extended period of time. Sugar
maple, in contrast, predominates the smaller
diameter classes and has probably been
increasing steadily during the past century. The
large number of sugar maple seedlings,
saplings, and smaller diameter trees suggests a
continuation of this trend.

Table | indicates when sugar maples
began to increase in importance in Baber
Woods. In nearly every quadrat, sugar maple
increased in number, size, and importance from
1965 to 1983. In addition, the number, size, and
importance of sugar maple decreased from the
northwestern corner of the woods, becoming
smaller and less common toward the southeast-
ern corner. This pattern suggests that sugar
maple probably occurred in the ravines that
exist just to the north and west of the woods,
where in presettlement times it was probably
protected from fire due to the rough topogra-
phy. With the cessation of fire, this fire-
sensitive species has been able to invade the
upland forests that still exist in the area.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 4

Another indication of the increase of
sugar maple in Baber Woods is the distribution
of this species and the oak species by diameter
classes for the 1965 and 1983 surveys (Table
2). Sugar maple increased in all diameter
classes between 1965 and 1983, particularly in
two diameter classes, 10-19 and 20-29 cm.
Sugar maple showed an overall increase of
nearly 30 trees per hectare between the two
surveys. In contrast, oak species decreased in
numbers, dramatically so in the lower diameter
classes, with increases occurring only in classes
60—69 cm in diameter and above (Table 2).
Overall, species density increased in the
woodlot, from 258.6 stems/ha in 1965 to 277.3
stems/ha in 1983. Most of this increase is due
to sugar maple and other mesic species that are
tolerant of shade and sensitive to fire. Presently
the oaks are common in the larger diameter
classes because of recruitment from the smaller
diameter classes. Oak reproduction is sparse
(McClain and Ebinger 1968; Newman and
Ebinger 1985), and as the veteran trees die,
fewer oaks are available to fill the canopy gaps.
In contrast, sugar maple, with its high gap-

Total basal area (square feet)

250
All species
200
150
100- Oaks
50, 2 YS, Sy
Pe maple ‘
WS

10. 15 20) 25. 30" 35) Se
Diameter class (inches)
Figure 1. Smooth curves of basal area by diameter
class for sugar maple, all oak species combined, and
all species combined at Baber Woods, Edgar
County, Illinois. Source: Ebinger 1986b.

April 1991 Symposium Proceedings: Our Living Heritage 379

phase replacement potential, is able to take within 4 m of the ground. They are probably
advantage of these canopy openings (Ebinger remnants from a time when this forest was an
1986). open, upland savanna. The average diameter of
Within Baber Woods are a number of these open-grown white oaks is 101.6 cm dbh,
large open-grown white oaks. In a walk- and two that had died recently were cut and
through survey conducted during the early aged at 313 years. Both had fire scars at 65 and
spring of 1990, twenty-six large, open-grown 77 years, indicating that in the past fires were
white oaks were observed. All have open, probably common in the area. Five other oaks
round crowns and large lower branches, some that had died recently were also cut and aged.

Table 1. Distribution of sugar maple in Baber Woods Nature Preserve, Edgar County, Illinois, for the surveys
of 1965 (McClain and Ebinger 1968) and 1983 (Newman and Ebinger 1985). The following information is
given for each quadrat (1 ha): the number of stems present (above 10 cm dbh), the number of stems exceeding
40 cm dbh, the average diameter (cm), and the importance value (relative density and relative dominance) for
sugar maple. Highest possible importance value is 200. The northern edge of the woods is represented in
quadrats | through 4.

Quadrat | Quadrat 2 Quadrat 3 Quadrat 4
1965 1983 1965 1983 1965 1983 1965 1983
Number of individuals 140 153 158 152 104 124 82 102
Number <40 cm dbh 8 14 12 17 3 4 2 9
Average diameter (cm) 23 23.6 2271 Psy) 19.3 20.9 20.1 22.7
Importance value 78.1 86.2 82.4 98.6 52.6 68.1 42.1 57.5
Quadrat 5 Quadrat 6 Quadrat 7 Quadrat 8
1965 1983 1965 1983 1965 1983 1965 1983
Number of individuals 98 134 9] 138 90 100 45 70
Number <40 cm dbh 7 9 5 6 3 6 l 4
Average diameter (cm) 20.6 20.5 19.1 19.4 18.9 21.6 19.5 20.4
Importance value S12 71.9 45.9 66.7 45.8 58.3 25.1 37.8
Quadrat 9 Quadrat 10 Quadrat 11 Quadrat 12
1965 1983 1965 1983 1965 1983 1965 1983
Number of individuals 60 95 29 101 38 74 34 58
Number <40 cm dbh 9 14 — 1 a 2 — l
Average diameter (cm) 23:5 21.1 15.6 ISS 19.8 20.2 18.5 20.9
Importance value 40.8 a3} 72 13.8 Shy) 25.4 45.8 20.4 34.5

Table 2. Density (number/ha) in broad diameter classes for sugar maple, oak species, and all other species in
Baber Woods Nature Preserve, Edgar County, Illinois, for the surveys of 1965 (McClain and Ebinger 1968)
and 1983 (Newman and Ebinger 1985).

Sugar maple Oak species Other species Totals
Diameter class 1965 1983 1965 1983 1965 1983 1965 1983
10-19 cm 42.6 58.9 7.6 Bu 50.0 62.1 100.2 124.7
20-29 cm 17.8 24.7 10.9 4.9 17.0 WZ) 45.7 47.3
30-39 cm 7.3 10.6 14.7 8.3 19.0 13.2 41.0 32.1
40-49 cm 2.4 4.6 Wee, 11.6 15.4 12.6 g555 28.8
50-59 cm 0.6 1.0 16.2 13.0 5.6 nS, 22.4 21.9
60-69 cm 0.1 0.3 Ie) 11.1 0.9 2.1 8.5 13.5
70-79 cm - - na 4.4 0.3 0.6 3.0 5.0
80-89 cm - - 1.1 23 0.1 0.2 1.2 2
90+ cm 7 1.1 1.5 - - 1.1 1.5

Total 70.8 100.1 79.5 60.8 108.3 116.4 258.6 277.3

380 Illinois Natural History Survey Bulletin

These were forest-grown trees with straight
trunks, no low branches, and an average
diameter of 68.2 cm. They varied in age from
140 to 158 years, with an average age of 148
years. In contrast were the increment cores
obtained from 30 sugar maples in various parts
of the woodlot. Those from the northwestern
part of the woods, where the largest individuals
occurred, averaged 44.7 cm dbh and had an
average age of 107.6 years. Sugar maples from
the northeastern and southeastern corners of the
woodlot were smaller and younger (Table 3).

The data suggest that before European
settlement, the area now known as Baber
Woods was an open, white oak savanna
maintained by periodic fires. This community
was probably parklike with an understory of
prairie grasses and forbs. With the cessation of
fire, the number of seedlings increased and
began to fill the gaps in the canopy between the
large open-grown oaks. As shade increased,
moisture levels within the forest probably
increased, creating a habitat for more mesic,
shade-tolerant, fire-sensitive species such as
sugar maple.

LITERATURE CITED

ApaMs, D., AND R.C. ANDERSON. 1980. Species
response to a moisture gradient in central Illinois
forests. American Journal of Botany 67:381—392.

ANDERSON, R.C. 1970. Prairies in the prairie state.
Transactions of the Illinois State Academy of
Science 63(2):214—221.

ANDERSON, R.C., AND D.E. ADAMS. 1978. Species
replacement patterns in central Illinois white oak
forests. Pages 284-301 in P. Pope, ed. Proceedings
of the Central Hardwood Forest Conference II.
Purdue University, IN.

ANDERSON, R.C., AND M.R. ANDERSON. 1975. The
presettlement vegetation of Williamson County,
Illinois. Castanea 40:345—363.

Vol. 34 Art. 4

ANDERSON, R.C., AND L.E. Brown. 1986. Stability
and instability in plant communities following fire.
American Journal of Botany 73:364—368.

Boccess, W.R. 1964. Trelease Woods, Champaign
County, Illinois: woody vegetation and stand
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Boccess, W.R., AND L.W. BaiLey. 1964.
Brownfield Woods, Illinois: woody vegetation and
changes since 1925. American Midland Naturalist
71:392-401.

Boccess, W.R., AND J.W. Geis. 1966. The Funk
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actions of the Illinois State Academy of Science
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Bourbo, E.A. 1956. A review of the General Land
Office Survey and of its use in quantitative studies of
former forests. Ecology 37:754-768.

Braun, E.L. 1950. Deciduous forests of eastern
North America. Blakiston, Philadelphia, PA. 596 p.

Cottam, G., AND J.T. Curtis. 1956. The use of
distance methods in phytosociological sampling.
Ecology 37:271-287.

Davis, A. 1977. The prairie-deciduous forest
ecotone in the upper Middle West. Annals of the
Association of American Geographers 67:204—213.

EBINGER, J.E. 1986a. Presettlement vegetation of
Douglas County, Illinois. Erigenia 7:15—22.

EBINGER, J.E. 1986b. Sugar maple, a management
problem in Illinois forests? Transactions of the
Illinois State Academy of Science 79(1 & 2):25-30.

EBINGER, J.E. 1987. Presettlement vegetation of
Coles County, Illinois. Transactions of the Illinois
State Academy of Science 80(1 & 2):15—24.

EBINGER, J.E., P.E. PHILLIPPE, AND L.R. PHILLIPPE.
1977. Woody vegetation of an upland forest in
Douglas County, Illinois. Castanea 42:285-293.

Table 3. Tree rings and diameters (dbh) of sugar maples at selected sites in Baber Woods Nature Preserve,

Edgar County, Illinois.

Diameter (cm)

Growth ring

Area Range
Northwest corner 37.4-S9.7
Northeast corner 26.0-35.8
Southeast corner 17.5-35.5

Average Range Average
44.7 101-116 107.6
30.4 52-91 70.6
25.7 51-71 61.1

April 1991

GLEASON, H.A. 1913. The relation of forest
distribution and prairie fires in the Middle West.
Torreya 13:173-181.

Grimm, E.C. 1984. Fire and other factors controlling
the big woods vegetation of Minnesota in the mid-
nineteenth century. Ecological Monographs
54:291-311.

Iverson, L.R., R.L. OLiver, D.P. TucKER, P.G.
Risser, C.D. BURNETT, AND R.G. RAYBURN. 1989.
Forest resources of Illinois: an atlas and analysis of
spatial and temporal trends. Illinois Natural History
Survey Special Publication 11. 181 p.

KiLBurn, P.D. 1959. The forest-prairie ecotone in
northeastern Illinois. American Midland Naturalist
62:206-217.

Kino, J.E. 1981. Late Quaternary vegetational
history of Illinois. Ecological Monographs
51:43-62.

KUcHLer, A.W. 1964. Manual to accompany the
map, potential natural vegetation of the contermi-
nous United States. American Geographical Society
Special Publication 36. American Geographical
Society, New York. 39+116 p.

McC alin, W.E., AND J.E. EBINGER. 1968. Woody
vegetation of Baber Woods, Edgar County, Illinois.
American Midland Naturalist 79:419—428.

McDowe LL, B., J. NEWMAN, AND J.E. EBINGER.
1983. Survey of the woody vegetation of the
Kankakee Sand Area Section of Indiana and Illinois.
Proceedings of the Indiana Academy of Science
93:187-193.

Miceut, J.C., G.L. RoLFe, D.R. PELZ, AND J.M.
EpGINGTON. 1977. Brownfield Woods, Illinois:
woody vegetation and change since 1960. American
Midland Naturalist 98:469—476.

Moran, R.C. 1976. Presettlement vegetation of
Lake County, Illinois. Pages 12-18 in D.C. Glenn-
Lewin and R.Q. Landers, Jr., eds. Proceedings of the
Fifth Midwest Prairie Conference. lowa State
University, Ames.

Newman, J.A., AND J.E. EBINGER. 1985. Woody
vegetation of Baber Woods: composition and change
since 1965. Pages 178-180 in J.O. Dawson and K.A.
Majerus, eds. Fifth Central Hardwood Forest
Conference. Society of American Foresters Publica-
tion 85-05. Department of Forestry, University of
Illinois at Urbana-Champaign.

Nuzzo, V.A. 1986. Extent and status of midwest oak
savanna: presettlement and 1985. Natural Areas
Journal 6:6—36.

Symposium Proceedings: Our Living Heritage 381

Petz, D.R., AND G.L. Rotre. 1977. Stand structure
and composition of a natural mixed hardwood forest.
Transactions of the Illinois State Academy of
Science 69(4):446—454.

Ropcers, C.S., AND R.C. ANDERSON. 1979.
Presettlement vegetation of two prairie peninsula
counties. Botanical Gazette 140:232—240.

SCHWEGMAN, J. 1973. Comprehensive plan for the
Illinois Nature Preserves System. Part 2. The natural
divisions of Illinois. Illinois Nature Preserves
Commission, Rockford, IL.

WELLS, P.V. 1970. Historical factors controlling
vegetation patterns and floristic distribution in the
Central Great Plains region of North America. Pages
211-234 in W. Dort, Jr., and J.K. Jones, Jr., eds.
Pleistocene and recent environments of the Central
Great Plains. University Press of Kansas, Lawrence.

WILLMAN, H.B., AND J.C. Frye. 1970. Pleistocene
stratigraphy of Illinois. Illinois State Geological
Survey Bulletin 94. 204 p.

Effects of Forest Fragmentation on Illinois Birds

Scott K. Robinson, Illinois Natural History Survey

Abstract. The forests in Illinois are among the
most fragmented in North America. Most re-
maining tracts are small, isolated, and domi-
nated by “edge” habitats. Populations of many
forest species, especially those that breed in the
forest interior, have been declining, and many
characteristic forest species do not occur in
woodlots below a certain minimum size. Data
from small woodlots (<65 ha; 170 acres) in the
Lake Shelbyville area of central Illinois suggest
that reproductive failure may be at least partly
responsible for these trends, especially among
the neotropical migrants that breed in Illinois
but winter in the tropics. Most nests fail
because of brood parasitism by brown-headed
cowbirds (Molothrus ater) (76% of all nests of
neotropical migrants) or because of nest
predation (80% of all nests).

Brown-headed cowbirds, which are
abundant throughout Illinois, pose a particu-
larly severe threat because they lay their eggs
in the nests of host species, which go on to
raise cowbirds instead of their own young.
Parasitized nests in the Lake Shelbyville area
averaged 3.3 cowbird eggs per parasitized nest.
All 19 wood thrush (Hylocichla mustelina)
nests were parasitized with an average of 4.6
cowbird eggs per nest. Only about 10% of the
birds of all species caught in midsummer were
juveniles. These data strongly suggest that the
reproduction of neotropical migrants in very
small woodlots is insufficient to compensate
for adult mortality, a result consistent with the
population declines observed in the Shelbyville
area.

Birds nesting in much larger tracts (up to
2,024 ha; 5,000 acres) in the Shawnee National
Forest appear to face similar problems. A crew
of 14 workers located over 400 nests in 1989
and discovered that cowbird parasitism and
nest predation rates were high, even deep in the
forest interior. In contrast to studies elsewhere,
cowbirds were found throughout each study
area, regardless of the proximity of edges. Over

382

55% of all nests were parasitized and an
average of 60% of all nests were destroyed by
predators. As in Shelbyville, wood thrushes
suffered most from cowbirds: 90% of all nests
parasitized and an average of 3.2 cowbird eggs
per nest. Other species that suffered high
(>70%) parasitism rates were the red-eyed
vireo (Vireo olivaceus) and the scarlet (Piranga
olivacea) and summer tanagers (P. rubra). A
few species reproduced successfully in spite of
the abundance of nest predators and cowbirds.
Worm-eating (Helmitheros vermivorus) and
Kentucky warblers (Oporornis formosus) hide
their nests effectively, and for these species
young outnumbered adults in midsummer
samples of birds caught in mist nests.

These results suggest that management
decisions will have to take into account
differences among species in susceptibility to
forest fragmentation. The cowbird situation is
more serious than has been anticipated and
apparently cannot be solved simply by mini-
mizing edges as has been proposed elsewhere
in the Midwest. At least a few species, for
example, the wood thrush, may be in serious
trouble throughout the Midwest and should
receive special management attention.

Session Two: Prairies and Barrens

The chance to find a pasque-flower is a right as inalienable as free speech—A\do Leopold

The first Europeans to see the Illinois country
had crossed a vast ocean, snaked their way
through a nearly impenetrable mountain range,
and forged a path through a thousand miles of
dense, primeval forest. They did it with indomi-
table spirit and by sheer force of will. Yet when
they reached the edge of the eastern deciduous
forest, approximated today by the Indiana—
Illinois border, they stopped in wonder. Here
was a landscape so different from those with
which they were familiar that they had no word
for it. In time this landscape came to be known
as “prairie,” a word derived from the French
word for meadow.

At first, early settlers avoided living on
the prairie because the treeless grasslands were
thought to be infertile. They did not provide
much needed building materials, fuel, and
water. Instead, they offered the prospect of
menacing prairie fires and howling winter
storms. Soon, however, the settlers realized that
prairie made excellent cropland, especially after
John Deere invented the moldboard plow that
allowed virgin prairie soil to be broken. The
wild prairies became cropland at an astonishing
rate—approximately 3.3% per year. Over
300,000 people settled on the prairie during the
decade of the 1830s, and by 1860 nearly all the
prairies had disappeared.

At least 23 different kinds of prairies are
found in Illinois—add barrens, savannas, and
glades and the list increases to over 30. These
various prairies once occupied nearly 22 million
acres of the state. Today they are confined to
about 3,000 acres, less than 0.01% of their
original extent. Unfortunately, it is easier to find
examples of the prairie’s influence in the
“prairie” state—Prairie Street, Prairie State
Games, Prairie Farms Dairy, Prairieview
Estates, Prairie Technology—than it is to find
an actual prairie. Prairie remnants persist,
however, along railroad lines, in pioneer
cemeteries, even on the grounds of industrial
complexes, growing in a forgotten corner of
some storage yard yet to be developed.

Over 200 species of plants characteristi-
cally inhabit Illinois prairies. Although this
number is relatively low compared with a
typical undisturbed woodland, a small prairie
remnant—as little as five acres—can be
surprisingly diverse with more than 120
species of plants. All present-day Illinois
prairies, however, are incomplete, fragmented
ecosystems and lack the large herbivores that
were so important in their development.

What if Illinoisans had had the foresight
to preserve only 100 square miles of virgin
prairie in central Illinois? What a tremendous
natural resource and botanical laboratory that
would be today! Inevitable though the destruc-
tion of the prairie may have been, it is truly
unfortunate that prairies will be visualized by
future generations as isolated pockets of native
vegetation, persisting in a world that passed
them by. Ironically, the French word for
meadow, so incongruous when applied to this
once vast grassland, now seems totally
appropriate.

The session opened with a broad
historical perspective of the tallgrass prairie.
The papers that followed focused tightly on
two aspects of that prairie—the remnant-
restricted prairie and savanna insects of the
Chicago region and the response of prairie
birds to habitat fragmentation.

383

Illinois Prairies: A Historical Perspective

Roger C. Anderson, Department of Biology, Illinois State University

The grasslands of central North America
originated in the Miocene—Pliocene transition,
about 7—S million years before present (YRBP)
and were associated with the beginning of a
drying trend. The Miocene uplift of the Rocky
Mountains created a partial barrier between
moist Pacific air masses and the interior portion
of the continent. The spread of the Antarctic ice
sheet, by tying up atmospheric moisture, also
contributed to increased aridity. Woody plants
are generally less well adapted to drought than
most grass species, and the spread of grasslands
consequently occurred at the expense of forests.
As the grassland expanded, numbers of grazing
and browsing animals increased, an indication
that the association of grasses and grazers
occurred over a long period of time (Stebbins
1981; Axelrod 1985).

The prairies of Illinois are part of the
central grassland, a large triangular-shaped area
that has its base along the foothills of the
Rocky Mountains from the Canadian provinces
of Saskatchewan and Manitoba southward
through New Mexico into Texas. The apex of
the triangle, the prairie peninsula (Transeau
1935), extends eastward into the Midwest and
includes the prairies of Illinois, Iowa, Indiana,
Minnesota, Missouri, and Wisconsin with
scattered outliers in southern Michigan, Ohio,
and Kentucky. Because the Rocky Mountains
intercept moist air masses moving westward
from the Pacific Coast, the grassland lies in the
partial rain shadow to the east. From west to
east within the central grasslands, annual
precipitation increases from 25—38 cm to
75—100 cm and becomes more reliable;
potential evapotranspiration decreases, the
number of days with rainfall increases, and
periods of low humidity and periodic droughts
in July and August decrease (Risser et al.
1981). Associated within this climatic gradient
is a shift in the grassland species dominating
the vegetation.

384

Ecologists traditionally have separated
the central grassland into three major west-east
divisions. The arid western shortgrass prairie is
dominated by such species as buffalo grass
(Buchloe dactyloides), blue grama (Bouteloua
gracilis), and hairy grama (B. hirsuta) that
reach heights of only 30—45 cm. The mixed-
grass prairie occupies the middle sector of the
central grassland and is dominated by grasses
that are 60—120 cm tall, including little
bluestem (Schizachyrium scoparium), needle-
grasses (Stipa spartea and S. comata), and
wheatgrasses (Agropyron smithii and A.
dasystachyum). The prairies of Illinois are in
the eastern portion of the remaining division of
the central grassland, the tallgrass prairie
(Figure 1). In this area of relatively high
rainfall, the dominant grasses on mesic sites
include big bluestem (Andropogon gerardi),
Indian grass (Sorghastrum nutans), and
switchgrass (Panicum virgatum)—grasses that
reach heights of 1.8—3.6 m. On poorly drained
sites supporting wet prairies, prairie cordgrass
(Spartina pectinata) and bluejoint grass
(Calamagrostis canadensis) are dominant
species; little bluestem and sideoats grama
(Bouteloua curtipendula) are important grasses
on dry sites (Weaver 1954; Risser et al. 1981;
Bazzaz and Parrish 1982). Figure 2 indicates
how these major grass species follow a soil
moisture gradient.

Illinois prairies, which dominated about
60% of the state prior to the extensive settle-
ment and alteration of the landscape by
Europeans, developed since the last glacial
advance. According to King (1981), as the last
of the Wisconsinan age ice sheet retreated from
the northeastern portions of the state, mesic
deciduous forests dominated most of the
landscape. A drying and warming trend began
about 8,700—7,900 YRBP, and prairie began to
replace deciduous forests in southern Illinois.
Prairie influx into central Illinois occurred

April 1991

about 8,300 YRBP and concomitantly
oak-hickory forest began to replace mesic
forest in the northern portion of the state.
Prairies occupied much of the state during the
Hypsithermal Period (8,000—6,000 YRBP),
which was the hottest and driest part of the
Holocene. The climate became cooler and more
moist following the Hypsithermal, but prairie
stabilized throughout much of Illinois (King
1981).

Because of increased rainfall and reduced
evapotranspiration, the climate is increasingly
favorable for the growth of trees from west to
east in the central grassland. Consequently, in
Illinois and the rest of the prairie peninsula, the
average climate for approximately the past
5,000 years appears to have been more favor-
able for forest than for grassland. However, this
region has had periodic droughts during which
the forest retreated and the grasslands advanced
or were maintained. To understand factors
influencing the persistence of grasslands in this
region, we must consider the extremes of
climate and not the average. Britton and
Messenger (1970) suggested that the droughts
that are most detrimental to woody species are
those that do not permit deep recharge of soil
moisture during the winter months. On soils

Figure 1. Presettlement distribution of the tallgrass
prairie. Adapted from National Geographic (1980)
157(1):43.

Symposium Proceedings: Our Living Heritage 385

without drainage restrictions, trees generally
root at greater depths than grasses and rely on
moisture stored deep in the soil during
droughty periods in midsummer. Interestingly,
Britton and Messenger (1970) presented data
showing that areas of the Midwest that did not
experience deep soil moisture recharge during
the drought of 1933-1934 approximately
corresponded to the prairie peninsula (Figure 3).

Relative Abundance

J Jit Sars 54

Dry Mesic
Moisture Gradient

Wet

Figure 2. Generalized distribution of major grass
Species across a soil moisture gradient: (1) sideoats
grama, Bouteloua curtipendula; (2) little bluestem,
Schizachyrium scoparium; (3) Indian grass, Sorghas-
trum nutans; (4) big bluestem, Andropogon geradi;
(5) switchgrass, Panicum virgatum; (6) bluejoint
grass, Calamagrostis canadensis; (7) prairie
cordgrass, Spartina pectinata. Adapted from Parrish
and Bazzaz 1982.

Figure 3. Area in which complete recharge of soil
moisture did not occur between the summer of 1933
and the summer of 1934 is shown in dark grey; light
grey indicates the area of complete recharge. From
Britton and Messenger 1970.

386 Illinois Natural History Survey Bulletin

Ecologists generally recognize that
climate is the most important factor influencing
the distribution of vegetation. However, most
ecologists believe that prairie vegetation in the
eastern United States would have largely
disappeared during the past 5,000 years had it
not been for the nearly annual burning of the
prairies by the North American Indians and the
prairie fires set by lightning (Komarek 1968).
The role of Indians in maintaining the prairies
and the reasons they burned these grasslands
have been discussed and documented by
various authors (e.g., Stewart 1951, 1956;
Curtis 1959; Pyne 1986).

Although many woody species, for
example, oaks (Quercus spp.), readily resprout
after being top-killed by fire, prairie species are
generally better adapted to burning than are
most woody plants. The adaptation that protects
grasses and forbs from fire is their annual
growth habit: the plant dies back to its under-
ground organs each year, exposing only dead
material above ground (Gleason 1922). Prairie
fires become very hot above ground and on the
surface of the soil (83 to 680 C) (Wright 1974;
Rice and Parenti 1978) but because they move
quickly and soil is a good insulator, little heat
penetrates the soil. The same adaptation that
protects prairie plants from fire also protects
them from drought and grazing. Growing
points beneath the surface of the soil permit
regrowth after intense grazing and protect
perennating organs from desiccation during
periods of drought or from fire at any time of
the year (Gleason 1922; Tainton and Mentis
1984; Anderson 1982,1990).

Grasses generally produce more biomass
annually than can be decomposed in a year.
This production of excess herbage probably
evolved in response to grazing; however, the
productivity of grasslands declines when excess
plant litter is not removed by fire or grazing
(Golley and Golley 1972). Thus, grasslands
evolved under conditions of periodic drought,
fire, and grazing and are adapted to all three
(Owen and Wiegert 1981; McNaughton 1979,
1984; Anderson 1990).

In presettlement Illinois, the vegetation
was primarily a shifting mosaic of prairie,
forest, and savanna that was largely controlled
by the frequency of fire under climatic condi-
tions that were capable of supporting any of
these vegetation types. The frequency of fire
was largely determined by topography and the

Vol. 34 Art. 4

occurrence of such natural firebreaks as
waterways and dissected landscapes. Fires
carry readily across landscapes that are level to
gently rolling, but in hilly and dissected
landscapes the spread of fire is more limited
(Wells 1970; Grimm 1984). Fire tends to carry
well uphill because rising convection currents
encourage its spread. But as fire moves down
slopes, the convection currents tend to retard it
by rising upward and working against the
downward direction of the moving fire.

The importance of waterways in deter-
mining the distribution of forest and prairie in
presettlement IIlinois was demonstrated by
Gleason (1913) through the use of the Govern-
ment Land Office Records for selected Illinois
counties. He found that prairies were more
associated with the west sides of streams and
bodies of water than with the east sides, and
forests were generally found bordering the east
sides. Gleason attributed this pattern to
prevailing westerly winds that carried fires
from west to east; the west sides of waterways,
therefore, burned more frequently than the east
sides. Forests were most abundant in presettle-
ment Illinois in the northeast Morainal Section
(Schwegman 1973) and in the three unglaciated
areas of Illinois (driftless area of Jo Daviess
and Carroll counties in northwest Illinois,
Calhoun County and portions of Pike County in
west-central Illinois, and the far southern
portion of the state) (Figure 4). In these areas,
the dissected nature of the topography and/or
the presence of waterways decreased the
frequency of fire and encouraged the growth of
forests and savannas. Similarly, the Illinoian till
plain, which is older and more dissected than
the Wisconsinan till plain, supported more
forest than the Wisconsinan till plain, espe-
cially in the southern portion (Figure 4).

The relationship between topographic
relief and vegetational patterns in Illinois has
been recently reexamined. Using a map
showing the distribution of prairies and timber
(forest and savanna) for Illinois, based on the
Government Land Office Records (Anderson
1970), and a map of the average slope range for
the state (Fehrenbacher et al. 1968), Anderson
(1991) determined the simultaneous occurrence
of slope categories and vegetation. Most of the
prairie vegetation (82.3%) occurred on land-
scapes with slopes of 2—4%; only 23.0% of the
timbered land, usually on floodplains, was
associated with this slope category. In contrast,

April 1991

77% of the timbered land occurred on sites that
had slopes greater than 4% (4—7% slope =
35.2% timber and >7% slope = 41.8% timber)
(Figure 5). Iverson (1988) also showed that
presettlement forests were positively correlated
with sloping landscapes.

The relationship between vegetational
patterns and topography is illustrated by the
presettlement vegetation of McLean County,
which is located in the Grand Prairie Division
(Schwegman 1973). That relationship is shown
in Figure 6 (Rogers and Anderson 1979). Prior
to settlement by Europeans, the county was
90% tallgrass prairie, which occupied relatively
level landscapes. Savannas and open forests
that were dominated by relatively shade-
intolerant but moderately fire-resistant oaks
(burr, Quercus macrocarpa; white, Q. alba;
and black, Q. velutina) occurred on slopes and

Driftless

Morainal Section \_

Wisconsinan

Kansan

Unglaciated

Unglaciated

Figure 4. Areal distribution of the dominant till
formations and unglaciated portions of Illinois.
Adapted from Willman and Frye 1970.

Symposium Proceedings: Our Living Heritage 387

ridges of glacial moraines. These areas were
subject to periodic fires but less frequently than
the prairies. Sheltered areas, such as ravines
and stream valleys, contained oaks and
hickories but also a high component of meso-
phytic, shade-tolerant, and fire-susceptible tree
species—elms (U/mus spp.), ashes (Fraxinus
spp.), and maples (Acer spp.).

The presettlement prairies of Illinois were
drastically altered by the influx of European
settlers who converted essentially all of the
prairie lands to agriculture. The earliest settlers
entered the unglaciated southern portion of the
state. This was a familiar landscape for these
people who were mostly hunters and trappers
from forested regions of Tennessee, Kentucky,
and West Virginia. As they migrated north-
ward, they followed the fingerlike traces of
forest along the major waterways and initially
avoided the larger tracts of prairie. For a variety
of reasons, the larger tracts of prairie were
avoided in favor of smaller tracts of prairie
adjacent to waterways and timber. The settlers
needed water for their livestock and to turn
waterwheels, and timber was needed for fuel
and building materials. In addition, the large
tracts of prairie exposed the settlers to the force
of winter storms. Timber was considered such
an important commodity on the prairie that
counties were not allowed to form as govern-
mental units until residents could demonstrate
that they had access to timber to support
development (Prince and Burnham 1908).

Ironically, some of the earliest settlers
believed that prairie soils were infertile. They
had been familiar with life in the forest and
thought that soil incapable of supporting trees
would not be productive for crops. Further-
more, turning over the thick prairie sod was an
almost insurmountable obstacle to early prairie
farmers until John Deere invented the self-
scouring steel plow in 1836. Even after settlers
had learned of the fertility of the prairie soil
and could raise large crops, many of the larger
tracts of prairie remained unsettled because the
lack of transportation to get crops to distant
markets inhibited expansion onto the prairie.
With the coming of the railroads in the
1850—1860s, however, prairies were rapidly
converted to cropland (Anderson 1970).

As the prairies were converted to an
agricultural landscape, fires, which had swept
nearly annually across the prairie in presettle-
ment times, were actively stopped by settlers

388 Illinois Natural History Survey Bulletin

who viewed them as a threat to economic
security. According to Gerhard (1857: 278),
“The first efforts to convert prairies into forest
land were usually made on the part of the
prairie adjoining to the timber. . . . three
furrows were ploughed all round the settle-
ments in order to stop the burning of the
prairies . . . ; whereupon the timber quickly
grows up.” The settlers also indirectly stopped
the fires by creating plowed fields and roads
that acted as firebreaks.

Cessation of these nearly annual prairie
conflagrations furthered the demise of the
prairies, and many of them were converted to
forests or savanna by invading tree species, the
distribution of which was no longer restricted
by periodic fires. Prairies continued to persist
along railroad rights-of-way. Railroads had
been in place before the landscape was exten-

Vol. 34 Art. 4

sively disturbed and the rights-of-way, which
usually extended for 100 feet on either side of
the track, were fenced to keep off livestock. In
addition, the rights-of-way were managed with
fire. Those fires along with many accidental
fires prevented the invasion of woody species
and exotic weeds. In the last 10 to 20 years,
however, many of the remnant prairies along
railroads have disappeared because herbicides
are used to manage rights-of-way rather than
fire. Then too, abandoned rights-of-way, which
often contained the only example of native
prairie vegetation in areas as large as a county,
have frequently been purchased by an adjacent
landowner and converted to cropland.

Within Illinois, tallgrass prairie was the
dominant grassland community. Variation in
topography, drainage patterns, and soil texture
resulted in a variety of prairie community

Figure 5. The distribution of native forest-savanna vegetation and prairie (left) compared to average slope
categories (right) in Illinois. Native prairie vegetation is shown as black; native forest-savannah vegetation ts
shown as white. A slope of 24% is shown as white, 4—7% as stripes, and >7% as black. From Anderson 1991.

April 1991

types. Hill prairies occur in scattered locations
along the generally forested bluffs of the major
river systems, especially the Illinois and
Mississippi. These prairies are relatively small,
occupying areas from less than a fraction of an
acre to as many as 12 or 13 acres. These xeric
prairies often occur on west to southwest facing
slopes and are dominated by species such as
little bluestem and sideoats grama that are
dominant components of the arid mixed-grass
and shortgrass prairies to the west of Illinois
(Evers 1955). Despite the xeric nature of these
sites, many presettlement hill prairies have
been eliminated or greatly reduced in area as a
result of the exclusion of fire and the subse-
quent encroachment of woody plants (Kilburn
and Warren 1963; Anderson 1972; Ebinger
1981; McClain 1983). Many hill prairies have
also experienced a decline in quality as a result
of grazing by cattle (Evers 1955).

Sand prairies occur on the deep Pleisto-
cene sand deposits along the Illinois River that
were laid down by glacial meltwaters during
the Woodfordian substage of the Wisconsinan
glacial advance (Willman and Frye 1970).
These coarse textured sandy soils have little
water-holding capacity and favor the growth of
plant species adapted to the droughty condi-
tions that characterize this habitat (Gleason
1907; Vestal 1913). Dominant plant species on
sand prairies include little bluestem grass, sand
lovegrass (Eragrostis trichodes), and sand
reedgrass (Calamovilfa longifolia). The sand
prairie community is more resistant to distur-
bance than the tallgrass prairie. Many agricul-
tural weeds are adapted to mesic sites and are
not effective competitors on sand prairie sites.
When weeds become established on tallgrass
prairie, however, they can prevent recoloniza-

Symposium Proceedings: Our Living Heritage 389

tion by tallgrass prairie species (Curtis 1959).
Until the expanded use of fertilizers and
irrigation, sustained agriculture had not been
possible on these droughty, low-nutrient sites
and as they were abandoned, the native sand
prairie flora frequently became reestablished.

Of the 22 million acres of tallgrass prairie
that once covered the Illinois landscape, only
about 2,300 acres of high-quality prairie remain
(White 1978). The prairie community inadver-
tently provided the incentive for its own
demise. In a grassland community, about two-
thirds of the plant mass is located beneath the
surface of the soil in the form of roots and other
underground organs. As these belowground
portions of the plant die, they decay in place
and greatly enrich the soil with organic matter.
The rich and productive soils of most of the
Midwest cornbelt, some of the most agricultur-
ally productive soils in the world, had their
genesis under prairies. Once the European
settlers learned of the fertility of the prairie soil,
had the plow that could effectively turn the sod,
and could transport their crops to distant
markets, the prairies of Illinois disappeared
quickly.

Today, however, there is growing interest
by the scientific community and the general
public in saving and restoring the prairie. The
esthetic values of prairie landscapes are being
appreciated by a growing number of persons
and the potential value of prairie plants in a
system of sustainable agriculture is drawing
attention from several sources. Efforts are
being made to develop one of the native grasses
(eastern gama grass, Tripsacum dactyloides)
into a perennial grain crop (Eisenberg 1989)
and to expand the use of warm-season native
grasses as a source of forage in combination

Tallgrass prairie (90%)
< | tree per 5 acres

Savanna (5.4%)
| tree per 5 acres
to < 19 trees per acre

Open forest
(1.8%)

> 19 trees per acre
but < 40 trees

per acre

Closed forest (3.3%)
> 40 trees per acre

Figure 6. Presettlement vegetation of McClean County, Illinois, in relation to topography about 1820.

Adapted from Anderson 1990.

390

with cool-season domestic grasses. The cool
season domestic grasses, such as orchard grass
(Dactylis glommeratus) and smooth brome
(Bromus inermis), provide forage during the
early and late (cool) portions of the growing
season. The warm-season prairie grasses, which
maximize growth in July and August, produce
a high-quality forage in the middle of the
summer when the productivity of the cool-
season species is low. As a result, cattle are
provided with abundant, good-quality forage
throughout the growing season.

It is interesting to note that such cool-
season grasses as the exotic Kentucky blue-
grass (Poa pratensis) were favored over native
grass species by the European settlers as forage
for livestock. Bluegrass provided forage a
month earlier in the spring and a month later in
the fall than the native species and was favored
for this reason (Prince and Burnham 1908).
Because the native grasses had evolved under a
system of intermittent grazing pressure, they
were eliminated when exposed to continuous
grazing. After a couple of years of continuous
grazing, native species declined, and the
Kentucky bluegrass invaded and dominated.

In Illinois, the tallgrass prairie ecosystem
is gone. Yet, the interest in preserving the
remaining remnant prairies is strong, including
the efforts of such private groups as the Grand
Prairie Friends and The Nature Conservancy
and such governmental agencies as the Illinois
Department of Conservation and the Depart-
ment of Transportation. Plantings of prairie
grasses now diversify the vegetation along
many interstate highway rights-of-way. An
increasing number of native prairie forbs, the
nongrass plants (“flowers”) of the prairie, and
prairie grasses are being sold by commercial
nurseries and seed growers. These forbs include
blazing star (Liatris spp.), purple cone flowers
(Echinacea pallida and E. purpurea), yellow
cone flower (Ratibida pinnata), and others.
These efforts ensure that future generations of
Illinoisans, like the earliest visitors to the state,
will have the opportunity to observe prairie life
and be inspired by the pleasant colors of tall
prairie grasses in the fall and shooting stars
(Dodecatheon media) and lavender phlox
(Phlox pilosa) in the spring.

Illinois Natural History Survey Bulletin

Vol. 34 Ant. 4

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Transactions of the Illinois State Academy of
Science 63(2):214-221.

ANDERSON, R.C. 1972. Prairie history, management,
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ANDERSON, R.C. 1982. An evolutionary model sum-
marizing the roles of fire, climate, and grazing
animals in the origin and maintenance of grasslands:
an end paper. Pages 297-308 in J. Estes and J.
Brunken, eds. Grasses and grasslands: systematics
and ecology. University of Oklahoma Press,
Norman.

ANDERSON R.C. 1990. The historic role of fire in the
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and L. Wallace, eds. Fire in North American tallgrass
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ANDERSON, R.C. 1991. The presettlement forests of
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Woodland Workshop (in press).

AXELROD, D. 1985. Rise of the grassland biome,
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Britton, W.A., AND A.S. MESSENGER. 1970. Com-
puted soil moisture patterns in and around the prairie
peninsula during the great drought of 1933-1934.
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Science 62(2):181—187.

Curtis, J.T. 1959. The vegetation of Wisconsin.
University of Wisconsin Press, Madison. 657 p.

EBINGER, J.E. 1981. Vegetation of glacial drift hill
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115-121.

EISENBERG, E. 1989. Back to Eden. The Atlantic
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Evers, R.A. 1955. Hill prairies of Illinois. Illinois
Natural History Survey Bulletin 26:368—446.

FEHRENBACHER, J., B. RAY, AND J. ALEXANDER.
1968. Illinois soils and factors in their development.
Pages 165-175 in R.E. Bergstrom, ed. The Quater-
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Agriculture, University of Illinois, Urbana.

GERHARD, F. 1857. Illinois as it is. Keen and Lee,
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G.eason, H.A. 1907. A botanical survey of the
Illinois River valley and region. Part 2. Pages
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biology of sand areas of Illinois. Illinois State
Laboratory of Natural History Bulletin 7.

April 1991

Geason, H.A. 1913. The relation of forest distribu-
tion and prairie fires in the middlewest. Torreya 13:
173-181.

GLEASON, H.A. 1922. Vegetational history of the
middlewest. Annals of the American Association of
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GoLLey, P.M., AND F.B. GOLLey, EDS. 1972. Papers
from a symposium on tropical ecology with an
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Grimm, E. 1984. Fire and other factors controlling
the Big Woods Vegetation of Minnesota in the mid-
nineteenth century. Ecological Monographs 54:
291-311.

Iverson, L.R. 1988. Land-use changes in Illinois,
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KivpurNn, P.D., AND D.K. WarREN. 1963.
Vegetation-soil relationships in hill prairies.
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Symposium Proceedings: Our Living Heritage 391

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131-137 in G. Clambey and R. Pemble, eds. The
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Illinois sand prairie. Illinois State Laboratory of
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Publishing Co., Lincoln, NB. 348 p.

WELLS, P. 1970. Postglacial vegetational history of
the great plains. Science 167:1574—1582.

White, J. 1978. Illinois natural areas inventory
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Illinois Natural Areas Inventory, Urbana. 426 p.

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Management 27:5—1 1.

Prairie and Savanna-restricted Insects of the Chicago Region

Ron Panzer, Northeastern Illinois University

Abstract. Numerous remnants of the presettle-
ment prairies and savannas of the Chicago
region have survived. Unfortunately, most are
very small and degraded. Nearly all are isolated
within vast expanses of human-dominated
landscape. For the past nine years, I have
surveyed grasshopper, katydid, froghopper,
leafhopper, treehopper, butterfly, and macro
moth (in part) communities on a variety of
these remnants in an attempt to gauge the status
and site size requirements of the remnant-
restricted members of these groups.

Few of the species considered in this
study (probably less than 5%) have been
extirpated. Most, perhaps as many as 80—90%,
have adapted to our degraded modern land-
scape and can be found in a variety of human-
dominated settings. Among the 10 to 20% that
are restricted to native grassland remnants,
roughly half are seemingly secure, surviving on
at least a dozen protected sites. Approximately
one-fifth of the remnant-restricted species are
known from fewer than six sites and may be
endangered within this area.

Most of the remnant-restricted insects
considered in this study have survived on
relatively small sites. One-third have been
found on sites smaller than 5 hectares. Two-
thirds have been found on sites of less than 40
hectares. More than four-fifths have been
recorded on two or more sites of less than 300
hectares. (Even sites as small as 1 hectare can
support a few restricted species.) Site size is
clearly an important determinant of butterfly
diversity on smaller remnants (1—60 ha) in this
region.

392

Prairie Birds of Illinois:

Population Response to Two Centuries of Habitat Change

James R. Herkert, Department of Ecology, Ethology, and Evolution,
University of Illinois at Urbana-Champaign

The landscape of Illinois has changed consid-
erably over the last two hundred years. The
once extensive, unbroken stretches of prairie
have given way to agricultural crops, and this
shift has had a substantial impact on the state’s
bird fauna. The purpose of this paper is
twofold: to examine how the prairie bird fauna
of Illinois has responded to changes in the
state’s landscape and to discuss how a highly
fragmented landscape may be affecting prairie
bird populations.

POPULATION STUDIES 1800-1900

Prior to European settlement, prairie occupied
approximately 8.5 million hectares in Illinois,
nearly two-thirds of the state (Anderson 1970).
The area of prairie was over 1.5 times that of
forests, which at approximately 5.5 million
hectares was the next most abundant habitat
type (Graber and Graber 1963). The composi-
tion of the presettlement bird fauna in Illinois is
not well known. Current data, however, show
that prairies support relatively low densities of
breeding birds. Bird densities in tallgrass
prairie habitat average roughly 1.8 pairs per
hectare (Cody 1985). Comparable densities for
eastern deciduous forests are 8.7 pairs per
hectare (obtained from 87 breeding bird studies
published in American Birds, volumes 37 and
38). Because of the low density of birds in
prairie habitat, Graber and Graber (1963)
estimated that only 35—40% of the presettle-
ment bird fauna of Illinois was composed of
prairie birds; forest birds, however, may have
accounted for as much as 55—60%.
Unfortunately, by the time much of the
early ornithological work was conducted within
Illinois (1850—1900), considerable losses of
prairie habitat had already occurred. By 1850
prairie habitat had been reduced to 2.1 million
hectares (Graber and Graber 1963), a reduction
of almost 75% or roughly 3.5% per year since
1810. We can, therefore, reasonably assume

that some changes in the prairie bird fauna had
occurred prior to any detailed study. Neverthe-
less, the works of Ridgway (1873, 1889, 1895)
for central and southern Illinois and Nelson
(1876) for northern Illinois can be used to
estimate prairie bird abundances in the state
prior to 1900 (Table 1).

A number of prairie bird species initially
benefited from the conversion of prairie to
farmland. Those that benefited most include the
horned lark, vesper sparrow, and greater
prairie-chicken. The increase in horned larks
and vesper sparrows was largely due to their
ability to colonize and breed in cultivated
habitats, which by 1900 had become the most
abundant habitat type in the state (Graber and
Graber 1963). The initial opening of the
prairies and forests to agriculture produced an
intermixed pattern of food and cover that was
beneficial to many species of upland game,
including the greater prairie-chicken (Weste-
meier and Edwards 1987). This shift in habitat
coupled perhaps with a reduction in the
abundance of predatory animals (due to fur
trapping and hunting) allowed the prairie-
chicken to reach a peak abundance within
Illinois of approximately 10 million birds by
1860 (Westemeier 1986; Westemeier and
Edwards 1987). Prairie-chickens started to
decline soon after reaching their peak abun-
dance. Nelson (1876) listed them as once
excessively abundant but now rather scarce in
the Chicago region and as less numerous in all
the more settled areas of the state due to egg
collection by humans, unrestricted hunting, and
loss of habitat.

POPULATION STUDIES 1900-1950

During 1906-1909, a systematic survey of the
state’s birds was conducted by Alfred Gross
and Howard Ray of the Illinois State Labora-
tory of Natural History (Forbes 1913; Forbes
and Gross 1922). These surveys provided the

393

394 Illinois Natural History Survey Bulletin

first quantitative estimates of breeding bird
populations within Illinois. A summary of the
relative abundances of the most common
grassland species encountered by Gross and
Ray in ungrazed grass, mixed-hay, and pasture
from the north and central regions of Illinois
are shown in Table 1. Gross and Ray found
bobolinks and meadowlarks (eastern and
western) to be the most common bird species,
accounting for more than 50% of all birds en-
countered in these habitats. Of the birds listed
as abundant or very common by Ridgway
(1889, 1895) and Nelson (1876), the greater
prairie-chicken, upland sandpiper, and
Henslow’s sparrow apparently experienced the
greatest declines between the mid-1800s and
the censuses of Gross and Ray. All three of
these species were uncommon or rare by 1906
(Table 1).

In the first paper addressing changes in
the bird fauna of Illinois, Ridgway (1915)
discussed changes that had taken place in the
half century preceding 1915. He cites three
prairie birds—the greater prairie-chicken,

Vol. 34 Ant. 4

upland sandpiper, and dickcissel—as experi-
encing serious declines during this period. The
greater prairie-chicken and upland sandpiper
were considered on the verge of elimination
within Illinois because of shooting and destruc-
tion of nests by dogs and cats. The dickcissel
had also dramatically declined during this
period for “unknown reasons” (Ridgway 1915).
Ridgway first noted the dickcissel’s decrease
around 1885 and stated that by 1915 this
species never reached more than one-fourth and
usually less than one-tenth its former numbers.
Coincidentally, Fretwell (1986) documented a
sevenfold increase in grazing pressure between
1870 and 1884 on the dickcissel’s primary
wintering grounds in Venezuela, a factor that
he believed could significantly affect winter
resources and, in turn, dickcissel numbers.

POPULATION STUDIES 1950-1989

In 1956-1958, the census routes of Gross and
Ray were repeated by Graber and Graber
(1963) of the Illinois Natural History Survey

Table 1. Relative abundance of prairie birds within Illinois 1850-1989.

1906-19097 1956-1958° 1987-1989 USFWS*
Species Prior to 1900! %e Rank %o Rank % Rank % change
Eastern meadowlark® Abundant 29 2 20.0 2 11.8 2 -67.0
Dickcissel Abundant 13.1 3 8.7 = qed 5 —46.7
Grasshopper sparrow Abundant 5.9 5 a5) 6 8.6 - 56.0
Bobolink Abundant 25.8 1 OF, 3 114 3 —90.4
Henslow's sparrow Abundant <1.0 15 <1.0 14 1.6 12 7
Red-winged blackbird Very common Le - 36.2 1 26.8 ] -18.8
Greater prairie-chicken Very common <1.0 13 0.0 16 0.0 16 is
Upland sandpiper Very common DS 9 <1.0 12 <1.0 13 -16.8
Vesper sparrow Common 1.3 11 1.4 10 <1.0 15 +12.1
Horned lark Common 4.9 6 4.8 7 <1.0 14 0.0
Field sparrow Common 4.0 7 2.9 9 5.6 7 52.6
Song sparrow Common 2.6 8 1.0 11 3.3 10 29.3
Savannah sparrow Common 2.3 10 5.8 5 35 9 —58.9
American goldfinch Common 12 12 3.1 8 4.7 8 42.8
Common yellowthroat © Common <1.0 14 <1.0 15 5.8 6 —88
Sedge wren Common <1.0 16 <1.0 13 2.8 11 —22.5

‘Relative abundance prior to 1900 based on the works of Nelson (1876) and Ridgway (1873, 1889, 1895).

Relative abundance 1906-1909 based on the censuses of Gross and Ray from approximately 380 ha of ungrazed grass,
mixed-hay, and pasture, located in northern and central Illinois (Forbes 1913; Forbes and Gross 1922).

‘Relative abundance 1956-1958 based on the censuses of Graber and Graber (1963) from approximately 290 ha of ungrazed
grass, mixed-hay, and pasture, located in northern and central Illinois.

‘Relative abundance from the present study (1987-1989) based on censuses of approximately 400 ha of ungrazed prairie and
agricultural grasslands in northeastern and east-central Illinois.

‘Estimated population change within Illinois between 1967-1989 based on United States Fish and Wildlife Service's
breeding bird survey (USFWS, unpublished data).

°For 1906-1909 and 1956-1958, relative abundance estimates are for eastern and westem meadowlarks combined.

*Present on too few routes for accurate trend analysis.

April 1991

(Table 1). The Grabers believed that the red-
winged blackbird, horned lark, and dickcissel
had shown large statewide population increases
between 1909 and 1956.

Red-winged blackbird numbers had
almost doubled since the earlier censuses of
Gross and Ray due to the ability of this species
to invade nearly all terrestrial habitats within
the state (Graber and Graber 1963). Ridgway
(1889) noted that although very common, the
nests of red-winged blackbirds were always in
or in very close proximity to a swamp or
marsh. Gross and Ray, however, found red-
winged blackbirds in all the grassland habitats
they censused in 1906-1909, although 60% of
the state’s population of these birds still nested
in marshes (Graber and Graber 1963). From
1909 to 1956, red-winged blackbird densities
within grassland habitats in Illinois increased
nearly tenfold. The species had become far
more common in grasslands than in marshes,
with individuals inhabiting marshes accounting
for less than 3% of the state’s population
(Graber and Graber 1963).

The statewide increase in horned larks
between 1909 and 1956 corresponded to their
shift from primarily grassland to cultivated
habitats, especially row-cropped fields. This
switch from a rapidly declining to a rapidly
increasing habitat greatly benefited the horned
lark, which Graber and Graber (1963) recog-
nized as the species that had increased most
dramatically between 1909 and 1956. The
Grabers attributed the dickcissel’s statewide
increase to an expansion in acreage of agricul-
turally disturbed grasslands, a type of habitat
that this species may prefer over true prairie
(Kendeigh 1941; Graber and Graber 1963;
Zimmerman 1971). Most species of prairie
birds, however, had shown either little or no
statewide population change between 1909 and
1956 (Graber and Graber 1963). The bobolink,
song sparrow, and savannah sparrow showed
slight increases, the upland sandpiper and field
sparrow slight decreases, and the vesper
sparrow, grasshopper sparrow, and American
goldfinch no change.

Between 1987 and 1990, I conducted
research on the breeding birds of Illinois
grasslands; however, my field methods differed
from those used by Gross and Ray and the
Grabers and direct comparisons are therefore
not possible (see Herkert 1991 and Graber and
Graber 1963 for descriptions of methods).

Symposium Proceedings: Our Living Heritage 395

Nevertheless, a comparison of relative abun-
dances of these species indicates that the
current composition of grassland bird fauna is
probably very similar to that of the late 1950s
(Table 1). Red-winged blackbirds remain the
most common species, outnumbering the next
most abundant species, the eastern meadow-
lark, by more than two to one. In fact, four of
the five most abundant species are the same in
my censuses and in those of Graber and Graber
(Table 1).

An estimate of how prairie bird numbers
have changed since the Grabers’ census can be
obtained from data collected by the United
States Fish and Wildlife Service’s cooperative
breeding bird survey (unpublished data). These
data from Illinois for 1967-1989 show that
nearly all prairie bird species have experienced
population declines during this 23-year interval
(Table 1). Some of the formerly most abundant
prairie bird species, for example, the bobolink,
have shown declines as high as 90% during this
period. The causes of these recent population
declines are not well understood but probably
are a consequence of continued loss of grass-
land habitat within Illinois.

Although the initial loss of prairie habitat
within Illinois was rapid and extensive, the
reduction of prairie habitat has continued in
recent decades. By 1978, less than 1,000
hectares of high-quality prairie remained in the
state (Schwegman 1983). The loss of prairie
habitat was originally offset by the creation of
secondary grasslands such as hayfields and
pastures, habitats which the majority of prairie
birds found suitable for breeding (Graber and
Graber 1963). In fact, none of the characteristic
birds of the eastern tallgrass prairie region are
considered endemic to prairie habitat (Risser et
al. 1981). Acreage of these secondary grassland
habitats, however, has also recently declined.
For example, the amount of hay within Illinois
was reduced by more than half, from 850,000
to 400,000 hectares, between 1960 and 1989
(Illinois Agricultural Statistics Service 1988,
1989). The amount of pasture within Illinois
has also been greatly reduced, with pasture
occupying only 607,000 hectares in 1987 (U.S.
Department of Commerce Bureau of the
Census 1989) compared with 2.5 million
hectares in 1906. The continued loss of both
native and agricultural grassland habitats in
Illinois has contributed to an increasingly
fragmented landscape.

396 Illinois Natural History Survey Bulletin

HABITAT FRAGMENTATION

The process of habitat fragmentation sets off

a series of events that can ultimately have a
major effect on breeding bird communities.
Changes associated with increased fragmenta-
tion include a decrease in the total amount of
habitat, a decrease in the average size of habitat
patches, increased patch isolation, and an
increase in the ratio of edge to interior habitat,
all of which may have important consequences
for breeding birds (Wiens 1989).

The most important consequence of
habitat fragmentation is the loss of large
amounts of habitat and the resulting losses of
individuals, local populations, and possibly
even species. Surprisingly, only three species
of prairie birds have been extirpated from
Illinois despite the extensive loss of prairie
habitat (Table 2). Bowles et al. (1980) origi-
nally listed four species as extirpated from
Illinois, but the sandhill crane has returned to
the state as a breeding species (Kleen 1988).
The remaining three species (sharp-tailed
grouse, swallow-tailed kite, and whooping
crane) were extirpated prior to or very shortly
after 1900 (Bowles et al. 1980). Another 13
prairie bird species are now considered to be
threatened or endangered within Illinois (Table
2), primarily as a direct result of extensive
habitat loss. A number of these endangered and
threatened species may be on the verge of
extirpation within Illinois. The greater prairie-
chicken, for example, once one of our most
abundant prairie birds, now has a statewide
population of less than 100 individuals
(R. Westemeier, pers. comm.).

The reduction of the average patch size
that accompanies habitat fragmentation also has
serious consequences for breeding birds. Small
patches may be too small to meet the minimum
territory requirements for a species or may lack
essential resources necessary for the establish-
ment of populations (Diamond 1975). The
responses of individual species to reductions in
patch size are variable, but nearly all bird
species exhibit a minimum area threshold
below which they never occur (e.g., Lynch and
Whigham 1984; Hayden et al. 1985; Robbins et
al. 1989). Six prairie bird species were never
encountered during my research within Illinois
on areas of less than 10 hectares (Table 3),
despite the fact that the average territory for
four of these species (bobolink, savannah
sparrow, grasshopper sparrow, and Henslow’s

Vol. 34 Art. 4

sparrow) is typically less than 2.5 hectares
(Wiens 1969). Many prairie bird species avoid
small areas, and small grasslands have been
shown to support impoverished breeding bird
faunas (Samson 1980; Howe et al. 1985:
Herkert 1991). The number of breeding bird
species in grassland fragments is strongly
related to fragment size, with large fragments
supporting significantly more species than
small fragments (Samson 1980; Herkert 1991).
In addition, small habitat patches generally
support small numbers of individuals, thus
greatly increasing the influence of stochastic
events on population demography. As a result,
small isolated bird populations have been
shown to exhibit relatively high turnover rates
(e.g., Diamond 1969; Diamond and May 1977;
Morse 1977) and therefore a higher probability
of local population extinction.

In Illinois, the natural areas inventory
(1975-1978) identified only 253 remnants,
totaling 950 hectares, of high-quality prairie

Table 2. Extirpated, endangered, and threatened
birds of Illinois prairies (from Bowles et al. 1980).

Endangered Threatened
American bittern Loggerhead shrike
Yellow rail Henslow's sparrow
Black rail Brewer's blackbird

Bachman's sparrow
Greater prairie-chicken
Swainson's hawk
Short-eared owl
Northern harrier
Upland sandpiper
Sandhill crane

Extirpated
Sharp-tailed grouse
Whooping crane
Swallow-tailed kite

Table 3. Minimum areas of encounter for 17 grass-
land bird species from 24 grassland fragments
located in northeastern and east-central Illinois
(1987-1989). Grasslands ranged from 0.5 to 650
hectares.

<10 hectares 10—30 hectares

Field sparrow Bobolink
American goldfinch Savannah sparrow
Song sparrow Grasshopper sparrow

Dickcissel
Ring-necked pheasant
Sedge wren

Common yellowthroat
Red-winged blackbird
Northern bobwhite
Eastern meadowlark
Vesper sparrow

Henslow’s sparrow

>30 hectares
Upland sandpiper
Northem harrier

April 1991

within the state (Schwegman 1983). The
majority of these remnants were small, most
less than 20 hectares, and would therefore be
expected to support very few, if any, prairie
bird species. Grasslands of 100 hectares or
more may be necessary to support just five
prairie interior species (Herkert 1991).

Increases in patch isolation can also
increase the probability of local population
extinctions due to decreased immigration rates.
Island biogeography theory predicts that
immigration rates will be affected by both
patch isolation and size, with the lowest
immigration rates occurring on patches that are
small and well isolated from a colonizing
source (MacArthur and Wilson 1967). Whether
mainland fragments act as true islands with
respect to immigration, however, is open to
question because mainland fragments are not
surrounded by totally inhospitable habitat as
are true islands and therefore might not show
immigration rates that are strongly dependent
on patch isolation. A number of studies
conducted in the eastern deciduous forests of
North America have demonstrated that isola-
tion does have a significant effect on species
richness within forest fragments (Robbins
1980; Howe 1984; Lynch and Whigham 1984;
Askins et al. 1987). Researchers working in
forests on other continents, however, have
found no evidence supporting isolation as a
significant factor affecting species richness
within fragments (Kitchener et al. 1982; Howe
1984; Opdam et al. 1985). The effects of
isolation on immigration rates in midwestern
grasslands have not been studied to date.

Harris (1984) points out that island
biogeography theory assumes that islands
always have a mainland source pool for
immigration; for terrestrial fragments, however,
the “mainland” source may be lost as a result of
the fragmentation process. In this case, the
recolonization of mainland fragments must
occur between habitat patches. The integrity of
the whole system would then depend on the
existence of areas large enough to produce
enough surplus individuals to provide dispers-
ers as well as maintain stable populations
within a particular preserve.

Another consequence of habitat fragmen-
tation is an increase in the ratio of edge to
interior habitat as patch size decreases (Butcher
et al. 1981; Temple 1986). This increase may
result in the loss of species that require interior
habitats and an increase in the abundance of

Symposium Proceedings: Our Living Heritage

397

edge species (Whitcomb et al. 1981; Ambuel
and Temple 1983; Temple 1986). Small
grasslands are usually dominated by such
nonprairie species as red-winged blackbirds
and common yellowthroats and support few
prairie interior bird species (Herkert 1991).
Moreover, the increase in the ratio of edge to
interior habitat may lead to lower reproductive
success for nesting grassland birds. Levels of
both nest predation and parasitism have been
shown to be higher in edge habitats than in
grassland interiors, especially if the edge is a
field—woodland or field—shrubland border (Best
1978; Gates and Gysel 1978; Johnson and
Temple 1986, 1990; Burger 1988).

Finally, we must remember that loss of
prairie and grassland habitat in Illinois, and
throughout the Midwest, affects birds primarily
during the breeding season. The majority of
prairie bird species are migratory and spend
only a fraction of any given year on the
breeding grounds. Similar alterations of
wintering and possibly migratory habitat may
also significantly affect these bird species. The
degree to which events off of the breeding
grounds affect prairie birds are not well known.
For such species as the dickcissel, however,
events on the wintering grounds and migratory
routes may be the most important factors
affecting distribution and abundance patterns
on the breeding grounds in the Midwest
(Fretwell 1986). The fact that processes
operating outside the boundaries of Illinois
affect bird populations within the state does not
excuse us from being concerned about events
occurring within Illinois, but rather should alert
us to the year-round needs of these species. If
conservation efforts to preserve prairie birds
are to succeed, management efforts must
address not only processes operating on the
breeding grounds within Illinois but the
migratory and wintering needs of these species
as well.

ACKNOWLEDGMENTS

This research was funded in part by the

S. Charles Kendeigh Memorial Fund, Illinois
Nongame Wildlife Fund, Sigma Xi, and the
University of Illinois at Urbana-Champaign.
G.C. Sanderson, S.K. Robinson, R.E. Warner,
and two anonymous reviewers provided helpful
comments and editorial assistance on previous
drafts of the manuscript.

398 Illinois Natural History Survey Bulletin

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Session Three: Wetlands

What would the world be, once bereft

Of wet and wildness? Let them be left,

O let them be left, wildness and wet;

Long live the weeds and the wilderness yet.

—Gerard Manley Hopkins

While most Illinois residents may not consider
their state to be particularly wet, early settlers
had a very different impression. Writing in
1833, the year Chicago was incorporated as a
village, Colbee Benton observed that Chicago
“stands on the highest part of the prairie, and in
the wet part of the season the water is so deep
that it is necessary to wade from the town for
some miles to gain the dry prairie. Notwith-
standing the water standing on the prairie and
the low, marshy places, and the dead-looking
river, it is considered a healthy place.”

The retreat of the glaciers left numerous
large and small streams with many associated
wet areas. Much of northeastern Illinois had
abundant diverse wetlands, and central Illinois
was a montage of wet prairies and marshes.
Extensive tracts of tupelo-cypress swamps
could be found in the far southern part of the
state.

Wetlands are diverse and complex places.
The most common wetlands in Illinois are
marshes and sedge meadows, although ponds,
fens, seeps, wet prairies, swamps, and bogs are
also present. Marshes form where water is
above the soil surface for all or nearly all of the
year—along the margins of ponds, lakes, or
rivers, in places sheltered from strong currents
and waves. Sedge meadows are usually associ-
ated with fens. Here the water level is near or
just below the surface most of the year, and this
habitat often merges into marshes as the water
depth increases. The surface of the vegetation
hides countless tussocks or humps formed by
the tussock sedge, and these vary in height from
a few inches to over a foot. The terms bog and
fen are often used inconsistently, even inter-
changeably, and considerable confusion has
been the result. In general, bogs are acidic and
poor in minerals, with most of the water coming
from rainfall and surface runoff and most of the
new peat developing from sphagnum moss.
Fens range from acidic to alkaline and are rich
in minerals; much of the water comes from

400

groundwater that has percolated through
calcareous bedrock or gravel. Peat is produced
primarily by sedges and grasses. Seeps are
characterized by groundwater that has reached
the surface in a diffuse rather than a concen-
trated flow. Seeps form when groundwater that
has percolated down through porous sand or
gravel reaches a layer of impermeable material
and flows outward, usually at the base of a
bluff or ravine. Swamps are areas where the
soil is saturated or covered with surface water
for most of the growing season; woody
vegetation dominates.

What was formerly looked upon as
sources of disease and pestilence, “sacred to
the ague and fever,” are currently viewed in a
new light. The importance of wetlands is only
now being realized: they store runoff after
major rains and slowly release it; they filter silt
and pollutants from water; and they are tre-
mendously productive, providing habitat for a
diversity of plants and animals.

Illinois originally had an estimated 8
million acres of wetlands. Since Illinois
became a state in 1818, more than 95% of
these have been drained with a concomitant
loss in the natural processes that wetlands
provide. High-quality wetlands that reflect
presettlement conditions are exceedingly rare
today; only about 6,000 acres remain.

The papers presented at this session re-
viewed the state of our wetlands, documenting
what has been lost as well as what must be
restored or preserved. Particular attention was
given to the plants and animals that depend on
the unique habitats of wetlands.

Aquatic and Wetland Plants of Illinois

John E. Schwegman, Natural Heritage Division,
Illinois Department of Conservation

Abstract. Over 100 of the 172 families of
vascular plants growing without cultivation in
Illinois have species adapted to aquatic or moist
soil habitats. These wetland plants range from
ferns and their allies to conifers to flowering
plants. Growth forms include herbs, shrubs, and
trees, any of which may function as the domi-
nant species of a plant community or as minor
components. Some important wetland plant
families in Illinois are the sedge family
(Cyperaceae), grass family (Poaceae), pond-
weed family (Potamogetonaceae), duckweed
family (Lemnaceae), smartweed family
(Polygonaceae), and sunflower family (Aster-
aceae). In providing for their own growth and
reproduction, these plants make up the vegeta-
tion component of wetlands and provide much
of the food, nesting cover, and escape cover for
wetland animals.

Common aquatic and emergent species of
wetland communities in Illinois include
coontail (Ceratophyllum demersum) beneath
the surface of calm waters, duckweeds (Lemna
sp.) floating on the surface, bulrushes (Scirpus
sp.) and cattail (Typha latifolia) in marshes,
buttonbush (Cephalanthus occidentalis) in
shrub swamps, and bald cypress (Taxodium
distichum) and water tupelo (Nyssa aquatica)
in wooded swamps. A wider variety of species
occupy moist soil communities as opposed to
aquatic communities.

401

Breeding Biology and Larval Life History of Four Species of
Ambystoma (Amphibia: Caudata) in East-central Illinois

Michael A. Morris, Cuivre Island Field Station, Western Illinois University

Abstract. Temporary aquatic habitats, whether
roadside ditches, flooded fields, or woodland
ponds are essential in maintaining the biodiver-
sity of Illinois. Nineteen species of Illinois
amphibians (50% of the state’s species) depend
on such habitats for breeding. Two species of
reptiles breed in those habitats, and 8 to 10
more use them as foraging areas. In addition,
these temporary aquatic habitats are important
for many invertebrate species.

Kickapoo State Park, located in Vermil-
ion County, Illinois, provides just such tempo-
rary aquatic habitats, and this paper records my
observations of the breeding biology and larval
history of four species of salamanders, genus
Ambystoma (Amphibia: Caudata) in that setting
from 1973-1984.

Ambystoma opacum migrated to the dry
beds of two vernal hilltop ponds at Kickapoo
State Park in late September or October. The
females oviposited under the mat of leaf litter
that covered the pond beds and abandoned the
eggs in late fall. Ambystoma platineum, A.
texanum, A. maculatum, and A. platineum X A.
texanum hybrids migrated to the ponds under
stimulus of rains in February and March,
provided groundwater was sufficient to fill the
ponds to a depth of at least 25 cm. Ambystoma
maculatum migrated 3—7 days later than the

other spring-breeding species. In years when no

standing water was present in the ponds, spring
migration was prolonged or involved few
animals. Ambystoma texanum and A.
maculatum males deposited beds of spermato-
phores in different locations on the pond
bottoms. The gynogenetic A. platineum used
sperm from the A. fexanum spermatophores to
initiate cleavage of their eggs, and fertilization
occasionally occurred. Ambystoma platineum
and A. texanum laid eggs in water less than 30
centimeters deep; A. maculatum laid eggs in
water at least as deep as 30 centimeters.
Ambystoma opacum larvae hatched
within 24 hours after the ponds filled in the

402

spring. Eggs of the other species hatched in
3-6 weeks. Larvae grew little for 2 weeks and
then grew rapidly for about 1.5 months. Little
further growth occurred before transformation.

Larvae usually transformed in late May
(A. opacum) or late June (the other species).
Ambystoma opacum larvae were always able to
transform, but in most years the ponds dried
before most, if not all, of the larvae of the other
species could transform. Larvae are opportunis-
tic feeders, and their food included volvocids,
ostracods, branchiopods, annelids, insects, and
in the case of A. opacum, the larvae of other
salamanders.

Ecological Integrity of Two Southern Illinois Wetlands

M. Ann Phillippi, Department of Zoology, Southern Illinois University at Carbondale

Palustrine and riverine wetlands in Illinois are
increasingly rare ecosystems. Unfortunately,
the declining wetland habitat in Illinois is not
an isolated phenomenon (Mitsch and Gosselink
1986; Illinois Department of Conservation
1988). Wetlands across this country are in
jeopardy due to drainage for a variety of human
endeavors, primarily agriculture, or to the
associated and chronic but less dramatic threat,
soil erosion.

The presettlement area of wetlands in this
country is difficult to ascertain, and estimates
vary from 51 to 87 million hectares (Greeson et
al. 1979). The rapidity with which our wetlands
disappeared is difficult to comprehend. By the
early 1950s, 35% of the wetlands in this
country had already been drained. Federal
bureaucracies were given unbridled authority to
drain any wetland deemed a nuisance. In the
eastern United States, the U.S. Army Corps of
Engineers and the USDA Soil Conservation
Service played major roles in the destruction of
wetlands. The Army Corps destroyed large
wetlands while the Soil Conservation Service
destroyed smaller ones. Drainage tiles were
installed throughout wetlands and quit dis-
charging only when no more water was left to
drain. Large and small ditches were dug to
expedite drainage and are dramatically illus-
trated on most topographic maps of southern
Illinois. Many of the largest ditches were given
quaint yet telling names, for example, Post
Creek Cutoff, which was dug in the early 1900s
and continues to disrupt the natural hydrologi-
cal dynamics of the wetlands along the Cache
River of southern Illinois. Smaller ditches
generally remain unnamed, such as the one dug
in an as yet incomplete effort to drain Lovets
Pond, a remnant of the once vast Mississippi
River floodplain wetlands of southern Illinois.
That ditch was most likely dug overnight
during the fall of 1986. From 1950 to 1970
another 8.5% of the nation’s wetlands were
lost, approximately 186,000 hectares per year

over the twenty-year period and an area almost
twice the size of the Shawnee National Forest.

Most (95%) of the wetlands in the United
States are inland and those are incredibly
diverse, ranging from the upland, subalpine
swamp-meadows of Yosemite to the lowland
pitcher-plant bogs of southern Alabama. Of all
wetland types (see Cowardin et al. 1979), none
is more threatened than the emergent wetlands,
those characterized by erect, rooted, herbaceous
hydrophytes (excluding mosses and lichens), or
the forested wetlands, those characterized by
woody vegetation at least 6 meters tall. The
former is found in Lovets Pond, and the latter
along the Cache River. Nationwide these two
wetland types disappeared at a rate approaching
10% each year from 1950 to 1970. This rate
has diminished but not nearly enough.

Illinois has the regrettable distinction of
having lost more of its wetlands than most
other states, and only 5% of our original
wetlands are left. Obviously, Illinois needs to
preserve all of its remaining wetlands. To do so
would provide greater assurance that the state's
biodiversity would not decrease to exclude
even fairly common but uncelebrated species
like the crawling water beetles (Peltodytes and
Haliplus spp.). One cannot be optimistic about
future preservation efforts because federal and
state laws and their implementation are “too
little too late” to prevent even state agencies
from destroying wetland habitat. To illustrate,
the Illinois Department of Conservation is
currently entertaining a proposal to destroy an
old-growth bottomland forest wetland in
Horseshoe Lake Conservation Area in Alexan-
der County. Public opposition to the project
may prevail, but current law and regulation
would make that destruction legal.

An immediate response is essential. We
need to identify and prioritize the Illinois
wetlands in greatest jeopardy, a task not easily
accomplished. Many practical and theoretical
questions must be answered in the process, for

403

404 Illinois Natural History Survey Bulletin

example, “What size do wetlands need to be to
assure their integrity and to preserve maximum
biodiversity?” Given present understanding and
adequate financial resources, the best answer is
to preserve the largest areas possible. In
Illinois, however, most of the remaining
wetlands are small, isolated islands such as
Lovets Pond. Small as these are, they cannot be
ignored, and we cannot allow ecological theory
to be used as an excuse for not preserving or
protecting them. If we accept that only large,
nearly pristine areas should be placed on a
priority list, we assure further decreases in the
state’s biodiversity because small wetlands do
harbor diverse communities, and in many cases
those communities appear to be stable. In fact,
small wetlands like Lovets Pond may presently
have greater ecological integrity than larger,
heavily silted ones like those along the Cache
River. By ecological integrity I mean the
relative disparity between the abundance and
diversity of the aquatic fauna in a given system
relative to that which could reasonably be
expected to occur in the same system if it were
undisturbed. A close look at the macro-
invertebrate communities of Lovets Pond and
the Cache River wetlands (Figure 1) provides
evidence for this contention. Acknowledging
the value of small wetlands does not of course
mean that we should not fight for the greatest
protection possible for larger areas such as the
Cache wetlands. Although these areas may be
seriously compromised, they nevertheless
contain pockets of diversity that might serve as
epicenters of re-invasion for an entire area if
allowed to do so.

Lovets Pond was once part of a wetland
system that covered a large area of the Missis-
sippi River bottoms of southern Illinois

Lovets
Pond

Figure 1. Location of Lovets Pond and the Cache
River, the two Southern Illinois sites in this study.

Vol. 34 Ant. 4

(Jackson County and others). Now, this once
vast ecosystem is reduced to a 16-ha remnant
that is surrounded by a lowland forest that
increases its size to 65 ha. This island is totally
enclosed by intensive agriculture. When I
began to investigate the ecological integrity of
Lovets Pond, I shared the bias of many
biologists who are convinced that preserving
small areas does not protect enough biodiver-
sity to justify the cost. This contention may be
true for large organisms but what about small
ones? In long- and short-term scenarios, many
species not in need of large areas may perhaps
be protected within small, isolated systems.

The Cache River wetlands were also once
part of a much larger system (114,000 ha).
Only 1% of this vast wetland complex remains,
with Heron Pond, a beautiful state nature
preserve, the best-known area. At the present
time, about 14,000 ha are being considered for
inclusion in the proposed Cypress Creek
National Wildlife Refuge. The area is an
important wintering ground for migrating
waterfowl and contains other unique features,
including several bald cypress trees over 1,000
years old that represent the oldest living
organisms east of the Mississippi River.
Agricultural activity occurs throughout the area
and forms the borders of most of the remaining
wetlands.

SITE DESCRIPTIONS AND METHODS

Lovets Pond. The investigation of the macro-
invertebrate communities of Lovets Pond
during 1986 focused on two questions (Phil-
lippi and Peterson 1986). Are the communities
diverse and distinct from one another? And if
so, are the communities distributed to corre-
spond to the vascular plant communities?
Because vascular plants are the major substrate
for the attachment of nonbenthic macroinverte-
brates, distinctiveness among the macroinverte-
brate communities might well be realized along
a gradient similar to that observed for the
vascular plants.

In order to answer these questions, one
site was selected for investigation in each of the
four major plant communities: open pond,
shrub swamp, true swamp, and marsh (Figures
2-5). During 1986, these communities were
connected by water for varying amounts of
time. The open pond and the shrub swamp were
connected the longest, and the true swamp was

April 1991

connected to the previous two for a shorter
period. The marsh was isolated from the other
three for most of the year. The open pond
community is edged with buttonbush (Ceph-
alanthus occidentalis), and by early summer
the surface is almost totally covered with
yellow pond lily (Nuphar luteum). Water in the
open pond community was about |—2 m deep.
The shrub swamp community surrounds the
open pond and is dominated by an impenetrable
thicket of buttonbush with a few black willows
(Salix nigra) scattered throughout. Thick stands
of lizard’s-tail (Saururus cernuus) occur along
its edge. In general, 20-30 cm of water covered
this community during the winter and spring.

Symposium Proceedings: Our Living Heritage 405

The true swamp is fully forested with a variety
of tree species, including pumpkin ash (Frax-
inus profunda), red maple (Acer rubrum), and
water locust (Gleditsia aquatica). Water
covered the forest floor (10—12 cm) only during
the winter. The marsh, dominated by graminoid
plants, is the smallest (1 ha) and most isolated
of the four communities. It is maintained by
periodic fires set by farmers to prevent the
lowland forest from encroaching onto their
fields. The amount of silt covering the bottom
of each of the four communities was minimal.
Two unit-effort dipnet samples of the
macroinvertebrate community were taken from
each plant community on six dates at four- to

Figure 2. Open pond community of Lovets Pond in
mid-April 1986. The thick growth of yellow pond
lily (Nuphar luteum) obscures the coontail (Cerato-
phyllum demersum) and pondweed (Potamogeton
spp.) that are scattered throughout. Photo by author.

Figure 3. Shrub swamp community of Lovets Pond
in mid-June 1986. The almost impenetrable growth
of buttonbush (Cephalanthus occidentalis) in the
background is surrounded primarily by lizard’s tail
(Saururus cernuus). Photo by author.

Figure 4. True swamp community of Lovets Pond in
mid-June 1986. New growth of arrow arum
(Peltandra virginica), foreground, covers the lowest
points in this community. A variety of tree species
are seen in the background, including pumpkin ash
(Fraxinus profunda), water locust (Gleditsia
aquatica), and red maple (Acer rubrum). Photo by
author.

Figure 5. Marsh community of Lovets Pond in mid-
May 1986. Such graminoid plants as bur reed
(Sparganium eurycarpum), giant bulrush (Scirpus
tabernaemontanii), and common cattail (Typha
latifolia) surround the marsh edge. Duckweeds
(Spirodela spp. and Lemna spp.), water meal
(Wolffia sp.), and sponge plant (Limnobium spongia)
cover the surface by summer. Photo by author.

406 Illinois Natural History Survey Bulletin

six-week intervals, January through June 1986.
Samples were preserved and later sorted and
identified to the lowest practical taxon.

Cache River and Wetlands. During the
summer of 1986 a team of biologists (Phillippi
et al. 1986) surveyed the aquatic fauna at 23
sites within the Cache River drainage (Figure
6). Two dipnet samples were taken from a
representative portion of each of the sites and
the organisms sorted and identified to the
lowest practical taxon.

Figure 6. Large bald cypress (Taxodium distichum)
along the Cache River and its wetlands provide a
major attraction for canoeists. Photo by Marti
Crothers.

Vol. 34 Art. 4

RESULTS AND DISCUSSION

Lovets Pond. The true swamp and marsh com-
munities of Lovets Pond contained the highest
number of macroinvertebrate taxa; the lowest
number was found in the open pond (Table 1).
Samples taken from the true swamp and shrub
swamp communities yielded the largest number
of individuals; once again, the open pond
yielded the lowest number (Table 1).

The number of taxa and individuals in
each community fluctuated in a roughly similar
fashion across the seasons; however, no pattern
within or across the four communities in regard
to the diversity (H’) of macroinvertebrates was
discernible (Figure 7). No single plant commu-
nity always harbored the highest or lowest
species diversity. Even so, the four plant
communities contained distinct macroinverte-
brate assemblages, at least qualitatively, and
this distinction was demonstrated using
Jaccard’s similarity coefficients and group
average clustering (Figure 8). Cluster 1 is
predominated by the shrub swamp macro-
invertebrate community, cluster 2 by the true
swamp, and cluster 4 by the open pond
community. The macroinvertebrate community
inhabiting the marsh is indistinct from those of
the other three communities even though the
marsh is the most isolated of the four commu-
nities. These data suggest that this small
wetland harbors distinct and diverse macro-
invertebrate communities—communities that
are known to be dramatically affected by
human-caused changes in substrate and water
quality (Greeson et al. 1979). From the
practical viewpoint of conservation biology, the
ecological integrity of Lovets Pond can be
considered good and thus worthy of protection.

Cache River and Wetlands. Approxi-
mately 230 aquatic and semiaquatic macro-
invertebrate taxa were collected from the 23
sites. The number of taxa and individuals at

Table 1. Total number of taxa and individuals for the four major plant communities of Lovets Pond. Ranges

are given in parentheses.

Open pond Shrub swamp True swamp Marsh
Total number of taxa 37 52 58 59
(Number per sample) (8-20) (14-30) (10-30) (11-26)
Total number of individuals 1,042 4,034 4.807 2,200
(Number per sample) (113-241) (526-982) (257-1,259) (113-769)

April 1991

each site ranged from 21—66 and 212—2,735,
respectively. Only 7% (17 taxa) were found at
10 or more sites. Of those 17 taxa, 6 were
crustaceans (aquatic sowbugs, sideswimmers,
shrimps, and crayfishes) and 6 were surface or
water-column dwelling beetles (Coleoptera) or
bugs (Heteroptera). Over 20,500 individuals
were examined, excluding those taken from
qualitative samples. The clubtail dragonfly
(Ariogomphus maxwelli) was observed and/or
collected at 4 of 23 sites. This species was
known from only a few Gulf Coast states until
June of 1985 when a single adult male was
collected at Mermet Lake in Massac County,
Illinois. Thus, the Cache population may be the
only viable one in the state. Sampling also
yielded such rare to uncommon bugs as the
water scorpion (Nepa apiculata) and such
common but hard to collect bugs as the marsh
treader or water measurer (Hydrometra
martini). In the sites most disturbed with a
heavy silt load, at least a few surface-dwelling
insects (for example, Gerris marginatus and
Trepobates spp.) were found. Gerris margi-
natus is perhaps the most common strider in the
Cache system.

To assess the ecological integrity of the
various Cache sites, species diversity measure-
ments (H’) were calculated and can be com-
pared with those found at Lovets Pond. Four
sites have a relatively high species diversity
(0.898—1.131): the Cache River at Highway 37,
Snake Hole, Eagle Pond, and Long Reach. The
Cache River at Highway 37 is a highly dis-
turbed site. The north bank has been cleared
and a levee built. The channel has been dredged
and carries a very heavy silt load. Long Reach
is also a heavily silted portion of the main
channel. Snake Hole is a well-shaded pond
located at the base of a rocky-boulder cliff in an
area known as Little Black Slough. This state-
owned site is generally the least silted of any of
the Cache wetlands. Eagle Pond, also heavily
silted, is a popular canoeing destination
because of its picturesque cypress knees and
buttonbush thickets. Sites with moderate
macroinvertebrate species diversity
(0.65 1—0.834) are heavily silted, including
Wildcat Bluff/Watson Pond and Short Reach,
both owned by the Illinois Department of
Conservation. The other 5 sites with moderate
diversity are privately owned. The remaining
12 sites have low species diversities

Symposium Proceedings: Our Living Heritage

407

ef Pond
1.0—— im Shrub swamp

|_| True swamp

Marsh

0.8

0.6

0.4

0.0

Jan Feb Mar Apr Jun

Sampling date

May

Figure 7. Shannon diversity (H’) values (N=2) for
the macroinvertebrate communities inhabiting the
four major plant communities of Lovets Pond.

open pond {
shrub swamp {
Ca marsh —
true swamp
shrub swamp4
true swamp 4
Cy L
marsh 4
shrub swamp-
C, marsh —
C.4 open pond+
c true swamp —
5 marsh-

T =T T a

ot 6 2) 4 3

Jaccard's Coefficient
Figure 8. A clustering of the macroinvertebrate

communities inhabiting the four major plant
communities of Lovets Pond using group average
clustering of the Jaccard’s coefficients of similarity.

408 Illinois Natural History Survey Bulletin

(0.170—0.612), including Limekiln Spring and
Slough which is owned by The Nature Conser-
vancy and is generally considered “protected.”
That site exemplifies the major threat to all the
remaining Cache wetlands—excessive habitat
destruction due to siltation from agricultural
endeavors. Even the integrity of the areas
“protected” by the state, by The Conservancy,
or by other private groups is being threatened
by siltation, which is obliterating most of the
available aquatic habitat. The quality of the
adjacent terrestrial habitat is variable; some
sites are cleared of all vegetation and others
have mature, high-quality forests or swamps.
Sites with the most disturbed terrestrial
component generally have the least diverse
aquatic component. Even though the data
reveal that macroinvertebrate species diversity
is generally low, enough islands of diversity
seem to exist to reclaim the area if it were
protected from further siltation and other
degrading influences. The ecological integrity
of the Cache and its wetlands cannot, however,
be considered good, especially in light of the
excessively silted substrate of the areas I
visited.

CONCLUSIONS

I have examined the ecological integrity of two
southern Illinois wetlands: one small, Lovets
Pond, and a much larger one, the Cache. I have
concluded that if drastic measures are not
immediately initiated (such as the proposed
Cypress Creek National Wildlife Refuge), the
future of the Cache River system is bleak,
primarily due to excessive siltation. On the
other hand, Lovets Pond appears adequately
protected from siltation by a forest buffer.

We should act now to preserve both
systems and all other Illinois wetlands, regard-
less of size. Large, disturbed systems such as
the Cache may recover, thereby preserving a
large portion of the biodiversity of Illinois.
Small systems such as Lovets Pond also serve
to preserve their share of biodiversity.

ACKNOWLEDGMENTS

Portions of this research were funded by the
Department of Zoology, College of Science,
Southern Illinois University at Carbondale, and
by the Illinois Nongame Wildlife Conservation
Fund. Melvin L. Warren, Jr., graciously
reviewed an early draft of the manuscript.

Vol. 34 Art. 4

LITERATURE CITED

CowarbDIN, L.M., V. CARTER, F.C. GOLET. AND E.T.
LaRoe. 1979. Classification of wetlands and
deepwater habitats of the United States. U.S. Fish
and Wildlife Service, Washington, DC. 103 p.

GrEESON, P.E., J.R. CLARK, AND J.E. CLARK, EDS.
1979. Wetland functions and values: the state of our
understanding. Proceedings of the National Sympo-
sium on Wetlands. American Water Resources
Association, Minneapolis, MN. 674 p.

ILLINOIS DEPARTMENT OF CONSERVATION. 1988.
A field guide to the wetlands of Illinois. Illinois
Department of Conservation, Springfield. 244 p.

Mitscn, W.J., AND J.G. GossELINK. 1986. Wetlands.
Van Nostrand Reinhold Co., Inc., New York. 539 p.

PHiLuippl, M.A., B-M. Burr, AND R.A. BRANDON.
1986. A preliminary survey of the aquatic fauna of
the Cache River in Johnson and Pulaski counties,
Illinois. Illinois Department of Conservation,
Springfield. 414 p.

PHILLippi, M.A., AND M. PETERSON. 1986. A pre-
liminary investigation of the aquatic macro-
invertebrate community of Lovets Pond in Jackson
County, Illinois. Illinois Department of Conserva-
tion, Springfield. 33 p.

Status and Distribution of Wetland Mammals in Illinois

Joyce E. Hofmann, Illinois Natural History Survey

Wetlands are highly productive and diverse
habitats that supply important resources for
many mammalian species (Fritzell 1988). The
objectives of this paper are to list the mammals
that are found in the wetlands of Illinois, to
identify species that are threatened or endan-
gered, and to discuss the distribution of wetland
mammals within the state, especially those
restricted to wetland habitats. Only palustrine
wetlands, rather than riverine or lacustrine
systems, are considered. These shallow water
habitats are categorized as palustrine emergent
(sedge meadow, marsh, bog, and fen), palus-
trine scrub-shrub, and palustrine forested
(swamp and seasonally or temporarily flooded
forested wetland) wetlands (Cowardin et al.
1979). Illinois mammals that inhabit these
types of wetlands are listed in Table 1.

Most of the mammals in Table | are
terrestrial or semiaquatic. Bats are not typically
considered wetland mammals, although any
Illinois species might well forage above
marshes or bogs or along the edges of swamps.
Research conducted by the Illinois Natural
History Survey and the Illinois Department of
Conservation revealed that forested wetlands in
southern Illinois provide roosting sites for three
species of bats. In May 1988, a radio-tagged
pregnant Indiana bat was found roosting behind
loose bark on a dead American elm (U/mus
americana) in a wetland created by subsidence
in Saline County. A lactating southeastern bat
was radio-tracked to the hollow base of a living
tupelo gum (Nyssa aquatica) in Little Black
Slough in Johnson County during the summer
of 1989; she shared this roost with at least 100
other individuals. Four Rafinesque’s big-eared
bats were also found roosting in a tupelo gum
in the slough during that summer. To stress the
importance of palustrine forested wetlands to
these three endangered species, I have listed
them in Table |. Other species of bats also
roost in trees during the summer, although little
is known about their specific habitat prefer-

ences (Barbour and Davis 1969; Hoffmeister
1989). Species likely to roost in forested
wetlands include the silver-haired bat
(Lasionycteris noctivagans), northern long-
eared bat (Myotis septentrionalis), and evening
bat (Nycticeius humeralis).

Table | includes one federally endan-
gered species, the Indiana bat (Endangered
Species Act, 16th U.S. Congress, docket 1531);
three state endangered species, the southeastern
bat, Rafinesque’s big-eared bat, and river otter;
and three state threatened species, the marsh
rice rat, golden mouse, and bobcat (Illinois
Administrative Code, Title 17, Chapter I,
subchapter c, part 1010.30, as amended March
17, 1989). These seven species and the swamp
rabbit (Kjolhaug et al. 1987) are uncommon in
Illinois; all other species in Table 1 range from
relatively common to abundant (Hoffmeister
1989). The beaver and white-tailed deer are
now common even though both species had
been nearly extirpated from the state by the end
of the 19th century (Pietsch 1954; Pietsch
1956; Hoffmeister 1989).

Some of the species in Table 1 have
restricted ranges within Illinois. The southern
short-tailed shrew, big-eared bat, southeastern
bat, swamp rabbit, marsh rice rat, and golden
mouse occur only in the southern portion of the
state (Ellis et al. 1978; Feldhamer and Paine
1987; Kjolhaug et al. 1987; Hoffmeister 1989;
Illinois Natural Heritage Database). The main
breeding population of river otters is along the
Mississippi River north of Rock Island (Jo
Daviess, Carroll, Whiteside, and Rock Island
counties); a smaller population may occur in
the Heron Pond-—Little Black Slough area of the
Cache River drainage (Johnson County) in
southern Illinois (Anderson 1982). Most
bobcats probably occur in the northwestern and
southernmost portions of Illinois where
relatively large expanses of suitable habitat
remain (Illinois Natural Heritage Database).
The Virginia opossum, southern flying squirrel,

409

410 Illinois Natural History Survey Bulletin

beaver, white-footed mouse, woodland vole,
muskrat, house mouse, meadow jumping
mouse, gray fox, raccoon, mink, and white-
tailed deer, on the other hand, occur throughout
the state (Hoffmeister 1989). The remaining
species in Table 1 have ranges that cover much
of Illinois. The meadow vole and least weasel
occur in the northern half of the state, and the
northern short-tailed shrew is found primarily
in the northern two-thirds (Hoffmeister 1989).
The southeastern shrew and southern bog
lemming occur in the southern two-thirds of
Illinois, although bog lemmings have been
caught in Carroll County (Hoffmeister 1989).
The Indiana bat, though rare, has been found in

Vol. 34 Art. 4

20 counties in central and southern Illinois
during the summer (Illinois Natural Heritage
Database). The masked shrew may have a
discontinuous distribution in Illinois, occurring
primarily in the northern third of the state but
also in at least two southern counties (Hoff-
meister 1989).

Many species of mammals are habitat
generalists. The home ranges of larger mam-
mals, such as the bobcat and white-tailed deer,
typically consist of a mosaic of forested areas
interspersed with open areas that could include
wetlands (Schwartz and Schwartz 1981). Many
smaller mammals may be found in a variety of
habitats. The masked shrew, for example, is

Table 1. Wetland mammals of Illinois. Terrestrial and semiaquatic species are included if their
activities (e.g., foraging, nesting) are conducted entirely or partly within palustrine wetlands; bats
are included if they are known to roost in wetlands.

Common name

Virginia opossum

Scientific name Habitat’
Didelphis virginiana FW

Masked shrew Sorex cinereus M SM B FW
Southeastern shrew Sorex longirostris M SW FW
Northern short-tailed shrew Blarina brevicauda M SM B
Southern short-tailed shrew Blarina carolinensis M

Indiana bat Myotis sodalis SW FW
Southeastern bat Myotis austroriparius SW
Rafinesque’s big-eared bat Plecotus rafinesquit SW

Swamp rabbit Sylvilagus aquaticus SS SW FW
Southern flying squirrel Glaucomys volans FW

Beaver Castor canadensis M SW FW
Marsh rice rat Oryzomys palustris M SS SW
White-footed mouse Peromyscus leucopus M SM SS FW
Golden mouse Ochrotomys nuttalli SS SW FW
Meadow vole Microtus pennsylvanicus M SM
Woodland vole Microtus pinetorum M FW
Muskrat Ondatra zibethicus M SW
Southern bog lemming Synaptomys cooperi M

House mouse Mus musculus M FW
Meadow jumping mouse Zapus hudsonius M SM

Gray fox Urocyon cinereoargenteus FW

Raccoon Procyon lotor M SS SW FW
Least weasel Mustela nivalis M

Mink Mustela vison M FW

River otter Lutra canadensis SW FW
Bobcat Felis rufus SS SW FW
White-tailed deer Odocoileus virginianus M SS SW FW

'Palustrine wetland habitats used by these species are coded as follows:

M = marsh
SM = sedge meadow
B= bog

SS = scrub-shrub wetland
SW = swamp

FW = seasonally or temporarily flooded forested wetland
Sources on habitat use: Barbour and Davis 1974; Schwartz and Schwartz 1981; Mumford and Whitaker 1982;

Jones and Birney 1988; and Hoffmeister 1989.

April 1991

abundant in sedge meadows and marshes in
northern Illinois but also inhabits sand prairies,
flatwoods, fencerows, pastures, and succes-
sional fields (Mumford and Whitaker 1982;
Mahan and Heidorn 1984; Szafoni 1989). The
white-footed mouse has been trapped in sedge
meadows and marshes (Mahan and Heidorn
1984; Szafoni 1989) but is more typically an
inhabitant of upland forests and shrublands. In
fact, few species of mammals are specifically
adapted for living in wetland environments
(Fritzell 1988). Most of the species listed in
Table | are not restricted to wetlands and,
therefore, their distribution and abundance are
not indicative of or significantly limited by the
status of wetlands in Illinois. The swamp rabbit
and marsh rice rat are the Illinois mammals that
are most limited to palustrine wetlands. The
beaver, muskrat, and river otter are also closely
associated with wetlands but are more aquatic
in their habits and could be considered species
of rivers, streams, lakes, or ponds. The swamp
rabbit and rice rat are uncommon and have
limited distributions within the state; the
remainder of this paper will discuss their
distribution and status in more detail.

The swamp rabbit is a representative of
the Eastern-Austral faunal element, the group
of mammalian species whose distributions are
centered in the southeastern United States
(Jones and Birney 1988). Its northern limit is in
Illinois and Indiana and coincides with that of
the southern swamp forest community at
approximately the 24°C temperature isoline
(Chapman and Feldhamer 1981). Swamp
rabbits rarely occur far from water and inhabit
floodplain forests, cypress swamps, and
canebrakes (Cory 1912; Layne 1958; Barbour
and Davis 1974; Sealander 1979; Chapman and
Feldhamer 1981; Hoffmeister 1989). In
Indiana, swamp rabbits were found in areas
where low ridges were interspersed with small
wooded sloughs and grassy marshes (Terrel
1972).

In the early 1900s, the swamp rabbit was
known to occur in swamps along the Missis-
sippi and Ohio rivers in Illinois; its northern
limits were thought to be a few miles south of
Grand Tower in Jackson County and 5 miles
below Golconda in Pope County (Howell
1910). The earliest specimens were collected in
Alexander and Johnson counties (Cory 1912)
and Williamson County (Necker and Hatfield

Symposium Proceedings: Our Living Heritage 411

1941). Cockrum (1949) believed that the
swamp rabbit had extended its range during the
early twentieth century as far north as Jefferson
County. He reported that hunters had killed
swamp rabbits in Franklin County during
1935-1936 and in Jefferson County during
1936. More recently, specimens and possible
sightings have been recorded in several other
counties: Marion, Massac, Perry, Randolph,
and Union (Layne 1958); Bond, Calhoun,
Gallatin, Lawrence, Wabash, Washington, and
Wayne (Klimstra and Roseberry 1969); and
Edwards and White (Terrel 1969). These
findings indicate a range extending northward
to Calhoun, Bond, and Lawrence counties
(Figure |). Whether these new records repre-
sent a range expansion or improved reporting
is, however, uncertain.

Kjolhaug et al. (1987) of the Cooperative
Wildlife Research Laboratory conducted
intensive searches for swamp rabbits or their
sign (pellets on logs, vegetation clippings,
tracks) in 11 southern Illinois counties and
limited searches in three others during
1984-1985. Sign was recorded at 22 sites
along the Bay Creek and Big Muddy, Cache,
Mississippi, and Ohio River drainages in
Alexander, Franklin, Jackson, Johnson,
Massac, Pope, Pulaski, and Union counties
(Figure 1). No sign was found in Gallatin,
Lawrence, Saline, Wabash, Wayne, and
Williamson counties, although all but Saline
had earlier records. Other counties for which
previous records exist were not searched during
the study by Kjolhaug et al. (1987).

@ Kjolhaug et al. 1987
G Earlier records

Figure 1. Southern Illinois counties in which swamp
rabbit sign was found by Kjolhaug et al. (1987) and
earlier records for this species (Howell 1910; Cory
1912; Necker and Hatfield 1941; Cockrum 1949;
Layne 1958; Klimstra and Roseberry 1969; Terrel
1969).

412 Illinois Natural History Survey Bulletin

The results of the study by Kjolhaug and

his colleagues suggest that Alexander, Johnson,

Massac, Pulaski, and Union counties support
several secure populations of swamp rabbits,
whereas this species is present at low densities
and with limited distributions in Franklin,
Jackson, and Pope counties. Only 12,585 ha in
southern Illinois were found to support swamp
rabbits, although approximately 2,000 addi-
tional hectares of suitable habitat were identi-
fied. The state of Illinois was the most impor-
tant owner of swamp rabbit habitat. The
potential habitat for this species in Illinois and
neighboring states has been drastically reduced
by the construction of levees and drainage
ditches and the conversion of bottomlands to
agricultural use (Terrel 1972; Barbour and
Davis 1974; Korte and Fredrickson 1977;
Whitaker and Arbell 1986; Kjolhaug et al.
1987; Hoffmeister 1989). In Indiana, for
example, swamp rabbits are now restricted to a
single county (Whitaker and Arbell 1986).
Fragmentation of bottomland forest and swamp
has created islands surrounded by unsuitable
habitat, a condition limiting successful disper-
sal and reestablishment of extirpated local
populations. Kjolhaug et al. (1987) concluded
that swamp rabbits were unlikely to colonize
vacant areas of habitat and that existing
populations will continue to be extirpated.

The marsh rice rat (Figure 2) is the only
member of this predominantly Neotropical
genus with an extensive range in the United
States (Honacki et al. 1982). The southern
portion of Illinois is at the northern limit of its
range, although rice rats once occurred as far
north in the state as Peoria County, where their
remains have been found at an archeological
site (Baker 1936). Rice rats are common
throughout much of their range, where they
inhabit coastal and freshwater marshes and
swamps and areas along lakes, rivers, and
streams (Wolfe 1982).

The first modern specimens from Illinois
were collected at Olive Branch and Cache in
Alexander County (Cory 1912; Necker and
Hatfield 1941). McLaughlin and Robertson
(1951) collected two specimens in Johnson
County and concluded that rice rats were
limited to swampy areas within the Coastal
Plain Division of the state (Schwegman 1973).
More recently, rice rats have also been reported
from Franklin, Jackson, Massac, Pulaski,

Vol. 34 Art. 4

Union, and Williamson counties (Klimstra and
Scott 1956; Klimstra 1969; Klimstra and
Roseberry 1969; Rose and Seegert 1982;
Urbanek and Klimstra 1986; Illinois Natural
Heritage Database). In addition, the remains of
a rice rat were found in the stomach of a mink
collected from an unspecified location in
Washington County (Casson 1984). The recent
range of the rice rat, inferred from these limited
records, extends through the Ozark, Mississippi
River Bottomlands, and Shawnee Hills divi-
sions into the Mt. Vernon Hill Country Section
of the Southern Till Plain Division.

During 1986-1987 staff members of the
Illinois Natural History Survey live-trapped in
17 southern Illinois counties to assess the
current distribution of the rice rat (Figure 3;
Hofmann et al. 1991). A total trapping effort of
3,517 trap-nights resulted in 1,111 captures of
small mammals representing 13 species. Rice
rats were captured at 13 sites in 10 counties
(Figure 3). They were found for the first time in
Hamilton, Pope, Saline, and White counties and
were also trapped at new localities in Alexan-
der, Franklin, Jackson, Johnson, Massac, and
Williamson counties. Rice rats were not caught
in Pulaski, Union, and Washington counties,
although earlier records existed. Despite recent
trapping efforts, no rice rats have been captured
in Gallatin, Hardin, Perry, and Randolph
counties. These results suggest that rice rats
occur farther to the northeast in the state than
indicated by previous records (into the Wabash
Border Division). Rice rats may have expanded
their range within the state, perhaps using
waterways and wet areas along highway and
railroad rights-of-way as dispersal corridors:
more likely, they were present in Hamilton,

Figure 2.
during the distribution study of 1986-1987. Photo by
Marilyn Mortis.

A rice rat live-trapped in Franklin County

April 1991

Pope, Saline, and White counties but un-
reported due to limited sampling. Although
some potentially suitable habitat for rice rats
occurs in Perry, Randolph, and Washington
counties, their primary range appears to extend
only as far north as Franklin and Jackson
counties in southwestern Illinois. In addition to
the 10 counties in which rice rats were captured
during the Survey’s study, they may also occur
in Pulaski and Union counties. Existing
records, however, do not suggest that they
would be common in either county. The only
specimen known from Pulaski County was
found dead in a field in January 1987 (Illinois
Natural Heritage Database), and no rice rats
have been reported from Union County since
1958 (Klimstra and Roseberry 1969; Illinois
Natural Heritage Database).

During the Survey’s study, 132 rice rats
were captured, a number that includes at least
99 individuals. Nearly half (45—49 individuals)
were trapped at the Saline County site and
more than 70% (72-76 individuals) were
caught at just four sites in Alexander, Jackson,
Pope, and Saline counties. At the nine remain-
ing sites, the number of individuals trapped was

Figure 3. Trapping sites in southern Illinois,
1986-1987 are shown as circles; sites at which
captures of rice rats occurred contain dots (Hofmann
et al. 1990). The range of this species based on
earlier records is indicated in gray (Cory 1912;
Necker and Hatfield 1941; McLaughlin and
Robertson 1951; Klimstra and Scott 1956; Klimstra
1969; Klimstra and Roseberry 1969; Rose and
Seegert 1982; Casson 1984; Urbanek and Klimstra
1986; Illinois Natural Heritage Database).

Symposium Proceedings: Our Living Heritage

413

5 or fewer. Despite the fact that their range
within the state is more extensive than had been
thought, rice rats do not appear to be common
in Illinois and their continued status as a
threatened species appears to be warranted.

Areas where rice rats were captured were
characterized by standing water and a dense
cover of emergent herbaceous vegetation,
specifically sedges (Carex spp.), rushes (Juncus
spp.), bulrushes (Scirpus spp.), spike rushes
(Eleocharis spp.), or cattails (Typha spp.).
Trapping was most successful in roadside
ditches along county or state highways and
along the shores of ponds and lakes. Since
many extensive wetlands in southern Illinois no
longer exist, rice rats occupy islands of original
or manmade wetland habitat that are often
small and widely scattered. Such areas cannot
support large populations, and small popula-
tions are especially vulnerable to extirpation
due to environmental changes, disease, or
predation. As with the swamp rabbit, recoloni-
zation of a site could be hampered by the large
expanses of unsuitable habitat separating it
from other populations.

The remaining wetland habitat of the
swamp rabbit and marsh rice rat needs to be
protected. Such protection should be the
highest priority, but habitat enhancement and
recreation may also warrant consideration.
State and federally owned forested bottomlands
could be managed to increase their quality as
swamp rabbit habitat (Kjolhaug et al. 1987).
Modern surface-mining reclamation techniques
have the potential to create habitat suitable for
rice rats (Ohlsson et al. 1982; Klimstra and
Nawrot 1985). There is no guarantee, however,
that such areas would be colonized because
existing populations are widely dispersed.
Relocation of animals to newly created or
existing wetlands may be a useful management
procedure. Whitaker and Arbell (1986) recom-
mended reintroduction of swamp rabbits into
areas with suitable habitat in Indiana, and the
feasibility of relocating rice rats is currently
being studied by the Illinois Natural History
Survey in southern Illinois. Finally, the fact that
most other mammals that use wetlands are
flexible in their habitat choices does not mean
that there is reason for complacency about the
loss of remaining Illinois wetlands.

414

ACKNOWLEDGMENTS

The study on rice rat distribution was supported
by the Illinois Endangered Species Protection
Board and the Illinois Department of Transpor-
tation. The following individuals assisted with
that study: Doug Carney, Monica Cox, Barbara
Frase, Jean Karnes, Dennis Keene, Patti
Malmborg, Ray Smith, Mary Kay Solecki, John
Taft, Karen Tyrell, Eric Ulaszek, and Mark
Wetzel. Bat data were collected by James E.
Gardner, Illinois Natural History Survey; James
D. Garner, Illinois Department of Conserva-
tion; and the author with assistance from
Randall Collins, Rebecca Porter, and Kelley
Neelley, and with support from the Illinois De-
partment of Transportation. Donald F. Hoft-
meister and John O. Whitaker, Jr., provided
valuable reviews of the manuscript.

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Session Four: Streams and Caves

Who hears the fishes when they cry?—Henry David Thoreau

More than half of the 13,200 miles of streams in
Illinois have been dredged, channelized,
dammed, or altered in other ways. Our rivers
and streams suffer from pollution, siltation, and
the introduction of exotic organisms. The
Illinois River, described by Thomas Jefferson as
“a fine river, clear, gentle, and without rapids,”
has served as Chicago’s sewer, a waterway for
untold numbers of barges made navigable only
by numerous dams, and a repository for much of
the eroded topsoil from central Illinois farm-
land. The “typical” stream in east-central Illinois
is a narrow ditch lined with mowed grass,
weeds, or row crops, stretching across the
landscape and disappearing into the distance.
The Cache River in southern Illinois was
diverted in 1916 via the Post Creek Cutoff.
Designed to alleviate flooding, it cut the river in
two, allowing a portion to drain directly into the
Ohio River. As a result, the Lower Cache has
become a sluggish trickle that even flows
backwards upon occasion.

Surprisingly, a few high-quality streams
remain in Illinois. The Biological Stream
Characterization, an index of stream quality
completed in 1989, identified 24 stream seg-
ments of excellent quality throughout the state.
These total somewhat less than 500 miles, about
4% of the stream mileage in Illinois. Included in
this group are segments of the Kishwaukee in
northern Illinois, the Vermilion in east-central
Illinois, and Lusk and Big creeks in the
Shawnee National Forest.

Caves in Illinois have fared somewhat
better. Four areas where caves are typically
found correspond to major outcroppings of
calcareous rocks. More than 480 caves were
identified during the 1988 inventory conducted
by the Illinois State Museum.

The remarkably stable, insulated environ-
ments of caves support a unique biota. For the
most part, these organisms are adapted to little
or no light and limited food resources. Caves are

416

regarded as natural zoological laboratories
where, because of the relative simplicity of the
ecosystem, important biological and evolution-
ary questions can be studied.

One presentation at this session surveyed
the nature of Illinois streams—what we have,
what we have lost, and what can yet be done
by way of restoration and preservation. Two
speakers focused on inhabitants of that stream
system, the surprisingly diverse and dynamic
Illinois fish fauna and the varied mussel
populations. The fourth paper described the
cave environment and ecosystem, noting the
often overlooked values of this unique natural
resource.

The Fishes of Illinois: An Overview of a Dynamic Fauna

Brooks M. Burr, Department of Zoology, Southern Illinois University at Carbondale

Just over ten years ago, Smith (1979) published
the most recent comprehensive summary of the
Illinois fish fauna. His review revealed 199 fish
species, 186 of which were considered native to
the state. A major finding was that the Illinois
fish fauna is dynamic and that the distributions
of many species have changed considerably
since the first comprehensive survey of Illinois

fishes by Forbes and Richardson ([{1908], 1920).

Because of introductions of alien species, dis-
coveries of species new to Illinois, and redis-
coveries of species formerly thought to be
extirpated, the composition of the Illinois fish
fauna is in need of clarification.

In the past decade, the greater redhorse,
Moxostoma valenciennesi (Seegert 1986), and
the cypress minnow, Hybognathus hayi (Burr
and Mayden 1982; Warren and Burr 1989),
which were thought to have been extirpated
from Illinois, were rediscovered. Examination
of collections made prior to Smith’s survey and
recent collecting have documented previously
unreported records for the bluehead shiner,
Preronotropis hubbsi (Burr and Warren 1986),
and the pallid shiner, Hybopsis amnis (Warren
and Burr 1988). Three fishes were recently
added to the state fauna; in addition, new
localities for ten other uncommon species were
reported by Burr et al. (1988) and by Dimmick
(1988). The introduced rainbow smelt, Osmerus
mordax, has recently and rapidly extended its
range in Illinois (Burr and Mayden 1980). The
white perch, Morone americana, previously un-
recorded from Illinois, has dispersed into the
Illinois portion of Lake Michigan (Savitz et al.
1989a). The bighead carp, Hypophthalmichthys
nobilis, silver carp, Hypophthalmichthys
molitrix, and rudd, Scardinius erythrophthal-
mus— three Eurasian exotics unknown in
Illinois streams during Smith's (1979) survey—
are being captured at a number of localities,
particularly big rivers and reservoirs.

My purpose here is to review briefly the
Illinois fish fauna and record some of the

changes that have occurred in the composition
of Illinois fishes since Smith’s (1979) compre-
hensive study. I have used the term ‘alien’ to
encompass any fish species “of foreign origin”
that is either an exotic, a transplant, or a
recently invading species from more southern
latitudes.

HISTORICAL PERSPECTIVE

The history of ichthyological investigations in
Illinois is a rich one. At the time the Illinois
Natural History Society was established in
1858, approximately three-fourths of the Illinois
fish fauna had been named and described by
such distinguished ichthyologists as Samuel L.
Mitchill (1764-1831), Charles A. Lesueur
(1778-1846), Constantine S. Rafinesque
(1783-1840), Jared P. Kirtland (1793-1877),
Louis Agassiz (1807-1873), and Charles F.
Girard (1822-1895). Fourteen of the species
described were first discovered in Illinois.

The first regional list of Illinois fishes was
prepared by Robert Kennicott (1855), who
treated the fishes of the Chicago area. Compre-
hensive catalogs of fishes of the entire state
later appeared by Edward W. Nelson (1876),
David Starr Jordan (1878), Stephen A. Forbes
(1884), and Thomas Large (1903).

Intensive Illinois ichthyology, however,
began with Stephen Forbes (1844-1930; Figure
1), the first Director of the State Laboratory of
Natural History then in Normal, Illinois, and
later moved to Urbana-Champaign in 1885.
Sometime in the 1870s, Forbes developed the
idea of producing a well-illustrated and detailed
account of Illinois fishes. Year after year horse-
drawn wagon parties were sent to explore and
collect in different streams of the state until
finally records were available for virtually
every river in Illinois. The monumental effort
that went into the project represented the
patience and toil of 30 years. The final report,
The Fishes of Illinois, appeared in 1908

417

418 Illinois Natural History Survey Bulletin Vol. 34 An. 4

Figure 1. Stephen Alfred Forbes (1844-1930).
Photo courtesy of Illinois Natural History Survey.

Figure 2. Location of collections of fishes made
from 1876 to 1903. From Forbes and Richardson
[1908].

(although no publication date is given in the
volume) and was authored by Forbes and his
colleague Robert Earl Richardson (1877-1935).
A separate atlas of 103 range maps accompa-
nied the volume. At that time, The Fishes of
Illinois was considered by many to be the best
regional ichthyology ever published on fishes in
North America. Exceptionally skillful water
colors of many species (52 in the 1908 edition,
68 in the 1920 edition), some never before
published in color, were included and helped to
make the book an immediate classic. Most of
the copies of the initial edition were burned in a
warehouse fire, and a second edition was
produced in 1920.

The Forbes and Richardson data base
(Figure 2) included over 200,000 fish speci-
mens and 1,545 collections made from about
475 localities representing all major drainages
and 93 of the 102 counties of Illinois. A total of
142 presently valid species (Table 1) was
recorded from Illinois waters by Forbes and
Richardson [1908], and only one (common
carp, Cyprinus carpio) of those was an alien
species. About 20,000 specimens used in the
original Fishes of Illinois are vouchered in the
collection of the Illinois Natural History
Survey. Clearly, the superb historical data base
for Illinois fishes is unique and unsurpassed by
that of any other state or province in North
America.

Subsequent to the masterful Forbes and
Richardson treatise appeared works by Meek
and Hildebrand (1910) on fishes of the Chicago
region and another list of Illinois fishes by
O'Donnell (1935), which added a few species
to the known fauna of the state. A large number
of collections made during the 1940s by Aden
C. Bauman, a student of Carl L. Hubbs,
contributed many significant records of Illinois
fishes, particularly from the southern half of the
state. Bauman’s collections are at the University
of Michigan Museum of Zoology and have only
recently been used (Lee et al. 1980; Burr and
Mayden 1982; Warren and Burr 1989).

In about 1950, Philip W. Smith (1921-
1986; Figure 3), former head of one of the
Illinois Natural History Survey’s scientific
sections and author of The Amphibians and
Reptiles of Illinois (1961), undertook to
resurvey the fishes of the state. This task
provided a unique opportunity for comparing
modern-day distributional data with the classic
work of Forbes and Richardson. The bulk of

April 1991

Smith’s fieldwork began in the summer of 1962
and continued until the mid-1970s. During this
period Smith published an account of the fishes
of Champaign County (Larimore and Smith
1963), an annotated preliminary list of Illinois
fishes (Smith 1965), an assessment of Illinois
streams based on fish distribution data (Smith
1971), a key to Illinois fishes (Smith 1973), and
finally, a new Fishes of Illinois (Smith 1979)
that summarized the identification, biology, and
distribution of the Illinois fish fauna.

Smith and his colleagues found 199
species in Illinois (Table 1), made over 3,000
collections from over 2,000 localities in all of
the drainages of the state and in all of the 102
counties (Figure 4), and preserved as vouchers
approximately 400,000 specimens deposited at
the Illinois Natural History Survey. When he
compared his data with those of Forbes and
Richardson, Smith (1971:8) found that about 70
Illinois fishes clearly showed patterns of range
decimation or extirpation from the state and that
13 alien species occupied Illinois waters.

Since the publication of Smith’s (1979)
treatise, state fish biologists have continued to
collect data on the Illinois ichthyofauna.
Particularly active have been ichthyologists and
fish biologists from the state’s universities, the
Illinois Natural History Survey, the Illinois
Department of Conservation, and several
consulting firms. Additional discoveries of
exotic species, native species previously
unreported, and the invasion of more southerly
species into Illinois waters emphasize the
dynamic nature of the Illinois fauna and the
need for continued collections of fishes even in
presumably well-surveyed areas.

Symposium Proceedings: Our Living Heritage 419

DYNAMIC NATURE OF THE ILLINOIS
FAUNA

Illinois has many drainage systems and is
bounded on the west by the Mississippi River,
on the south by the Ohio River, on the east by
the Wabash River, and on the northeast by Lake
Michigan. The numerous interior streams,
glacial lakes in Lake County, and cypress—
tupelo swamps in southern Illinois account for
the richness of the fauna. Illinois has the lowest
average elevation of the north-central states.
More than 90% of the state lies within the
Central Lowlands Province, all of which was
glaciated except the Driftless Area in extreme
northwestern Illinois. Although well-watered,
Illinois has lost many aquatic habitats to
agriculture, stream impoundments, industrial
and domestic pollution, and other modifications
of watersheds.

Disappearance of Native Species

As noted previously, Smith (1971:8) docu-
mented range decimation or rarity for approxi-
mately 70 Illinois fishes; later, Smith

(1979: xviii—xix) revised this number to include
52 species, some of which probably were rare
even prior to European settlement. For about
120 species, no range change was detected.
According to Smith (1971), several factors are
primarily responsible for the disappearance of
native Illinois fishes: 1) excessive siltation has
caused the extinction or decimation of at least
16 species through loss of water clarity,
disappearance of aquatic vegetation, and
deposition of silt over rocky or sandy sub-
strates; 2) drainage of wetlands has shrunk the
ranges of at least 13 species; 3) desiccation

Table 1. Composition of Illinois fishes over the past century.

—_e—ee—e—e—eee————————eeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee

Total no. of species

No. of aliens No. extirpated

Forbes and Richardson [1908]
Smith (1979)
Present Information (1990)

142! (141 native) l
199 (186 native) 13
209° (187 native) 223 127

Not applicable
9

' Forbes and Richardson [1908] recognized 150 species, 142 of which are considered valid today.
* Additions since Smith (1979) include Atlantic salmon, bighead carp, silver carp, rudd, taillight shiner, inland
silverside, threespine stickleback, striped mullet, white perch, and Rio Grande cichlid.

“The number of alien species also includes three relatively recent invaders from the south (threadfin shad, inland silverside,
and striped mullet); the first two of these are also stocked as forage in Illinois reservoirs.

* Extirpations since Smith (1979) include bluehead shiner, bigeye chub, harlequin darter, northern madtom, and alligator gar.

The cypress minnow and greater redhorse, both included as extirpated by Smith (1979), have been rediscovered recently in

Illinois, as noted in the text.

420

during drought, which has dried up once
permanently flowing streams, stopped the flow
in seeps and springs, and temporarily reduced
the size of formerly larger rivers, has shrunk the
ranges of at least 12 species; 4) interactions
between species, including the effects of
introduced species on native ones, competitive
supplantation, and aggressive dispersal by
ecologically labile species, has caused the
extinction or decimation of at least 9 species; 5)
industrial, domestic, and agricultural pollution
has caused the decimation of at least 5 species;
6) dams and impoundments are responsible for
the decimation of at least 4 species through the
loss of a large variety of habitats and the
blocking of natural migration; 7) higher water
temperatures now than formerly, chiefly the
result of stream channelization and the removal
of marginal vegetation, have caused the
decimation of at least 1 species. No single
factor has as yet been identified for the extirpa-
tion of the muskellunge, Esox masquinongy,
from northern Illinois or the saddleback darter,
Percina ouachitae, from the Wabash River.

Since the publication of Smith’s book
(1979), the continued decline of several species
has been documented. Examples include the
pallid shiner, Hybopsis amnis, a species now
known to have been much more widespread in
Illinois than indicated on Smith’s (1979)
distribution map. It has disappeared from seven
major Illinois drainages where it was known to
occur from the late 1800s through the 1940s
(Warren and Burr 1988). It remains in the
Kankakee River drainage (Skelly and Sule
1983) and in the upper Mississippi River
(Warren and Burr 1988). The Mississippi
silvery minnow, Hybognathus nuchalis, was not
taken in the recent (late 1980s) survey of the
fishes of Champaign County and was rarely
taken in several recent surveys in southern
Illinois where suitable habitat was present. The
bigeye shiner, Notropis boops, continues to
disappear from sites of former occurrence but
survives in the Little Vermilion River and the
Clear Creek drainage of southern Illinois. Major
impoundments (Carlyle and Shelbyville
reservoirs) on the Kaskaskia River have
severely limited the habitat of the western sand
darter, Etheostoma clarum, which is now very
rare (if not extinct) in the drainage. The species
has, however, been taken recently in the
Mississippi River below the mouth of the
Missouri River (Dimmick 1988).

Illinois Natural History Survey Bulletin

Vol. 34 Art. 4

Figure 3. Philip Wayne Smith (1921-1986). Photo
courtesy of Illinois Natural History Survey.

Figure 4. Location of collections of fishes made
from 1950 to 1978. From Smith 1979.

April 1991

Another striking discovery emanating
from Smith’s (1979) survey and subsequent
work was the relatively large number of Illinois
fishes that have been extirpated since the
original Forbes and Richardson (1908) survey.
As of this writing, these include eight species:
Ohio lamprey, /chthyomyzon bdellium; blackfin
cisco, Coregonus nigripinnis; muskellunge,
Esox masquinongy; rosefin shiner, Lythrurus
ardens; gilt darter, Percina evides; saddleback
darter, Percina ouachitae; crystal darter,
Crystallaria asprella; and spoonhead sculpin,
Cottus rice.

Even more alarming is the number of
species that have disappeared since Smith
(1979) began his survey in the 1960s. Examples
include the bluehead shiner, Preronotropis
hubbsi, last collected in Illinois waters in 1974
(Burr and Warren 1986) and the bigeye chub,
Hybopsis amblops, last collected in 1961
(Smith 1979; Warren and Burr 1988). In
addition, the harlequin darter, Etheostoma
histrio, known previously from the Embarrass
River, Cumberland and Jasper counties, is
almost certainly extinct in Illinois, probably
because of drainage alterations below Lake
Charleston dam. My recent attempts (1987,
1988) to collect the northern madtom, Noturus
stigmosus, in the Wabash drainage of Illinois
have been unsuccessful. The alligator gar,
Atractosteus spatula, has not been taken in
Illinois since 1965, although sufficient effort
has not been expended recently to clarify its
status.

On a positive note, at least two species
thought to have been extirpated at the time of
Smith’s (1979) survey have been rediscovered
in Illinois. The cypress minnow, Hybognathus
hayi, is now known with certainty to be
reproducing in the middle Cache River drainage
(and possibly in Horseshoe Lake) in southern
Iilinois but is still considered extirpated from
former sites of occurrence in the Big Muddy
River drainage (Warren and Burr 1989). The
drainage of wetlands that are used as nursery
areas by the species is thought to be the main
factor responsible for extirpation from the Big
Muddy River. The greater redhorse, Moxostoma
valenciennesi, thoughtto have been extinct in
Illinois since 1901, was collected in 1985 from
the Illinois River, rivermile 249 (Seegert 1986)
and again in 1989 from the Illinois River,
rivermile 270.5. These two individuals must be
part of a population residing somewhere in the
upper basin.

Symposium Proceedings: Our Living Heritage 421

Native Species Previously Unrecorded

One native fish has been added to the state
ichthyofauna since Smith’s (1979) report. The
taillight shiner, Notropis maculatus, was
discovered for the first time in Illinois in a
wetland in Massac County in 1987 (Burr et al.
1988). This species was captured at only | of 22
wetlands sampled on the lower Wabash and
Ohio rivers (Burr and Warren 1987) and should
be recognized as endangered in Illinois and
given highest priority for protection.

Species Expanding Their Ranges
Because the Illinois fish data base is extensive,
covers two broad historical periods, and is well
vouchered, it allows us to be reasonably
confident of the ranges of most native, nongame
fishes within the confines of Illinois. While
many species have experienced range reduc-
tions in the last 90 years, a few others have
expanded their ranges in response to wide-
spread modification of habitats. An outstanding
example is the red shiner, Cyprinella lutrensis,
a species tolerant of wide fluctuations in pH,
dissolved oxygen, and thermal shock (Matthews
and Hill 1977}. Additionally, its adaptable
feeding habits and reproductive capability
(Matthews and Hill 1977) in combination with
its tolerance for the above-mentioned parame-
ters undoubtedly account for its success in
Illinois. This species has expanded its range
north into Wisconsin, up the Ohio River
drainage of southern Illinois into Kentucky and
the lower Wabash River, and beginning in the
1960s crossed over from Mississippi River
drainages into the upper Vermilion River
drainage (Page and Smith 1970), where it has
continued to move downstream to Champaign
County. Another example is the silverjaw
minnow, Ericymba buccata, which has ex-
panded its range chiefly in the Illinois River
drainage. This pioneering species quickly
disperses into newly dredged ditches with sandy
substrates. Because Illinois streams tend to be
wider and shallower than formerly (Larimore
and Smith 1963), suitable habitat for species
tolerant of these conditions has increased.
Nearly all game/sport fishes and some
forage species (e.g., golden shiner,
Notemigonus crysoleucas, and fathead
minnow, Pimephales promelas) have had their
ranges expanded by numerous introductions
which continue unabated in Illinois. The
mosquitofish, Gambusia affinis, has been

422

22 Illinois Natural History Survey Bulletin

widely transplanted in efforts to control
mosquito outbreaks. The inland silverside,
Menidia beryllina, was collected in 1978 from
the Mississippi River at Grand Tower (a record
included in a footnote by Smith [1979:211]).
Beginning in 1980, this fish has been stocked as
a forage species in several southern IIlinois
ponds and impoundments (Stoeckel and
Heidinger 1989). Examples of game/sport
fishes recently captured in the Illinois waters of
Lake Michigan and not reported in Smith
(1979) include the channel catfish, /ctalurus
punctatus, and the black crappie, Pomoxis
nigromaculatus (Savitz et al. 1990). Smith
(1971:8) lists another five native species whose
ranges have expanded in recent times.

New Records of Rare or Geographically
Limited Species

Collections of Illinos fishes made during the
1940s by A.C. Bauman and those made during
the 1980s have revealed new records for rare or
geographically limited Illinois species that
expand the information in Smith (1979). For
example, the lake sturgeon, Acipenser fulves-
cens, not reported from the Mississippi River
since 1966, is known from three recent records
in the Mississippi (Burr et al. 1988) and Ohio
rivers (Burr et al. 1990). New localities for eight
other uncommon Illinois fishes were included
in Burr et al. (1988). Dimmick (1988) reported
the first Illinois records of the western sand
darter, Etheostoma clarum, from the Missis-
sippi River south of the mouth of the Missouri
River; Savitz et al. (1989b) recorded the first
record of the quillback, Carpiodes cyprinus, in
the Illinois waters of Lake Michigan. Examina-
tion of voucher specimens from several U.S.
museums has resulted in a reassessment of the
ranges of the bigeye chub and pallid shiner
(Warren and Burr 1988) as originally presented
in Smith (1979).

The Alien Component and Recent Southern
Invasions

Since Smith’s (1979) survey, three exotics, the
bighead carp, silver carp, and rudd, in addition
to the four Smith reported, have been found at
several localities in Illinois and, if not already
established, almost certainly will be within a
few years. The potential ecological effects of
introduced and exotic fishes on native aquatic
communities include habitat alterations (e.g.,
removal of vegetation, degradation of water
quality); introduction of parasites and diseases;

Vol. 34 Art. 4

trophic alterations (e.g., predation, competition
for food); hybridization; and spatial alterations
(e.g., overcrowding) (Taylor et al. 1984).

Twenty-two (10.5 %) of the total of 209
fish species in Illinois are not native to the state
(Table 2). Of these, at least 13 were probably
intentionally introduced, 5 spread through
manmade canals in the Great Lakes drainage to
the Illinois portion of Lake Michigan, | was an
unintentional introduction, and 3 euryhaline
species recently invaded from more southern
latitudes.

The presence of new species raises
questions as to their source, their ecological role
in Illinois, and their importance to human
welfare. Among the 22 species, 7 are introduc-
tions from Europe or Asia; 3 are from western
North America; 8 are from eastern fresh waters
of the Atlantic Coast, of which 3 are introduced
and 5 used canals; 3 are native to the lower
Mississippi basin or Gulf Coast and have
entered the state naturally or by human transfer;
and | (the cichlid) was presumably introduced
accidentally with other sport fishes. Several,
probably many, additional species have in the
past been introduced into Illinois waters but are
not known to persist. Thousands of Atlantic
salmon, Salmo salar, were introduced into the
Mississippi River in the late 1800s (Carlander
1954). Apparently the stockings were not
successful, although several individuals
collected in 1986 from the Mississippi River
near Chester (Burr et al. 1988) indicate that
illegal stockings have apparently occurred in
the river in recent decades. Grass, silver, and
bighead carps have been encountered at many
localities in Illinois, and the grass and bighead
carps are known to be reproducing in the upper
Mississippi River basin (Pflieger and Grace
1987; Pflieger 1989; Jennings 1989). A plethora
of tropical and subtropical aquarium fishes have
surely been released into Illinois waters (see
Smith [1965] for examples) only to perish in the
ensuing winter. One exception is the Rio
Grande cichlid, Cichlasoma cyanoguttatum,
released accidentally in the mid-1980s into
Powerton Lake near Pekin; individuals have
been observed setting up territories in that
thermally treated lake during summer months
(Rich Monzingo, pers. comm.). The threespine
stickleback, Gasterosteus aculeatus, captured
twice in 1988 from the Illinois portion of Lake
Michigan (at Trident Harbor and Cicero), is
apparently spreading rapidly through the upper
Great Lakes. It was first taken in Lake Huron in

April 1991

1982 (C. L. Smith 1985:276), but whether the
species is self-sustaining in Illinois waters is not
known.

Some of the alien species are localized
geographically, rare, or small and apparently
unimportant ecologically. In contrast, the
salmonids, striped bass, and recently introduced
carps are much valued as recreational species or
for weed control, and some are common and
becoming widespread. Another group of species
includes the locally abundant alewife and
goldfish, the widespread common carp, and the
rapidly spreading rainbow smelt and white
perch. These species are more or less controver-
sial, being variously valued as sources of food
or recreation but with negative ecological
attributes (e.g., periodic alewife die-offs,

Symposium Proceedings: Our Living Heritage

423

predation, unfavorable ecological interactions
with native species). The rainbow smelt, the
most numerous small species in some winter
seine samples from the Mississippi River for
over 10 years, has not been collected from June
through October and is probably not self-
sustaining in the Illinois portion of the Missis-
sippi River. The sea lamprey, an alien in Lake
Michigan, has played a major role in the history
and fisheries of the Great Lakes Basin.

One of the most surprising invasions in
Illinois was the appearance during the fall of
1989 of the striped mullet, Mugil cephalus, in
the Mississippi and Ohio rivers. This princi-
pally marine species had not been reported
previously from Illinois waters and was known
only in the published literature as far north in

Table 2. General distribution in Illinois of alien fish species and recent invaders from southern latitudes.
Numbers in parentheses indicate (1) exotics introduced directly into Illinois, (2) transplants from elsewhere in
North America, (3) species colonized after introduction elsewhere or through manmade access, and (4) species

that have recently invaded.

Fish species by family

Petromyzontidae
Petromyzon marinus, sea lamprey (3)
Clupeidae
Alosa pseudoharengus, alewife (3)
Dorosoma petenense, threadfin shad (2, 4)

Salmonidae
Oncorhynchus kisutch, coho salmon (2)
Oncorhynchus mykiss, rainbow trout (2)
Oncorhynchus tshawytscha, chinook salmon (2)
Salmo salar, Atlantic salmon (2)
Salmo trutta, brown trout (1)
Osmeridae
Osmerus mordax, rainbow smelt (3)
Cyprinidae
Carassius auratus, goldfish (1)
Ctenopharyngodon idella, grass carp (1)
Cyprinus carpio, common carp (1)
Hypophthalmichthys molotrix, silver carp (1)
Hypophthalmichthys nobilis, bighead carp (1)
Scardinius erythrophthalmus, rudd (1)
Ictaluridae
Ameiurus catus, white catfish (2)
Moronidae
Morone americana, white perch (3)
Morone saxatilis, striped bass (2)
Atherinidae
Menidia beryllina, inland silverside (2, 4)
Gasterosteidae
Gasterosteus aculeatus, threespine stickleback (3)
Mugilidae
Mugil cephalus, striped mullet (4)
Cichlidae
Cichlasoma cyanoguttatum, Rio Grande cichlid (2)

General distribution in Illinois

L. Michigan

L. Michigan
Ohio R., Mississippi R., Wabash R., southern
Illinois reservoirs

L. Michigan

northern half of Illinois

L. Michigan

Mississippi R.

northern Illinois, L. Michigan
L. Michigan, Illinois R., Mississippi R., Ohio R.
Illinois and Rock R. drainage
big rivers, reservoirs, ponds
statewide

big rivers, reservoirs, ponds
big rivers, reservoirs, ponds
northern Illinois; sporadic

Illinois R., Mississippi R., Kaskaskia R.

L. Michigan
Illinois reservoirs

southern Illinois reservoirs, Mississippi R.
L. Michigan
Ohio R., Mississippi R.

Powerton L., Pekin

424 Illinois Natural History Survey Bulletin

the Mississippi River as southern Arkansas
(Robison and Buchanan 1988). According to
William L. Pflieger (pers. comm.), striped
mullets were obtained from the Mississippi
River at New Madrid in 1983 and at Cape
Girardeau in 1988. The lower water levels in
the Mississippi River in 1989 may have created
water quality conditions (e.g., high dissolved
solids) favorable for striped mullet and allowed
them to reach the upper Mississippi River basin
(Burr et al. 1990).

ENDANGERED, THREATENED, AND
WATCH LIST SPECIES

In the approximately 150 years since Europeans
actively colonized the state of Illinois, changes
in the fish fauna have been profound. Of the
187 native species (Table 1), a few have
expanded their ranges and are now more
abundant and more generally distributed than
formerly, but many more have been decimated
to some degree by the widespread modification
of habitats and deterioration of water quality.
Prior to the passage of the federal Endangered
Species Act in 1973, attempts had been made
(e.g., Lopinot and Smith 1973) to list species as
rare or endangered on the basis of their natural
rarity, restricted distribution, and paucity of
habitat as well as on the basis of immediate or
potential threats to their existence within
Illinois (Smith 1979). After implementation of
the act, terminology was revised to include the
categories endangered and threatened. Since the
longjaw ciscoe, Coregonus alpenae, is no

Vol. 34 Art. 4

longer considered a valid species and was never
officially reported from the Illinois waters of
Lake Michigan, none of the Illinois species
qualifies as endangered (actively threatened
with extinction) in the sense of the federal
definition.

The Illinois Endangered Species Act of
1972 (amended in 1977) provides for some
protection of rare fishes. Lists (Smith and Page
1981; Illinois Endangezed Species Protection
Board 1990) of endangered and threatened
fishes have continued to be revised and up-
dated; however, potential threats to rare fishes
are always present and the status of each is
constantly subject to change. A change in status
can occur quickly, particularly in a peripheral or
relict population.

Thirteen of the 187 native species are
endangered and 15 are threatened (Table 3).
Eleven species have been placed on a watch list
(Table 4), an action that suggests they may be
recategorized as endangered or threatened
depending on changes that take place in Illinois.
A significant concern to conservation biologists
and others is the status and protection of those
species that are restricted to big, free-flowing
rivers (i.e., the Mississippi River). Some of the
species on the watch list are big river fishes:
however, because these species do not occur
generally within the “inland” waters of state
boundaries, they are not receiving the protec-
tion they warrant. Examples of big river fish
needing more formal protection in Illinois
include the pallid sturgeon, Scaphirhynchus
albus, the flathead chub, Platygobio gracilis,

Table 3. Fishes categorized as endangered or threatened in Illinois according to the Illinois Endangered
Species Protection Board (1990). Nomenclature has been modified where appropriate to follow Page and

Burr (1991) and Warren (1989).
Endangered

Northern brook lamprey, /chthyomyzon fossor
Bigeye chub, Hybopsis amblops

Pallid shiner, Hybopsis amnis

Pugnose shiner, Notropis anogenus

Weed shiner, Notropis texanus

Bluehead shiner, Preronotropis hubbsi
Cypress minnow, Hybognathus hayi
Greater redhorse, Moxostoma valenciennest
Northern madtom, Noturus stigmosus
Western sand darter, Etheostoma clarum
Eastern sand darter, Etheostoma pellucidum
Bluebreast darter, Etheostoma camurum
Harlequin darter, Etheostoma histrio

Threatened

Least brook lamprey, Lampetra aepyptera
Lake sturgeon, Acipenser fulvescens
Alligator gar, Atractosteus spatula

Cisco, Coregonus artedii (or artedi)

Lake whitefish, Coregonus clupeaformis
Bigeye shiner, Notropis boops

Ironcolor shiner, Notropis chalybaeus
Blackchin shiner, Notropis heterodon
Blacknose shiner, Notropis heterolepis
River redhorse, Moxostoma carinatum
Longnose sucker, Catostomus catostomus
Banded killifish, Fundulus diaphanus
Redspotted sunfish, Lepomis miniatus
Bantam sunfish, Lepomis symmetricus
lowa darter, Etheostoma exile

April 1991

the sturgeon chub, Macrhybopsis gelida, and
the sicklefin chub, Macrhybopsis meeki. These
four species are restricted in Illinois to the main
channel of the Mississippi River below the
mouth of the Missouri River. Intermittent
sampling in the Mississippi River below the
mouth of the Missouri River over a 12-year
period indicates that the three chub species are
naturally rare and sporadic in occurrence. Small
numbers of the sicklefin chub are still being
captured, but the flathead and sturgeon chubs
have been taken once each since 1985. The
pallid sturgeon is so rare throughout its range
that it is being considered for listing as a
federally endangered species.

If species that are considered extirpated
from Illinois and those on the endangered,
threatened, or watch lists are included, 46
species or 24% of the native fauna are experi-
encing trouble maintaining viable populations
in Illinois. The addition of the taillight shiner,
flathead chub, and sicklefin chub, which are
presently not on any formal list, brings the total
to 49 species or 26%.

RECOMMENDATIONS

Illinois is a model state in view of its excellent
data base on fish distributions over time.
Although we have learned a great deal about the
effects of human activities on the aquatic
environment in Illinois, we must continue to
conduct basic survey work on Illinois fishes and
document long-term changes in the fauna.
Because fishes are sensitive indicators of
environmental quality, continued collection of
data will aid in monitoring a variety of stream-
quality parameters and assist state agencies in

Table 4. Fishes placed on the watch list by the
Illinois Endangered Species Technical Advisory
Committee on Fishes. These species do not receive
protection under federal or state laws.

Pallid sturgeon, Scaphirhynchus albus

Round whitefish, Prosopium cylindraceum
Lake chub, Couesius plumbeus

River chub, Nocomis micropogon

Gravel chub, Erimystax x-punctatus

Sturgeon chub, Macrhybopsis gelida

Blacktail shiner, Cyprinella venusta

Northern starhead topminnow, Fundulus dispar
Fourhorn sculpin, Myoxocephalus quadricornis
Spoonhead sculpin, Coftus ricei

Cypress darter, Etheostoma proeliare

Symposium Proceedings: Our Living Heritage

425

identifying high-quality aquatic habitats in need
of protection.

Because of the number of species
extirpated or endangered in Illinois, we need to
establish a monitoring program and status
surveys of species on the watch list. Several of
the species on the Illinois endangered list are
probably already extirpated (e.g., bigeye chub,
bluehead shiner) and the most effective course
of action might be to allocate funds and efforts
on species that may be realistically recoverable.

Over the last several years, we have come
to recognize that we know comparatively little
about the fundamental life histories of nongame
fishes in contrast to the voluminous literature on
the biology of game or sport fishes. If we are
ever going to manage nongame species effec-
tively, more funding is needed for studies on
basic fish biology, especially those emphasizing
reproductive biology, trophic ecology, predator-
prey interactions, and parasites and diseases.

The purchase of critical habitat by The
Nature Conservancy, the Illinois Department of
Conservation, and other agencies has provided
islands of habitat where some rare fish species
can survive. For the taillight shiner, the pur-
chase of critical habitat may be the best measure
for protecting this rare and highly localized
species. Several rare Illinois fishes that occur in
relatively undisturbed and protected areas (e.g.,
LaRue-Pine Hills Swamp) continue to maintain
viable populations. Efforts to purchase critical
stream and wetland habitats in Illinois need to
increase.

Game and sport fishes have been stocked
in Illinois waters for many years. Within reason,
state agencies should now consider stocking
certain nongame fishes in an attempt to restore
viable populations. Pond culture of endangered
and threatened species should be continued in
Illinois because it has provided a useful
environment for studying aspects of the
fundamental life histories of rare species; this
information in turn leads to more effective
management.

Because siltation is still considered to be
the number one factor in decimation of native
fish populations, we must continue to work
creatively with farmers and others in protecting
the valuable prairie topsoil of Illinois. The
removal of gravel from headwater streams
should be discouraged because the process
increases erosion and destroys breeding sites of
headwater creek fishes. Reservoir construction

426

and stream channelization should also be
discontinued in Illinois because of the detrimen-
tal effects these practices have on large ex-
panses of aquatic habitat.

Finally, basic survey work on the big
rivers of Illinois is badly needed. While we
know comparatively little about the biology of
small stream species, we know next to nothing
regarding nongame, big river fishes. Unusual
Illinois species (e.g., the pallid sturgeon) may
disappear before we learn anything substantial
about them or can protect them.

ACKNOWLEDGMENTS

Iam grateful to Larry A. Jahn, William L.
Pflieger, and Lawrence M. Page for construc-
tive comments on an earlier draft of this report.
Kevin S. Cummings and Christine A. Mayer
helped compile information on Illinois fish
collections at the Illinois Natural History
Survey.

LITERATURE CITED

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rainbow smelt, Osmerus mordax, into the upper
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Burr, B.M., AND R.L. MAYDEN. 1982. Status of the
cypress minnow, Hybognathus hayi Jordan, in
Illinois. Chicago Academy of Sciences Natural
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Burr, B.M., AND M.L. WarrEN, JR. 1986. Status of
the bluehead shiner (Notropis hubbsi) in Illinois.
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Burr, B.M., AND M.L. WARREN, JR. 1987. Wetland
resources of the Ohio and lower Wabash rivers—an
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Burr, B.M., M.L. WARREN, JR., AND K.S.
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Burr, B.M., M.L. WARREN, JR., G.K. WEDDLE, AND
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April 1991

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the fish fauna of the lower Missouri River,
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of Oklahoma Press, Norman. viii + 310 p.

Rosison, H.W., AND T.M. BUCHANAN. 1988. Fishes
of Arkansas. University of Arkansas Press, Fayette-
ville. 536 p.

Savitz, J., C. AIELLO, AND L.G. BARDYGULA. 1989a.
The first record of the white perch (Morone ameri-
cana) in Illinois waters of Lake Michigan. Transac-
tions of the Illinois State Academy of Science 82

(1 & 2):57-S8.

Savitz, J.,S. ARANGO, L.G. BARDYGULA, AND L.
Scoma. 1990. The first records of three fish species
in Illinois waters of Lake Michigan: longear sunfish
(Lepomis megalotis peltastes), black crappie
(Pomoxis nigromaculatus), and the channel catfish
(Ictalurus punctatus). Transactions of the Illinois
State Academy of Science 83(1 & 2):114—115.

Savitz, J., L G. BARDYGULA, AND L. SCoMA. 1989b.
The first record of the quillback carpsucker
(Carpiodes cyprinus) in Illinois waters of Lake
Michigan. Transactions of the Illinois State Academy
of Science 82(3 & 4):1912

SEEGERT, G. 1986. Rediscovery of the greater
redhorse (Moxostoma valenciennesi Jordan)
(Cypriniformes: Catostomidae) in Illinois. Transac-
tions of the Illinois State Academy of Science 79
(3 & 4):293-294.

Symposium Proceedings: Our Living Heritage 427

SKELLY, T.M., AND M.J. SuLE. 1983. The pallid
shiner, Notropis amnis Hubbs and Greene, a rare
Illinois fish. Transactions of the Illinois State
Academy of Science 76(1 & 2):131—140.

Situ, C.L. 1985. The inland fishes of New York
state. New York State Department of Environmental
Education, Albany. 522 p.

Situ, P.W. 1961. The amphibians and reptiles of
Illinois. Illinois Natural History Survey Bulletin 28
(1):1-298.

Situ, P.W. 1965. A preliminary annotated list of the
lampreys and fishes of Illinois. Illinois Natural
History Survey Biological Notes 54. 12 p.

Situ, P.W. 1971. Illinois streams: a classification
based on their fishes and an analysis of factors
responsible for disappearance of native species.
Illinois Natural History Survey Biological Notes 76.
14 p.

Situ, P.W. [1973]. A key to the fishes of Illinois.
Illinois Department of Conservation, Division of
Fisheries, Fishery Bulletin 6. 43 p.

Situ, P.W. 1979. The fishes of Illinois. University
of Illinois Press, Urbana. 314 p.

Sit, P.W., AND L.M. PaGe. 1981. Endangered and
threatened fishes. Pages 5—20 in M. Bowles, ed.
Endangered and threatened vertebrate animals and
vascular plants of Illinois. Illinois Department of
Conservation. 189 p. + vi appendices.

STOECKEL, J.N., AND R.C. HEIDINGER. 1989. Repro-
ductive biology of the inland silverside, Menidia
beryllina in southern Illinois. Transactions of the
Illinois State Academy of Science 82(1 & 2):59-69.

TAYLOR, J.N., W.R. CouRTENAY, JR., AND J.A.
McCann. 1984. Known impacts of exotic fishes in
the continental United States. Pages 322-373 in W.R.
Courtenay, Jr., and J.R. Stauffer, Jr., eds. Distribu-
tion, biology, and management of exotic fishes.
Johns Hopkins University Press, Baltimore.

Warren, M.L., JR. 1989. Geographic variation of the
spotted sunfish, Lepomis punctatus complex (Cen-
trarchidae), with recognition of phylogenetic species.
Ph.D. thesis. Southern Illinois University at Carbon-
dale. 283 p.

Warren, M.L., JR., AND B.M. Burr. 1988. Re-
assessment of the Illinois ranges of the bigeye chub,
Hybopsis amblops, and the pallid shiner, Notropis
amnis. Ohio Journal of Science 88(5):181-183.

WarreEN, M.L., JR., AND B.M. Burr. 1989.
Distribution, abundance, and status of the cypress
minnow, Hybognathus hayi, an endangered Illinois
species. Natural Areas Journal 9(3):163—168.

The Aquatic Mollusca of Illinois

Kevin S. Cummings, Illinois Natural History Survey

Illinois has historically supported a diverse
aquatic molluscan fauna, numbering over 175
species and occupying almost every type of
aquatic habitat from the Great Lakes to wet-
lands, temporary woodland ponds, seeps,
springs, and streams. Two classes of mollusks
are represented in the waters of IIlinois:
Bivalvia, which includes the clams and
mussels, and Gastropoda, represented by the
snails and limpets. The native bivalves of
Illinois are members of three families: the Mar-
garitiferidae and Unionidae (the freshwater
mussels) and the Sphaeriidae (the fingernail
clams and peaclams). The gastropods are
divided into two subclasses, Prosobranchia and
Pulmonata. The Prosobranchs or the opercu-
lated, gill-breathing snails are represented in
Illinois by 37 species in six families. The
Pulmonates or the nonoperculated, lung-
breathing snails contain 37 species in four
families. A list of the species for each of the
families reported from the state is given on
pages 435—438. For the unionids, aspects of
their biology, commercial use, and status are
discussed. Information on identification,
distribution, and biology of the aquatic mollus-
can fauna of Illinois will appear in forthcoming
publications. An excellent monograph on the
freshwater snails of North America has been
published (Burch 1989) and should be con-
sulted for keys and figures of most of the
species found in Illinois.

The list of the freshwater mussels of
Illinois (pages 435—436) is based on the exami-
nation of specimens in collections housed in the
following museums: Academy of Natural
Sciences, Philadelphia; Chicago Academy of
Sciences; Field Museum of Natural History;
Illinois Natural History Survey; Illinois State
Museum; Museum of Comparative Zoology,
Harvard; Ohio State University Museum of
Zoology; University of Illinois Museum of
Natural History; University of Michigan
Museum of Zoology: and the United States

428

National Museum. The list for Sphaeriidae and
Gastropoda (pages 436—438) were compiled
from the literature on Illinois Mollusca,
primarily the publications of Baker (1900,
1901, 1902, 1906, 1922); Basch (1963): Burch
(1989); Dexter (1956): Ulffers (1855); and
Zetek (1918). Additional work is planned to
verify the sphaertid and gastropod lists by
examining specimens in museum collections.
Nomenclature in this paper, with three
exceptions, follows a list of common and
scientific names of mollusks prepared by the
Committee on Scientific and Vernacular Names
of Mollusks of the Council of Systematic
Malacologists, American Malacological Union
(Turgeon et al. 1988). Subspecies are not
recognized, nomenclature for members of the
Pleurobema cordatum species complex follows
Stansbery (1983), and nomenclature for the
family Hydrobiidae follows Hershler and
Thompson (1987) and Hershler et al. (1990).
The aquatic mollusks of Illinois have
been studied for over 150 years. Thomas Say,
the first scientist to work on mollusks in
Illinois, was one of America’s earliest natural-
ists. Say traveled to the Midwest as early as
1817 and in 1826 moved from Philadelphia to
the utopian community of New Harmony,
Indiana (Van Cleave 1951). While there, he
collected and described many of the mollusks
found in the Wabash River and its tributaries,
some of which are are still recognized today.
Few attempts have been made to compile
a list of the mollusk species found in Illinois. In
1906, Frank C. Baker published an annotated
checklist of the Mollusca of Illinois in which he
summarized the available data on the distribu-
tion of the species within the state. A prolific
writer, Baker published over 400 papers,
including many important works on the
molluscan fauna of Illinois (Baker 1897, 1898,
1899, 1900, 1901, 1902, 1906, 1922, 1926).
Baker’s papers remain the best source of
published information on the biology and

April 1991

distribution of aquatic mollusks in the state.
Other early workers on the freshwater mollusks
of Illinois included Kennicott (1855); Ulffers
(1855); Calkins (1874a, 1874b, 1874c); Strode
(1891, 1892): Wilson and Clark (1912);
Danglade (1912, 1914); Zetek (1918); and
Hinkley (1919).

Few papers were published on the aquatic
Mollusca of Illinois in the 1930s and 1940s.
During the late 1940s and 1950s, Dr. Max R.
Matteson of the University of Illinois collected
mussels at over 200 sites in Illinois and
amassed one of the largest and best docu-
mented collections that exists for any state in
the nation. Matteson’s surveys provided both
distribution and abundance data on mussels
from Illinois streams, many of which had not
been previously sampled. His collections, now
at the Illinois Natural History Survey, provide
an invaluable data set and serve as the bench-
mark for mussel surveys conducted today.

In 1967, Paul W. Parmalee of the Illinois
State Museum published The Fresh-water
Mussels of Illinois, which included many
original observations on the distribution and
habitat of unionids. This monograph, one of the
most frequently cited regional works on
freshwater mussels, is still the best guide
available on the mussels of the state. Other
papers on aquatic mollusks of Illinois in the
1950s and 60s include van der Schalie and van
der Schalie (1950); Dexter (1953, 1956);
Parmalee (1955, 1956); Matteson (1961);
Matteson and Dexter (1966); and Fechtner
(1963).

In the 1970s and 1980s, stream surveys
were conducted on the Illinois (Starrett 1971),
Kankakee (Lewis and Brice 1980; Suloway
1981), Kaskaskia (Suloway et al. 1981), and
Wabash rivers (Meyer 1974; Clark 1976).
These and current studies document the rapid
decline of the freshwater mussels of Illinois and
provide data on the status of rare species.

BIVALVIA: MUSSELS AND CLAMS

Freshwater mussels in the families Margariti-
feridae and Unionidae are found throughout the
holarctic region but reach their greatest
diversity in eastern North America, where they
number about 285 species (Turgeon et al.
1988). A total of 78 species in two families and
four subfamilies has been recorded from
Illinois and boundary waters (pages 435—436).

Symposium Proceedings: Our Living Heritage

429

Biology. Mussels filter-feed on plankton,
which they remove from the water as it
circulates through the animal via incurrent and
excurrent aperatures. In most freshwater mussel
species, the sexes are separate. Sperm are
released into the water and taken into the
female via the incurrent aperature. The eggs are
fertilized and develop into an intermediate
stage, the glochidium. Glochidia are stored in
the female’s gills, which function as brood
chambers. Nearly all unionids must pass
through a parasitic phase in order to complete
their life cycle. In the spring or summer,
glochidia are expelled into the water and must
come in contact with the appropriate host,
usually a fish, to which they attach and
metamorphose into a juvenile mussel.
Glochidia are either internal parasites on the
gills or external parasites on the fins. Some
species are host specific, but others are general-
ists and use a wide variety of fishes as hosts.
Mussels are long lived. Many species live as
long as 25 years, and some are reported to live
more than 50 years.

Commercial Use. In 1891 a German
immigrant, J.F. Boepple of Petersburg, Illinois,
realized that the mussels of the United States
could be used, as they had been in Europe, to
manufacture buttons. In the early part of the
twentieth century, enormous quantities of
mussels were harvested for the button industry,
with some beds in Illinois producing over 700
tons in a single year (Coker 1919). Mussel
shells were collected, cooked out, and shipped
to factories where they were cut into blanks,
sorted, polished, and finished into buttons.
Today freshwater mussel shells are exported to
Japan where they are converted into beads and
inserted into oysters where they serve as nuclei
for cultured pearls. The oysters are maintained
in cages under water, and over a period of
about a year, a layer of mother-of-pearl is
secreted around the bead to form the pearl.

From 1912 to 1914, roughly 15,000 tons
of shells were taken in Illinois and boundary
waters and sold at a price that varied from $4 to
$10 a ton. The increase in price over the last 75
years has been astronomical. In the 1940s, the
price of shells was about $25 a ton and re-
mained at that level until the button industry
collapsed in the late 1950s due to the advent of
plastics. As the demand for shells to manufac-
ture cultured pearls increased, so did the price,
from $45 a ton in the 60s, $800 in the 70s, and

430

$1,800 in the 80s, to $2,400 a ton this year

(N. Cohen, pers. comm.). At current prices, the
estimated harvest of 1912 to 1914 would be
worth about $36 million.

Status. Surveys across North America
have documented significant declines in
freshwater mussel populations. Recent surveys
for mussels in Illinois using the same methods
as those of previous studies have documented a
reduction in the fauna for all streams sampled
(Table 1). In 1966, William C. Starrett of the
Illinois Natural History Survey conducted an
in-depth study of the Illinois River. He col-
lected only 23 of the 47 species previously
reported from the Illinois (Starrett 1971). Two
of the 24 extirpated species were the butterfly,
Ellipsaria lineolata (Rafinesque 1820), a
species that has declined statewide in recent
years; and the Higgins eye, Lampsilis higginsi
(Lea 1857), now on the federally endangered
species list. Similar results were obtained in the
Kankakee River where Suloway (1981)
reported only 24 of the 32 species historically
known to inhabit the river. The Kankakee River
drainage continues to support some of the
richest mussel populations of the state, includ-
ing the state threatened bullhead, Plethobasus
cyphyus (Rafinesque 1820), and the ellipse,
Venustaconcha ellipsiformis (Conrad 1836). In
the Kaskaskia River, the decline in diversity
has been pronounced. Only 32 of the 39 species
recorded from the drainage were found in 1956,
and that number was reduced to 24 by 1978
(Suloway et al. 1981). In addition, the number
of individuals dropped from 2,595 to 498, an
80% reduction in just over 20 years. A survey
of the Sangamon River in 1988—1989 recoy-

Table 1. Selected streams in Illinois where recent
surveys have documented declines in the freshwater
mussel fauna. Data from Starrett 1971; Suloway et
al. 1981; Suloway 1981; and Cummings et al. un-
published.

Number of mussel species

Pre-1960 Post-1960

Mississippi River drainage

Illinois River 47 23

Kaskaskia River 39 24

Kankakee River 32 24
Wabash River drainage

Embarras River 44 27

Vermilion River 41 25

Little Wabash River 41 31

Illinois Natural History Survey Bulletin

Vol. 34 Art. 4

ered all of the species found in 1956-1960:
however, overall numbers collected per unit of
effort were much lower, and some, for
example, Elliptio dilatata (Rafinesque 1820)
and Megalonaias nervosa (Rafinesque 1820)
have been nearly extirpated (Schanzle and
Cummings 1991).

In the Wabash River drainage, even the
relatively undisturbed Vermilion River has
suffered a serious decline, with almost 40% of
the mussel species extirpated by the 1970s.
Although its species richness has declined, this
river supports the only known populations of at
least two state endangered species: the wavy-
rayed lampmussel, Lampsilis fasciola Raf-
inesque 1820, and the rabbitsfoot, Quadrula
cylindrica (Say 1817). The pattern is the same
in the Embarras River, where the number of
species has dropped from 44 to 27. A compari-
son of surveys done in 1956 and 1986 revealed
that the Embarras River continues to support a
fairly diverse fauna; however, the number of
individuals has declined over 80% in the last 30
years. Two state endangered species are found
in Illinois only in the Embarras: the
kidneyshell, Ptychobranchus fasciolaris
(Rafinesque 1820), and the snuffbox, Epiob-
lasma triquetra (Rafinesque 1820). The Little
Wabash River has suffered a similar decline,
and a 1988 survey revealed that only 31 of the
41 species known to have occurred in the
drainage are extant.

A variety of factors are responsible for
the decline of mussel populations. Foremost is
siltation from agricultural run-off due to poor
land management. Mussels are sedentary and
particularly susceptible to the smothering
effects of siltation. Channelization is detrimen-
tal because it eliminates habitat for mussels as
well as potential host fishes. Impoundments
often create good habitat directly below the
dam, but they also inundate large areas of the
stream and impede the migration of host
species. Herbicides, pesticides, and petroleum-
related pollution also have negative effects,
and competition from exotics has been impli-
cated in the decline of native mussels, although
the mechanisms involved are not entirely
understood.

One result of the status surveys con-
ducted in Illinois and other states in recent
years has been the addition of many mussel
species to state and federally endangered
species lists. Thirteen species are now consid-

April 1991

ered to be globally extinct, including four once
found in Illinois (Turgeon et al. 1988; see
listing on pages 435—436, this publication). On
the federal level, 37 mussels are listed as en-
dangered and another 56 are proposed or candi-
dates for listing (U.S. Department of the
Interior, Fish and Wildlife Service 1989a,
1989b). The Illinois Threatened and Endan-
gered Species List now contains 33 mussels
(29 endangered and 4 threatened), slightly over
40% of the species ever recorded from Illinois
(Illinois Endangered Species Protection Board
1990). Another 11 species are candidates or
species of special concern that may be listed in
the future. These bring the total number of rare,
endangered, or extirpated species in Illinois to
44 species—56% of the state’s known mussel
fauna. Other states have similar problems.
North Carolina, for example, recently reported
that half of its mussel species are disappearing
and in need of protection (Venters 1990). This
national decline has received some much
needed attention and funding has been provided
in recent years to begin to document and
address the problem.

The fingernail clams and peaclams of the
family Sphaeriidae are holarctic in distribution
and occupy a wide variety of habitats. Thirty-
eight species in four genera are found in North
America, and 26 species in three genera are
reported from Illinois (pages 436— 437). Al-
though little has been published on the distribu-
tion and status of these animals in Illinois since
Baker’s list of 1906, unpublished reports make
clear that many species have disappeared from
the streams in which they formerly occurred
and are declining throughout their range.
Sphaeriids are hermaphroditic and, unlike
freshwater mussels, have direct development,
with about 2 to 20 young produced per female.
Although sphaeriids have no direct economic
value, they are an important food source for
many animals, including fishes and diving
ducks.

The family Corbiculidae is represented in
Illinois by the exotic Asian Clam, Corbicula
fluminea (Miller 1774). Introduced in North
American in the 1920s (Counts 1981), this
species was first reposted in Illinois from the
Ohio River in southern Illinois in the early
1960s (Fechtner 1962). Since then it has spread
at least as far north as Rock Island and is
present in most if not all drainages in the state.

Symposium Proceedings: Our Living Heritage 431

As is the case with most established exotics,
Corbicula has had serious negative effects on
the environment. This extremely prolific clam
has caused major problems associated with the
fouling of cooling water intakes of power
plants (Isom 1986) and may outcompete native
species (Clarke 1988).

The family Dreissenidae is represented in
North American freshwaters by the zebra
mussel Dreissena polymorpha (Pallas 1771).
Although the zebra mussel is not currently
established in Illinois waters, it was recently
discovered in the Indiana portion of Lake
Michigan and its arrival here is imminent. This
exotic is causing tremendous economic
problems in Lake Erie and Lake St. Clair and
will negatively affect our native mussels by
smothering and suffocating them as it has in the
Great Lakes.

GASTROPODA: FRESHWATER SNAILS

Freshwater snails are basically herbivores and
detritivores and use their radulae to scrape
algae and diatoms from plants and rocks. About
500 species of freshwater snails are found in
North America, 350 Prosobranchs and 150
Pulmonates (Burch 1989). Of those, 85 or
about one-fifth of the species are candidates for
federal protection (U.S. Department of Interior,
Fish and Wildlife Service 1989b). A review of
the literature suggests that there are or were
about 74 species of freshwater snails in Illinois,
two of which were introduced and three that are
under consideration for federal listing (pages
437-438).

The subclass Prosobranchia is repre-
sented in Illinois by 37 species in six families:
Valvatidae, Viviparide, Bithyniidae, Hydro-
biidae, Pomatiopsidae, and Pleuroceridae.

The shells of North American Valvatidae
are relatively small (up to 5 mm) and flattened
in shape. Valvatids are egg layers and, unlike
most Prosobranchs, hermaphroditic. Five
species, all in the genus Valvata, have been
reported from Illinois.

The family Viviparidae is found on all
continents except Antarctica and South
America and occurs throughout eastern North
America. The sexes are separate, and as their
name implies, they are “live bearers” as
opposed to egg layers. Six species in three
genera are found in Illinois.

432

The family Bithyniidae is represented in
Illinois by the Mud Bithynia, Bithynia tentacu-
lata (Linnaeus 1758). This species also occurs
in Europe, and populations have been intro-
duced into North America where the species
has spread widely (Burch 1989). Bithynia ten-
taculata has been reported from Pleistocene
deposits in Chicago, and it may, therefore, have
been present in North America before Europe-
ans arrived.

The family Hydrobiidae is one of the
most common and widely distributed snail
families in the world. These small- to medium-
sized snails are a major component of the North
American fauna and number about 35 genera
and 170 species (Hershler and Thompson 1987;
Turgeon et al. 1988). Most live in fresh water,
although a few have been found in brackish
water. Twelve species in seven genera have
been reported from Illinois.

The family Pomatiopsidae is represented
in North America by six species, two of which
are found in Illinois. These snails are usually
regarded as amphibious, inhabiting river banks
or moist areas near streams.

The Pleuroceridae are widely distributed,
occurring in North, Central, and South America
and in Africa and Asia. They reach their
greatest diversity, however, in the southeastern
United States. Pleurocerids are extremely
sensitive to the effects of pollution and silta-
tion. At least 23 species are presumed extinct,
and many others are candidates for threatened
or endangered status (Turgeon et al. 1988; U.S.
Department of the Interior, Fish and Wildlife
Service 1989b). Eleven species in four genera
have been found in Illinois, three of which are
candidates for federal listing (page 437). Their
current status in Illinois is unknown and needs
investigation.

The subclass Pulmonata is represented in
Illinois by four families. Like the pleurocerids,
members of the family Lymnaeidae are found

‘worldwide but reach their greatest diversity in
North America. Fourteen species (1 introduced)
in six genera have been reported from Illinois.

The family Physidae is mainly a New
World family with a few species found in
Eurasia and Africa. Physids are found in a wide
variety of habitats and are the most widespread
and abundant snails in North America. They
appear to be the most pollution tolerant of all
freshwater mollusks and may be the only
species found in highly degraded waters.

Illinois Natural History Survey Bulletin

Vol. 34 Art. 4

The family Planorbidae is restricted to
fresh water and is worldwide in distribution.
Planorbids vary widely in size from about | to
30 mm. A few species are known to serve as
intermediate hosts for human parasites and
have been studied extensively; most others are
relatively unknown ecologically. Twelve
species (1 introduced) in six genera have been
found in Illinois.

The Ancylidae, or freshwater limpets, are
worldwide in distribution and are found in
many freshwater habitats. The family, revised
in 1963, is currently thought to contain about
13 species in four genera (Basch 1963; Turgeon
et al. 1988). Ancylids can usually be found
attached to aquatic vegetation or living on
stones or other debris. Little is known about the
biology of freshwater limpets, but they are
reported to be fairly intolerant of chemical
pollution (Basch 1963). Six species in three
genera have been found in Illinois.

The current distribution and status of
gastropods in Illinois are poorly understood,
and as a result we are unable to compile a list of
threatened or endangered freshwater snail
species for the state. Given the documented
decline in freshwater mussels and other aquatic
organisms, however, there can be little doubt
that Illinois has lost and is likely in danger of
losing many species of snails as well.

Conservation efforts in Illinois and other
states have thus far concentrated on preserving
or protecting terrestrial ecosystems and their
inhabitants. While the protection of prairies,
bogs, fens, glades, and forests is an extremely
important and worthwhile endeavor, we need to
protect aquatic habitats as well or we will most
certainly lose many of the fascinating and
unique species that are found in the fresh
waters of North America.

ACKNOWLEDGMENTS

I would like to thank the following curators and
collection managers for allowing me access to
collections under their care: Arthur E. Bogan
and George M. Davis, The Academy of Natural
Sciences of Philadelphia; Ron Vasile, The
Chicago Academy of Science; Margaret Baker
and the late Alan Solem, Field Museum of
Natural History; Tim Cashatt, Illinois State
Museum; Kenneth Boss, Silvard P. Kool, and
Richard I. Johnson, Museum of Comparative
Zoology, Harvard University; John B. Burch,

April 1991

Douglas J. Eernisse, and Walter R. Hoeh,
University of Michigan Museum of Zoology;
David H. Stansbery and Kathy G. Borror, Ohio
State University Museum of Zoology; Robert
Hershler, U.S. National Museum; and Lowell
Getz and Thomas Uzzell, University of Illinois
Museum of Natural History. Robert W.
Schanzle, Illinois Department of Conservation,
and Carol Stein, Ohio State University Museum
of Zoology, provided helpful comments on the
manuscript.

LITERATURE CITED

Baker, F.C. 1897. On a collection of mollusks from
Grand Tower, Illinois. Nautilus 11(3):28—30.

Baker, F.C. 1898. The Mollusca of the Chicago
area. Part 1: The Pelecypoda. Chicago Academy of
Sciences Bulletin 3(1):1—130.

BAKER, F.C. 1899. Notes on the mollusks of
Lilycash Creek. Nautilus 12(3):30-31.

Baker, F.C. 1900. A revision of the Physae of
northeastern Illinois. Nautilus 14(2):16—24.

Baker, F.C. 1901. A revision of the Limnaeas of
northern Illinois. Transactions of the Academy of
Science of St. Louis 11(1):1-24 + 1 plate.

Baker, F.C. 1902. The Mollusca of the Chicago
area. Part 2: The Gastropoda. Chicago Academy of
Sciences Bulletin 3(2):131—418 + 9 plates.

Baker, F.C. 1906. A catalogue of the Mollusca of
Illinois. Bulletin of the Illinois State Laboratory of
Natural History 7(6):53-136 + 1 map.

Baker, F.C. 1922. The molluscan fauna of the Big
Vermilion River, Illinois. Illinois Biological
Monographs 7(2):105—224 + 15 plates.

Baker, F.C. 1926. The naiad fauna of the Rock
River system: a study of the law of stream distribu-
tion. Transactions of the Illinois State Academy of
Science 19:103-112.

Bascu, P.F. 1963. A review of the recent freshwater
limpet snails of North America (Mollusca: Pulmon-
ata). Bulletin of the Museum of Comparative
Zoology, Harvard University 129(8):399—461.

Burcu, J.B. 1989. North American freshwater
snails. Malacological Publications, Hamburg, MI.
Vili + 365 p.

CALKINS, W.W. 1874a. The land and fresh water
shells.of LaSalle County, Ills. Proceedings of the
Ottawa Academy of Science. 48 p. + 1 plate.

Symposium Proceedings: Our Living Heritage

433

CaLkins, W.W. 1874b. Notes on freshwater
Mollusca, found in the vicinity of Chicago, Illinois.
Cincinnati Quarterly Journal of Science 1:242—244.

CALKINS, W.W. 1874c. Notes on the molluscan fauna
of northern Illinois. Cincinnati Quarterly Journal of
Science 1:321-325.

CLarK, C.F. 1976. The freshwater naiads of the
lower end of the Wabash River, Mt. Carmel, Illinois,
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434

Isom, B.G. 1986. Historical review of Asiatic clam
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April 1991 Symposium Proceedings: Our Living Heritage

The Aquatic Mollusca of Illinois. Species are arranged alphabetically within each family or in the case of
Unionidae within each subfamily. Abbreviations for status are as follows: (+) = extinct, X = extirpated from
Illinois, FE = federally endangered, FC = federal candidate, SE = state endangered, ST = state threatened,

SC = state candidate (watch list), I = introduced.

Scientific Name

CLASS BIVALVIA
OrDER UNIONOIDA

Family Margaritiferidae (1 species)
Subfamily Cumberlandinae
Cumberlandia monodonta (Say 1829)

Family Unionidae (77 species)

Subfamily Ambleminae
Amblema plicata (Say 1817)
Cyclonaias tuberculata (Rafinesque 1820)
Elliptio crassidens (Lamarck 1819)
Elliptio dilatata (Rafinesque 1820)
Fusconaia ebena (Lea 1831)
Fusconaia flava (Rafinesque 1820)
Fusconaia subrotunda (Lea 1831)
Hemistena lata (Rafinesque 1820)
Megalonaias nervosa (Rafinesque 1820)
Plethobasus cicatricosus (Say 1829)
Plethobasus cooperianus (Lea 1834)
Plethobasus cyphyus (Rafinesque 1820)
Pleurobema clava (Lamarck 1819)
Pleurobema cordatum (Rafinesque 1820)
Pleurobema plenum (Lea 1840)
Pleurobema rubrum (Rafinesque 1820)
Pleurobema sintoxia (Rafinesque 1820)
Quadrula cylindrica (Say 1817)
Quadrula fragosa (Conrad 1835)
Quadrula metanevra (Rafinesque 1820)
Quadrula nodulata (Rafinesque 1820)
Quadrula pustulosa (Lea 1831)
Quadrula quadrula (Rafinesque 1820)
Tritogonia verrucosa (Rafinesque 1820)
Uniomerus tetralasmus (Say 1831)

Subfamily Anodontinae
Alasmidonta marginata Say 1818
Alasmidonta viridis (Rafinesque 1820)
Anodonta grandis Say 1829
Anodonta imbecillis Say 1829
Anodonta suborbiculata Say 1831
Anodontoides ferussacianus (Lea 1834)
Arcidens confragosus (Say 1829)
Lasmigona complanata (Barnes 1823)
Lasmigona compressa (Lea 1829)
Lasmigona costata (Rafinesque 1820)
Simpsonaias ambigua (Say 1825)
Strophitus undulatus (Say 1817)

Subfamily Lampsilinae
Actinonaias ligamentina (Lamarck 1819)
Cyprogenia stegaria (Rafinesque 1820)
Ellipsaria lineolata (Rafinesque 1820)
Epioblasma flexuosa (Rafinesque 1820)
Epioblasma obliquata (Rafinesque 1820)
Epioblasma personata (Say 1829)

Common Name

Spectaclecase

Threeridge

Purple wartyback
Elephant-ear
Spike

Ebonyshell
Wabash pigtoe
Long-solid
Cracking pearlymussel
Washboard

White wartyback
Orange-foot pimpleback
Sheepnose
Clubshell

Ohio pigtoe
Rough pigtoe
Pyramid pigtoe
Round pigtoe
Rabbitsfoot
Winged mapleleaf
Monkeyface
Wartyback
Pimpleback
Mapleleaf
Pistolgrip
Pondhorn

Elktoe

Slippershell

Giant floater
Paper pondshell
Flat floater
Cylindrical papershell
Rock-pocketbook
White heelsplitter
Creek heelsplitter
Fluted-shell
Salamander mussel
Squawfoot

Mucket

Fanshell
Butterfly
Leafshell
Catspaw

Round combshell

Status!

FGISE

ST
Se
Se

Ee SCex
FE, SE, X

RE, SE ox
FE, SE
ST

FC, SE
SC

FE, SE, X
SC

SE
FC, SC, X

Sm

SE

Su

FC, SE

436 F Illinois Natural History Survey Bulletin Vol. 34 Art. 4

Scientific Name Common Name Status'
Epioblasma propinqua (Lea 1857) Tennessee riffleshell (+). SE. X
Epioblasma rangiana (Lea 1839) Northern riffleshell EGC, SC>:
Epioblasma sampsonii (Lea 1861) Wabash riffleshell (+), SE. X
Epioblasma torulosa (Rafinesque 1820) Tubercled blossom FE, SE{%
Epioblasma triquetra (Rafinesque 1820) Snuffbox SE
Lampsilis abrupta (Say 1831) Pink mucket EE, SEs
Lampsilis cardium Rafinesque 1820 Plain pocketbook
Lampsilis fasciola Rafinesque 1820 Wavy-rayed lampmussel SE
Lampsilis higginsi (Lea 1857) Higgins eye FE, SE
Lampsilis ovata (Say 1817) Pocketbook sc
Lampsilis siliquoidea (Barnes 1823) Fatmucket
Lampsilis teres (Rafinesque 1820) Yellow sandshell
Leptodea fragilis (Rafinesque 1820) Fragile papershell
Leptodea leptodon (Rafinesque 1820) Scaleshell FC, SE, X
Ligumia recta (Lamarck 1819) Black sandshell
Ligumia subrostrata (Say 1831) Pondmussel
Obliquaria reflexa Rafinesque 1820 Threehorn wartyback
Obovaria olivaria (Rafinesque 1820) Hickorynut
Obovaria retusa (Lamarck 1819) Ring Pink FE, SE, X
Obovaria subrotunda (Rafinesque 1820) Round hickorynut SE
Potamilus alatus (Say 1817) Pink heelsplitter
Potamilus capax (Green 1832) Fat pocketbook FE, SE
Potamilus ohiensis (Rafinesque 1820) Pink papershell
Potamilus purpuratus (Lamarck 1819) Bleufer sc
Ptychobranchus fasciolaris (Rafinesque 1820) Kidneyshell SE
Toxolasma lividus (Rafinesque 1831) Purple lilliput FC. SE
Toxolasma parvus (Barnes 1823) Lilliput
Toxolasma texasensis (Lea 1857) Texas lilliput
Truncilla donaciformis (Lea 1828) Fawnsfoot
Truncilla truncata Rafinesque 1820 Deertoe
Venustaconcha ellipsiformis (Conrad 1836) Ellipse sc
Villosa fabalis (Lea 1831) Rayed bean FC, SE, X
Villosa iris (Lea 1829) Rainbow SE
Villosa lienosa (Conrad 1834) Little spectaclecase SE

ORDER VENEROIDA

Family Sphaeriidae (26 species)
Musculium lacustre (Miller 1774)
Musculium partumeium (Say 1822)
Musculium securis (Prime 1852)
Musculium transversum (Say 1829)
Pisidium adamsi Prime 1851
Pisidium casertanum (Poli 1791)
Pisidium compressum Prime 1852
Pisidium conventus Clessin 1877
Pisidium cruciatum Sterki 1895
Pisidium dubium (Say 1817)
Pisidium equilaterale Prime 1852
Pisidium fallax Sterki 1896
Pisidium ferrugineum Prime 1852
Pisidium idahoense Roper 1890
Pisidium lilljeborgi (Clessin 1886)
Pisidium nitidum Jenyns 1832
Pisidium punctatum Sterki 1895
Pisidium punctiferum (Guppy 1867)
Pisidium rotundatum Prime 1852
Pisidium variabile Prime 1852
Pisidium walkeri Sterki 1895
Sphaerium fabale (Prime 1852)

Lake fingernailclam
Swamp fingernailclam
Pond fingernailclam
Long fingernailclam
Adam peaclam
Ubiquitous peaclam
Ridged-beak peaclam
Alpine peaclam
Omamented peaclam
Greater eastern peaclam
Round peaclam

River peaclam

Rusty peaclam

Giant northern peaclam
Lilljeborg peaclam
Shiny peaclam
Perforated peaclam
Striate peaclam

Fat peaclam
Triangular peaclam
Walker peaclam

River fingernailclam

April 1991 Symposium Proceedings: Our Living Heritage 437

Scientific Name Common Name Status!

Sphaerium occidentale (Lewis 1856)
Sphaerium rhomboideum (Say 1822)
Sphaerium simile (Say 1817)
Sphaerium striatinum (Lamarck 1818)

Family Corbiculidae (1 species)
Corbicula fluminea (Miiller 1774)

Family Dreissenidae (1 species)
Dreissena polymorpha (Pallas 1771)

CLASS GASTROPODA (74 species)
SUBCLASS PROSOBRANCHIA
OrDER MESOGASTROPODA

Family Valvatidae (5 species)
Valvata bicarinata Lea 1841
Valvata lewisi Currier 1868
Valvata perdepressa Walker 1906
Valvata sincera Say 1824
Valvata tricarinata (Say 1817)

Family Viviparidae (6 species)
Campeloma crassulum Rafinesque 1819
Campeloma decisum (Say 1817)
Lioplax sulculosa (Menke 1827)
Viviparus georgianus (Lea 1834)
Viviparus intertextus (Say 1829)
Viviparus subpurpureus (Say 1829)

Family Bithyniidae (1 species)
Bithynia tentaculata (Linnaeus 1758)

Family Hydrobiidae (12 species)
Amnicola limosa (Say 1817)
Amunicola pilsbryi Walker 1906
Amnicola walkeri Pilsbry 1898
Birgella subglobosus (Say 1825)
Fontigens aldrichi (Call & Beecher 1886)
Fontigens antroecetes (Hubricht 1940)
Fontigens nickliniana (Lea 1838)
Hoyia sheldoni (Pilsbry 1890)

Herrington fingernailclam
Rhomboid fingernailclam
Grooved fingernailclam
Striated fingernailclam

Asian clam

Zebra mussel

Two-ridge valvata
Fringed valvata
Purplecap valvata
Mossy valvata
Threeridge valvata

Ponderous campeloma
Pointed campeloma
Furrowed lioplax
Banded mysterysnail
Rotund mysterysnail
Olive mysterysnail

Mud bithynia

Mud amnicola

* Lake duskysnail

Canadian duskysnail
Globe siltsnail
Hoosier amnicola

Watercress snail
Storm hydrobe

Probythinella lacustris (Baker 1928) Delta hydrobe
Pyrgulopsis lustrica (Pilsbry 1890) Boreal marstonia
Pyrgulopsis scalariformis (Wolf 1870) Moss pyrg
Somatogyrus depressus (Tryon 1862) Sandbar pebblesnail

Family Pomatiopsidae (2 species)
Pomatiopsis cincinnatiensis (Lea 1840)
Pomatiopsis lapidaria (Say 1817)

Family Pleuroceridae (11 species)
Elimia costifera (Reeve 1861)
Elimia livescens (Menke 1830)
Elimia semicarinata (Say 1829)

Brown walker
Slender walker

Corded elimia
Liver elimia
Fine-ridged elimia

Leptoxis praerosa (Say 1821) Onyx rocksnail FC, SC
Leptoxis trilineata (Say 1829) Broad mudalia

Lithasia armigera (Say 1821) Armored rocksnail FC, SC
Lithasia obovata (Say 1829) Shawnee rocksnail

Lithasia verrucosa (Rafinesque 1820) Verrucose rocksnail FC)SC

Pleurocera acuta Rafinesque 1831
Pleurocera alveare (Conrad 1834)
Pleurocera canaliculata (Say 1821)

Sharp hornsnail
Rugged hornsnail
Silty hornsnail

438 Illinois Natural History Survey Bulletin Vol. 34 Art. 4

Scientific Name

Common Name Status '

SUBCLASS PULMONATA
ORDER BASOMMATOPHORA

Family Lymnaeidae (14 species)
Acella haldemani (Binney 1867)
Fossaria dalli (Baker 1907)
Fossaria humilis (Say 1822)
Fossaria obrussa (Say 1825)
Fossaria parva (Lea 1841)
Fossaria tazewelliana (Wolf 1870)
Lymnaea stagnalis Linnaeus 1758
Pseudosuccinea columella (Say 1817)
Radix auricularia (Linnaeus 1758)
Stagnicola caperatus (Say 1829)
Stagnicola catascopium (Say 1817)
Stagnicola elodes (Say 1821)
Stagnicola exilis (Lea 1834)
Stagnicola woodruffi (Baker 1901)

Family Physidae (5 species)
Aplexa elongata (Say 1821)
Physella gyrina (Say 1821)
Physella heterostropha (Say 1817)
Physella integra (Haldeman 1841)
Physella virgata (Gould 1855)

Family Planorbidae (12 species)
Biomphalaria glabrata (Say 1818)
Gyraulus deflectus (Say 1824)
Gyraulus parvus (Say 1817)

Helisoma anceps (Menke 1830)
Micromenetus dilatatus (Gould 1841)
Micromenetus sampsoni (Ancey 1885)
Planorbella armigera (Say 1821)
Planorbella campanulata (Say 1821)
Planorbella pseudotrivolvis (Baker 1920)
Planorbella trivolvis (Say 1817)
Planorbella truncata (Miles 1861)
Promenetus exacuous (Say 1821)

Family Ancylidae (6 species)
Ferrissia fragilis (Tryon 1863)
Ferrissia parallela (Haldeman 1841)
Ferrissia rivularis (Say 1817)
Laevapex diaphanus (Haldeman 1841)
Laevapex fuscus (Adams 1840)
Rhodacmea hinkleyi (Walker 1908)

Spindle lymnaea
Dusky fossaria

Marsh fossaria

Golden fossaria
Pygmy fossaria
Tazewell fossaria
Swamp lymnaea
Mimic lymnaea
Big-ear radix I
Wrinkled marshsnail
Woodland pondsnail
Marsh pondsnail
Flat-whorled pondsnail
Coldwater pondsnail

Lance aplexa
Tadpole physa
Pewter physa
Ashy physa
Protean physa

Bloodfluke planorb I
Flexed gyro

Ash gyro

Two-ridge rams-horn

Bugle sprite

Thicklip rams-horn
Bellmouth rams-horn

Marsh rams-hom
Druid rams-horn
Sharp sprite

Fragile ancylid
Oblong ancylid
Creeping ancylid
Cymbal ancylid
Dusky ancylid
Knobby ancylid

' Readers may be puzzled by such dual designations for a species as endangered and extirpated or endangered and extinct.
The current Illinois list of threatened and endangered mussels was compiled in 1987. Since that time, surveys have
determined that some of the species on that list are probably no longer extant. Future lists will reflect such changes and
species thought to be extirpated or extinct will be removed. At the present time, however, a species may continue to be
listed as endangered but considered by researchers to be extirpated or extinct.

Streams of Illinois

Lawrence M. Page, Illinois Natural History Survey

The recent increased interest in protecting
streams (Phillippi and Anderson 1989) is an
extremely welcome development. Until now,
little effort has been directed toward protecting
flowing bodies of water in Illinois, largely
because of the difficulties of the task. In
contrast, completion of a natural areas inven-
tory in Illinois and excellent efforts by the
Illinois Nature Preserves Commission and The
Nature Conservancy have resulted in safe-
guarding a number of prairies and other
terrestrial ecosystems.

To protect our streams, we need to gather
data and develop appropriate methodologies.
To organize this process, we need to address
the following questions in relation to streams:
What does Illinois have? What should we
protect? What are the major causes of stream
degradation? How do we protect streams?

WHAT DOES ILLINOIS HAVE?

Because Illinois has a large and complex
drainage pattern (Figure 1), it is considered a
well-watered state, particularly in relation to
most western states. It is bounded on the west
by the Mississippi River, on the south by the
Ohio, on the northeast by Lake Michigan, and
on the southeast by the Wabash. An excellent
discussion of the drainages of Illinois and their
characteristics at the turn of the century was
undertaken by C.W. Rolfe in Forbes and
Richardson’s The Fishes of Illinois [1908]. The
biogeography of the fishes of Illinois and other
states of the lower Ohio and upper Mississippi
River basins is discussed by Burr and Page
(1986).

The geological characteristics of Illinois
strongly influence the diversity and distribu-
tions of its aquatic biota, and the streams of
Illinois can be classified physiographically
according to Fenneman’s physiographic
provinces (Fenneman 1938):

I. Great Lakes: Lake Michigan Section
II. Mississippi River
A. Wisconsin Driftless Section
B. Till Plains Section
1. Wisconsin Glacial Till
2. Illinoian Glacial Till
C. Shawnee Hills—Ozark Plateaus Section
D. Coastal Plain Section

The streams over most of Illinois are
relatively recent products of glaciation. Those
flowing into Lake Michigan and those on the
Till Plains Section developed after Pleistocene
glaciers had receded and are less than 100,000
years old; those north of the Shelbyville
moraine, the southern terminus of the Wiscon-
sin glaciation, are less than 10,000 years old. In
contrast, streams in the unglaciated areas of
Illinois—the Wisconsin Driftless, Shawnee
Hills, and Coastal Plain sections—traverse
much older areas. Unglaciated areas exhibit
more topographic relief and have more bed-
rock; their streams are characterized by higher
gradients, and they often sustain unique aquatic
communities.

The Illinois portion of the Wisconsin
Driftless Section is found mostly in Jo Daviess
County. It escaped glaciation, and the streams
there are the product of millions of years of
geological evolution. Relict populations of
species otherwise eliminated from Illinois by
the glaciers (e.g., the Ozark minnow, Notropis
nubilus) remain there. The major stream of the
area is the Apple River.

The Till Plains Section is the vast area of
the state covered during the Pleistocene by one
or more glacial advances. During glaciation,
old river channels were filled with glacial drift.
As the glaciers receded, drift was laid down in
ridges that acted as dams holding back melt-
water and creating large lakes. Later, over long
periods of time, the lakes filled with deposi-
tional materials, drainage outlets formed in the
moraines, and the lakes transformed into
marshes and prairies. Water flowing through

439

440 Illinois Natural History Survey Bulletin

the marshes and prairies eventually cut the
drainage patterns that exist today. Nearly the
entire region covered by glacial till (Till Plains
Section) is drained by tributaries flowing
southwest into the Mississippi River (mainly,
the Rock, Illinois, Kaskaskia, and Big Muddy
rivers) and by tributaries flowing southeast into
the Wabash and Ohio rivers (the Vermilion,
Embarras, Little Wabash, and Saline rivers).

The Shawnee Hills are composed almost
entirely of Mississippian limestone and
sandstone and stand an average of about 400
feet above the surrounding land. Several of the
most interesting streams and aquatic organisms,
including species endemic to Illinois, such as
the Illinois crayfish (Orconectes illinoiensis),
occur in this region. The streams of the
Shawnee Hills—including Big, Lusk (Figure
2), Big Grand Pierre, and Clear creeks—are
small, clear rocky streams that are among the
most scenic in the state.

issipp,;

miss

Figure 1. Major streams of Illinois.

Vol. 34 Ant. 4

The Coastal Plain lies south of the
Shawnee Hills. Flat, sandy, and covered by
residual soils, it is drained almost entirely by
the Cache River and small tributaries of the
Ohio. Aquatic organisms found on the IIlinois
Coastal Plain tend to be restricted to this region
in Illinois, although they are also characteristic
of the Coastal Plain to the south of Illinois. Be-
cause the Illinois portion of the Coastal Plain is
small, many species found there are rare and
restricted and therefore protected in Illinois.

The present character of the streams of
Illinois is as much a function of human
activities as it is of the evolution of drainage
patterns. What we have done to the streams in
the last 200 years has had a major impact on
the distributional patterns and community
structure established during the millions of
years of geological history that preceded our
arrival. The questions now are, what does
Illinois have left and what should be protected
and from what?

Lusk Creek Canyon, Pope County, Illinois.

Figure 2.
Photo by Michael Jeffords.

April 1991

WHAT SHOULD WE PROTECT?

An element of scenic beauty apart from living
organisms is certainly worth preserving, but
generally we are interested in protecting life. In
deciding what to protect, we can concentrate on
biodiversity. The species that remain are of
interest and of value to us for a number of
reasons, and it seems clear that we as a society,
through the establishment of environmental
protection agencies and endangered species
lists, have stated emphatically that we want to
protect them. The reasons for protecting species
include vital as well as aesthetic and economic
considerations. Living organisms provide the
oxygen we breathe and the food we eat and are
the source of many of our medicines. We enjoy
the beauty and diversity of life and acknowl-
edge that our lives without wild places and wild
plants and animals would be much less
interesting and enjoyable. By maintaining a
diversity of plants and animals, we are also
maintaining a variety of choices for the
biological control of noxious species; surely
that option is more likely to result in a healthy
environment than is resorting to potentially
dangerous pesticides.

Because of the enormous modifications
of the Illinois landscape, we are faced with
protecting large numbers of species. Our
present list of endangered and threatened
animals and plants includes nearly 500 species.
In addition to these, which are considered to be
in risk of extirpation from the state, thousands
of others have disappeared or declined signifi-
cantly in abundance in the past 200 years. Ina
sense, because Illinois is so highly modified,
we are faced with protecting almost all native
species. Unfortunately, it is too late to protect
complete watersheds and other large areas (the
exception being Heron Pond-—Little Black
Slough Preserve in southern Illinois), and thus
we need to concentrate on identifying and
protecting streams with high species diversity
and those with rare species. Other parameters
that might be used to select streams to protect,
for example, water quality, land use, unusual
habitats, naturalness of the ecosystem, and
natural divisions, are reflected in the biodiver-
sity. If many species or rare species are present,
it is because the water quality has remained
good for a long time, because unusual habitats
are present, and so on.

Symposium Proceedings: Our Living Heritage 44]

How do we recognize streams with high
diversity and rare diversity? The best way is to
obtain data from large geographic, in this
instance statewide, data bases and compare
various localities with one another. Fortunately,
Illinois has more complete statewide data bases
on the diversity of aquatic organisms than any
other state. Burr (pages 417—427, this volume)
has discussed the surveys of fishes (Forbes and
Richardson [1908]; Smith 1979), and Cum-
mings (pages 428—438, this volume) has dis-
cussed past (Parmalee 1967; Starrett 1971) and
ongoing surveys of the mussels of Illinois. A
third important data base is that on crustaceans,
part of which was published (crayfishes and
shrimps) by Page (1985). Combined, these data
bases can be used to identify outstanding
streams by locating those that have the highest
diversity (most species) of fishes, crayfishes,
and mussels, and those that have the rarest
diversity (i.e., those that support populations of
threatened and endangered species).

Outstanding streams can also be identi-
fied by using the Biological Stream Characteri-
zation (BSC), a stream-quality classification
developed by the Illinois Department of
Conservation and the Illinois Environmental
Protection Agency (Hite and Bertrand 1989).
The classification is based on fish community
characteristics and the potential of a stream to
function as a fishery resource. Stream segments
are categorized from “A” (highest quality) to
“E” (lowest). Currently, 24 stream segments are
considered to belong in the “A” category and
about 184 in the “B” category.

This year, the Center for Biodiversity at
the Illinois Natural History Survey initiated a
study to enlarge and enhance the BSC with
statewide data on biodiversity. Fieldwork will
update existing statewide data bases, specifi-
cally those on endangered and threatened
species and on the diversity of mussel species.
These data, in turn, will be used to identify
outstanding streams in addition to those already
recognized by the BSC. The end product will
be a list of streams to be protected and man-
aged for their outstanding biological character-
istics. Although data continue to be gathered,
20 aquatic ecosystems, including 13 streams,
were identified as outstanding by Page, Burr,
and Cummings (1989) (Table 1), and they seem
certain to appear on subsequent lists of streams
in Illinois most deserving of protection.

442 Illinois Natural History Survey Bulletin

WHAT ARE THE MAJOR CAUSES OF
STREAM DEGRADATION?

The recognition of streams worthy of protec-
tion is a major accomplishment, but ultimately
it becomes a meaningless exercise unless we
identify the sources of degradation and initiate
actions to eliminate them. Smith (1971)
identified factors primarily responsible for the
disappearance of some and the decline of other
species of fishes in Illinois (Table 2). These
factors negatively affect other aquatic species
as well and are probably the principal! threats to
stream biodiversity.

Because of the pervasive nature of
agriculture in Illinois, siltation is undoubtedly
the major cause of stream degradation and has
affected at one time or another nearly every
stream in the state. Silt negatively affects
stream organisms in several ways and benefits
only a few species that are able to tolerate the
silt-laden habitats left behind when other
species die out. Silt inhibits the ability of
organisms to breathe by covering their gills and
preventing effective oxygen exchange. High
turbidity (silt suspended in water) for pro-
longed periods results in the suffocation of
many aquatic organisms—plants as well as
animals. When the primary producers (plants)
and primary consumers (e.g., many insect
larvae) are eliminated, fishes and other organ-
isms dependent on them for food die or perhaps
produce fewer offspring, and eventually species
disappear. Silt is unsuitable as a spawning
substrate for most fishes because eggs laid in

Table 1. Outstanding streams of Illinois based on
aquatic biodiversity.

. Kankakee River, Kankakee and Will counties

. Big Creek, Hardin County

. Embarras River, Jasper, Cumberland, and Coles
counties

. North Fork Vermilion River, Vermilion County

. Little Vermilion River, Vermilion County

. Crane Creek, Mason County

. Lusk Creek, Pope County

. Kishwaukee River, Winnebago, Boone, and
McHenry counties

10, Little Wabash River, Clay, Effingham, and

Shelby counties

11. Mississippi River, Rock Island County

12. Wabash River, White County

13. Clear Creek, Union County

hWnd —

wocaonrtawm

Source: Page, Burr, and Cummings 1989.

ee
. Middle Fork Vermilion River, Vermilion County

Vol. 34 An. 4

silt are unable to obtain an adequate oxygen
supply. Instead, fishes commonly lay their eggs
on gravel or among plants, where they are
hidden from predators and at the same time
remain in actively flowing water and thus in a
continuous supply of oxygen. In heavily silted
streams where gravel and plants are covered
with silt, reproductive success is reduced for
many species, and they disappear after a few
seasons. Mussels are especially vulnerable
because of their sessile habits and, as noted by
Cummings (pages 428—438, this volume), the
loss of mussel diversity in Illinois has been ex-
traordinarily large (21% of the species have
been extirpated and another 35% are in danger
of extirpation.)

“Drainage” as a factor contributing to the
loss of fishes (Smith 1971) refers to the
drainage of bottomland lakes that serve many
fishes as nurseries and some stream-dwelling
fishes as overwintering refuges and spawning
areas. In their natural condition, these lakes are
extraordinarily productive (Dodge 1989) and
favored areas for the growth and development
of small fishes. In Illinois, most of these lakes
were found along large rivers such as the
Mississippi and Illinois. Their loss resulted
from drainage to produce more farmland and
from filling with silt as sediment-laden rivers
overflowed during periods of flooding. If we
are to protect stream organisms, the remaining
bottomland lakes must be protected and, where
possible, others should be restored.

As more water is consumed in Illinois,
primarily for agricultural purposes, water tables

Table 2. Factors primarily responsible for the extir-
pation of 8 and decimation of 60 native species of
Illinois fishes.

Number of Number of

species species

extirpated decimated
Siltation 2 I4
Drainage 0 13
Desiccation during drought 0 12
Species interaction 2 7
Pollution 2 5
Dams and impoundments 0 a
Temperature 0 1
Unknown causes 2 -

88

Source: Smith 1971.

April 1991

are lowered in many places and stream desicca-
tion has become a major problem. Springs that
were formerly perennial are now ephemeral,
and species restricted to them die during
periods of drought. The disappearance of the
southern redbelly dace, Phoxinus erythrogaster,
from southern Illinois is thought to be a result
of the lower water table and the increased
frequency with which springs dry.

Detrimental interactions between exotic
and native species include competition,
predation, disease, and parasitism. Although
some species introduced into Illinois have
produced results perceived as beneficial (e.g.,
certain crops adopted from Europe), the vast
majority have proved detrimental to native
species. Familiar aquatic examples include the
common carp (Cyprinus carpio), which is
notorious for its ability to stir up stream
substrates and destroy otherwise suitable
feeding or spawning grounds for other fishes,
and the rusty crayfish (Orconectes rusticus),
which displaces native crayfishes in amazingly
short periods of time by means that are not
entirely understood. The most recent invader,
the zebra mussel (Dreissena polymorpha), is
now in the Great Lakes and likely to negatively
affect native mussels. It is already causing
major problems in water treatment and power
plants (Cummings 1990).

Much has been written about stream
pollution (e.g., Hynes 1960), and it is unneces-
sary to detail that discussion here. Briefly,
pollutants poison aquatic organisms. Major
progress has been made recently in reducing

Symposium Proceedings: Our Living Heritage

44:

point sources of pollution (Illinois Environ-
mental Protection Agency 1990), but such
nonpoint sources as the agricultural runoff of
pesticides remain a major problem.

Dams and impoundments convert large
segments of flowing water into standing water.
A few species are favored by the conversion,
but many more are eliminated. The pre-
impoundment list of species present in a
medium to large river in Illinois commonly
includes 30—40 species of fishes and 10-20
species of mussels. In contrast, an impound-
ment typically supports only 8—12 species of
fishes and 4—6 species of mussels. The nega-
tive impact of an impoundment on biodiversity
is compounded by the fact that species in the
impoundment are always common, for ex-
ample, largemouth bass (Micropterus salmoi-
des), gizzard shad (Dorosoma cepedianum),
and common carp (Cyprinus carpio); the
species lost, however, can include threatened
and endangered species. The battle in Illinois
over a proposed reservoir on the Middle Fork
of the Vermilion River (Figure 3) was in part
related to protection of the state-endangered
bluebreast darter (Etheostoma camurum), and
the battle in Tennessee over the proposed
Tellico Dam was in part related to the per-
ceived threat to the federally endangered snail
darter (Percina tanasi). Exacerbating the
negative impact of impoundments on biodiver-
sity is their tendency to fill with sediments
carried by the streams flowing into them.
Because they fill in, they are short-lived
relative to the potential life of a stream.

Figure 3. Middle Fork of the Vermilion River, Vermilion County, Illinois. Photo by Lawrence Page.

Dams negatively affect stream communi-
ties in addition to the direct effects of inunda-
tion. Many species of fishes migrate upstream
to spawn; when a dam blocks their passage,
they cannot reach suitable spawning areas. In a
relatively short time, populations decline and
sometimes disappear. A dam impedes and often
stops the flow of water downstream and causes
major alterations in the stream ecosystem.

In many streams, temperature elevation
results in the removal of riparian vegetation
that once shaded flowing water. With direct
sunlight for prolonged periods, the water is
warmed and becomes unsuitable for many
species. Another cause of warming is the
continuous lowering of the water table, with the
result that less groundwater reaches surface
streams. Fishes that generally prefer cool water
and species adversely affected by this warming
trend include trouts, nearly absent from Illinois,
and sculpins, which are becoming less common
and more restricted in distribution.

Channelization (or canalization) of
streams converts them from a series of riffles
and pools of varying characteristics into a ditch
of nearly uniform width, depth, velocity, and
substrate. Instead of providing the variety of
habitats available in an unchannelized stream,
a channelized stream offers only one habitat
and only those species capable of living in that
habitat persist. In addition, bankside vegetation
is usually removed to enable the large equip-
ment needed for channelization to gain access
to the stream. Loss of vegetation further
reduces biodiversity. The diversity of species in
a ditch is usually much lower than that in a
meandering stream.

HOW DO WE PROTECT STREAMS?

Given the major causes of degradation (Table
2) and the multiple uses of streams in Illinois, a
multifaceted approach to their protection is
imperative. Our goal is to keep the native biota
intact, and all approaches aimed at stream
protection must have as their objective to keep
the stream ecosystem as natural as possible.
Broadly considered, protection means that we
must prevent the harmful development of the
stream and the watershed and the deterioration
of the water quality.

A third alternative, restoration (e.g.,
eliminating the source of a pollutant or allow-
ing a channelized stream to return to a mean-

Illinois Natural History Survey Bulletin

Vol. 34 Art. 4

dering stream) is a reasonable and highly
desirable objective and is usually relatively
inexpensive. Such massive projects as the
restoration of wetlands, although desirable, can
be extremely expensive and inevitably fall
short of the goal of ecosystem restoration
because of the intervening loss of many species
previously present. Although a great deal of
interest and enthusiasm is being devoted to
restoration, if we must choose between
protecting the remaining “natural” ecosystems
(i.e., those least disturbed by man) and restor-
ing areas, the wiser course is to protect what we
have left rather than to devote limited resources
to restoring abused ecosystems.

Preventing development. Following the
enhancement of the Biological Stream Charac-
terization and the more complete listing of
outstanding Illinois streams, I anticipate that
the Illinois Nature Preserves Commission, The
Nature Conservancy, and other conservation
organizations will purchase easements, dedicate
preserves, or otherwise move to protect these
outstanding aquatic ecosystems. Designation of
streams and key portions of watersheds
(particularly headwaters) as nature preserves by
the Illinois Nature Preserves Commission, the
procurement of land by The Nature Conser-
vancy, and similar protective measures would
be major steps in keeping stream ecosystems
intact because the kinds of development that
negatively affect these systems would be
prevented.

In addition, when outstanding streams
appear on an official list (in this instance, the
list generated by the BSC), regulatory agencies
can require that development that might
negatively affect a stream or its watershed be
undertaken in ways that minimize these effects.
The identification of healthy and degraded
streams will result in a data base that can be
extremely useful in other studies on the patterns
and causes of stream degradation (e.g., land-use
studies).

Protecting water quality. Water quality
is protected by preventing the introduction of
contaminants such as pesticides and sewage.
One extremely important way to reduce the
most detrimental nonpoint pollutant, silt, is to
keep riparian vegetation intact. In central
Illinois, the recent practice of plowing to the
stream bank has resulted in stream bank failure
and permitted large amounts of silt to enter
streams. In addition to its value as a filter of

April 1991

silt, riparian vegetation shades the stream from
direct sunlight during the hottest part of the
year, thereby benefiting the many cool-water
species characteristic of Illinois streams.
Legislation is needed in Illinois to reduce
nonpoint pollution.

Other approaches to protecting streams
include the development of methods and
legislation to restrict introductions of exotic
species and to control the amount of water
diverted from streams for municipal, industrial,
and agricultural uses.

SUMMARY

The present characteristics and biota of the
streams of Illinois are the results of geological
and evolutionary history and the recent
modifications of streams and watersheds by
human activities. To protect Illinois streams,
we need to determine what aquatic biodiversity
remains, where it is located, and what compo-
nents need to be protected and from what. Then
we must develop the most effective means of
protection. By supplementing stream quality
ratings and statewide data bases on aquatic
organisms with fieldwork, we can identify
streams with outstanding (i.e., high and rare)
biodiversity. After outstanding streams appear
on an official state list (the BSC), regulatory
agencies can act to minimize environmental
damage.

Major threats to the integrity of Illinois
streams can be identified and protective
measures implemented even though streams,
which are affected by activities throughout
their watersheds, are clearly more difficult to
protect than are many terrestrial ecosystems.
Major threats to streams include siltation,
drainage of bottomland lakes, desiccation,
introductions of exotic species, pollution,
artificial impoundments, elevated temperatures,
and channelization. Protective measures
include the purchase of easements and the
dedication of preserves to prevent harmful
development of the stream and the watershed.
Water quality can be protected by preventing
the introduction of detrimental substances such
as silt, pesticides, and sewage. One extremely
important way to reduce siltation, the most
detrimental nonpoint pollutant of streams in
Illinois, is to leave riparian vegetation intact.
Legislation is needed in Illinois to reduce
nonpoint pollution, to restrict introductions of

Symposium Proceedings: Our Living Heritage 445

exotic species, and to control the amount of
water diverted from streams for municipal,
industrial, and agricultural uses.

LITERATURE CITED

Burr, B.M., AND L.M. PAGE. 1986. Zoogeography
of fishes of the lower Ohio—upper Mississipp1 basin.
Pages 287-324 in C.H. Hocutt and E.O. Wiley, eds.
The zoogeography of North American freshwater
fishes. John Wiley & Sons, New York.

Cummins, K.S. 1990. The zebra mussel: exotic
invader. Illinois Natural History Survey Reports 298.

n.p.

DopceE, D.P., ED. 1989. Proceedings of the Interna-
tional Large River Symposium (LARS). Canadian
Special Publication of Fisheries and Aquatic
Sciences 106. 629 p.

FENNEMAN, N.M. 1938. Physiography of eastern
United States. McGraw-Hill, New York. 714 p.

Fores, S.A., AND R.E. RICHARDSON. [1908]. The
fishes of Illinois. Illinois State Laboratory of Natural
History, Danville. cxxxvi + 357 p.

Hite, R.L., AND B.A. BERTRAND. 1989. Biological
stream characterization (BSC): a biological assess-
ment of the Illinois biological stream characteriza-
tion work group. Special Report 13 of the Illinois
State Water Plan Task Force. Illinois Environmental
Protection Agency, Division of Water Pollution
Control. IEPA/WPC/89-275. Springfield, IL.

Hynes, H.B.N. 1960. The biology of polluted
waters. Liverpool University Press, England. 202 p.

ILLINOIS ENVIRONMENTAL PROTECTION AGENCY.
1990. Illinois Water Quality Report 1988-1989.
Illinois Environmental Protection Agency, Division
of Water Pollution Control Planning Section. IEPA/
WPC/90-160. Springfield, IL.

PaGe, L.M. 1985. The crayfishes and shrimps
(Decapoda) of Illinois. Illinois Natural History
Survey Bulletin 33(4):335—448.

PaGe, L.M., B.M. Burr, AND K.S. CUMMINGS. 1989.
Outstanding aquatic ecosystems within Illinois based
on uniqueness of their fauna and environmental
quality. Pages 18-20 in M.A. Phillippi and B.D.
Anderson, eds. Preserving the aquatic biodiversity of
Illinois: inventory, research, regulation, and
protection. Proceedings of the Illinois Nature
Preserves Commission 25th Anniversary Sympo-
sium, Springfield.

PARMALEE, P.W. 1967. The fresh-water mussels of
Illinois. Illinois State Museum Popular Science
Series VIII, Springfield. 108 p.

446 Illinois Natural History Survey Bulletin

Puituipet, M.A., AND B.D. ANDERSON, EDS. 1989.
Preserving the aquatic biodiversity of Illinois:
inventory, research, regulation, and protection.
Proceedings of the Illinois Nature Preserves
Commission 25th Anniversary Symposium,
Springfield. 32 p.

Smitu, P.W. 1971. Illinois streams: a classification
based on their fishes and an analysis of factors
responsible for disappearance of native species.
Illinois Natural History Survey Biological Notes 76.
14 p.

Smit, P.W. 1979. The fishes of Illinois. University
of Illinois Press, Urbana. 314 p.

Srarrett, W.C. 1971. A survey of the mussels
(Unionacea) of the Ilinois River: a polluted stream.
Illinois Natural History Survey Bulletin 30(5):
267-403.

Vol. 34 Art. 4

Illinois Caves: A Unique Resource

James E. Gardner, Illinois Natural History Survey

Unlike neighboring Missouri with over 5,000
caves, Illinois is not known as a cave state. The
many glacial advances that extended far south
into the state buried the limestone bedrock that
is so conducive to the formation of caves.
Nevertheless, according to Oliver and Graham
(1988), at least 480 caves are found in Illinois.
They noted that the largest and most hydrologi-
cally active caves occur in the Sinkhole Plain
area of St. Clair and Monroe counties, one of
the four major cave areas in Illinois (Figure 1).
They also observed that biological activity
appears greatest in caves in the Shawnee Hills
Section.

The study of caves (speleology) encom-
passes a unique and intriguing world of
darkness, one that often extends far below the
earth’s surface. Because caves are devoid of
sunlight and green plants, they may appear
foreboding to any form of life. This perception,
combined with the difficult and oftentimes
hazardous obstacles for intrepid scientists to
overcome, would appear to make speleology an
unattractive field of study. To the contrary,
speleology is an exciting and rewarding pursuit.

The study of cave life (biospeleology) has
not been avoided simply because of potential
hazards to investigators. Scientific studies of
caves began as early as the 17th century in
Europe, when theories on cave hydrology were
introduced. Early biospeleology was limited
primarily to very general faunal surveys and to
descriptions of unpigmented animals (initially
thought to be albino) with degenerative eye
structures. In the United States, the first cave
studies were spearheaded by Europeans.
Constantine Rafinesque studied and named
cave animals in Mammoth Cave and other
caves near Lexington,,Kentucky, during his
visits around 1822. However, it wasn’t until the
late 1800s that interest in North American
caves and cave life were made fully manifest.

The history of biospeleology in Illinois
reaches back over a century when the founder
of the Illinois Natural History Survey, Stephen
A. Forbes, wrote on blind cave fishes and their
allies (Forbes 1881, 1882). The studies that
followed much later (Layne and Thompson
1952; Gunning and Lewis 1955; Weise 1957;
Smith and Welch 1978) were indirectly
associated with caves and springs and focused
on the spring cavefish (Chologaster agassizi).
By 1950, the mass of data that had been
collected by nonprofessional biospeleologists,
cave explorers, and surveyors encouraged more
complete systematic descriptions of taxonomic
groups of cave animals and their distribution.
Encouraged by a rapidly growing interest in
cave ecology and the physiology of caverni-
coles (animals found in caves), researchers
grew more interested. Peck and Lewis (1977)
provided the first and presently only compre-
hensive information on the occurrence of more
than 200 invertebrate species collected from
caves in Illinois. The only other studies of
invertebrate cave fauna in Illinois focused on
taxonomic descriptions (Yeatman 1964; Liang
1970; Steeves and Seidenberg 1971; Lewis and
Bowman 1981). Other Illinois studies involving
caves (or abandoned mines) did not consider
the larger subterranean ecosystem or its
inhabitants, but focused on bats that used caves
as roosts.

We gained a better understanding of cave
ecosystems through studies by Poulson and
White (1969), Barr (1968), Caumartin (1963),
and Poulson (1972). Perhaps the most compre-
hensive publication concerning natural cave
resources was The Life of the Cave by Mohr
and Poulson (1966). Biospeleology has now
become a recognized field of study. Universi-
ties offer degrees with emphasis on aspects of
biospeleology, and a number of nonprofit state
and national cave research and conservation

447

448 Illinois Natural History Survey Bulletin

organizations actively promote the study and
conservation of cave resources. State and
federal land management agencies have
undertaken studies involving cave resources
and the unique life forms associated with
them(Gardner 1984, 1986; Oliver and Graham
1988; J.D. Garner, pers. comm.).

There is a great need for more informa-
tion concerning Illinois caves and their associ-
ated fauna. It is my intent to introduce the
reader to the rich heritage of our unique cave
resources in the hope of fostering appreciation
and stimulating continued work. Lipman
(1965) commented that “speleology has a
definite place on the national conservation
scene,” and I share his hope that “as the need
for more detailed information about under-
ground conditions increases, the science of
speleology will grow.”

Driftless Area

Lincoln
Hills
Section

Figure |. The four major cave areas of Illinois.
Adapted from Oliver and Graham 1988.

Vol. 34 Ant. 4

THE VALUE OF CAVE RESOURCES

Our unique cave habitats and the diversity of
life they support are subjected to environmental
pressures that threaten their very existence. The
delicate balance of many cave ecosystems has
been needlessly destroyed by human activities.
Caves, springs, and other subterranean features
are a valuable part of our natural resources; yet
pollution of our subterranean water systems is
becoming increasingly evident, damaging the
resource and in the process threatening our
health and well-being. Cave explorers (spelun-
kers) must learn to be even more conscientious
in order to lessen the impact of their visits.
There must be caves left in Illinois free from
detrimental impacts, thereby conserving their
natural state for future studies.

Caves, like other more traditional natural
resources, have four basic values:

Intrinsic. In the most literal sense, caves
are a viable and important link in the great
environmental chain that binds our planet
together. Caves and the resources they contain
have an inherent value.

Aesthetic and cultural. Caves provided
dwellings for humankind long before recorded
history. Often they were sacred places associ-
ated with rites and ceremonies. Caves are
important historically and aesthetically. Their
mystery exists even today and the beauty of
untouched cave formations (speleothems)
cannot be denied. Caves are a valuable part of
our heritage.

Recreational. Spelunking is an increas-
ingly popular recreational sport. As cave
locations become known, explorers flock to see
them. Anyone who has met the challenge of
exploring passageways rarely or never seen
will have a memorable and deeply moving
experience. Caves have a recreational value but
they are also an economic asset, as documented
by the millions of tourists who buy tickets
yearly for commercial cave tours.

Scientific. Perhaps the most precious
value of our caves is found in the knowledge
we gain from studying them. Caves, like pages
in a history book, provide information on past
climate, paleontology, and archaeology.

Caves have perhaps been studied longest
by geologists, fascinated by the natural
processes of caves (Bretz 1938; Harris and
Allen 1952). Hydrologists and engineers have

April 1991

recognized the need to study caves and the
secrets of their formation (speleogenesis).
Caves provide a barometer whereby we can
measure environmental quality. Dye tracing
studies, with their subsequent determination of
water courses, have averted serious pollution
catastrophes and ensured water quality to many
communities. Finally, cave environments and
the animals associated with them provide living
systems to study. Many cave animals have
provided solutions to environmental and
medical problems; others serve as examples for
the study of basic ecological principles.

BIOSPELEOLOGICAL OBJECTIVES

Howarth (1981) argued that if cave inverte-
brates were to be targeted for conservation, top
priority should be given to conducting thorough
biological inventories and ecological studies in
threatened caves. He further emphasized that
the long-term goal in the conservation of cave
invertebrates must be the protection of suitable
cave habitats. Poulson (1975) addressed cave
management problems and their solutions,
noting the importance of baseline biological
data. Poulson and Kane (1976) provided an
excellent outline for the biological inventory of
caves, pointing out that most detrimental
impacts could be understood only if a baseline
inventory had been conducted before distur-
bances occurred. The prime objectives of a
biological resource inventory according to
Poulson and Kane (1976) are summarized
below.

Identifying species. As many species of
cave animals as possible should be identified
and recorded from each cave under investiga-
tion. This task is achieved by three methods.

1. A review of the literature. Investigators
must be familiar with the work that has
preceded their own if they are to conduct
inventories efficiently.

2. Identifications in the field. Recorded
observations provide a substantial amount of
data with minimum impact to the cave envi-
ronment. Cave invertebrates are among the
most difficult life forms to discover, observe,
and identify. Specific determinations of
invertebrate cave fauna often require a
taxonomic specialist. Identifications of
vertebrates do not usually require detailed
knowledge of microscopic taxonomic charac-
teristics. Bats, salamanders, and certain fishes

Symposium Proceedings: Our Living Heritage 449

can be readily identified by trained observers.
Collecting and preserving cave vertebrates for
the sole purpose of identification is an
unacceptable method of inventory. Several
species of cave-dwelling vertebrates are
protected by state and federal legislation that
prohibits their collection.

3. Identification through established collec-
tions. Identifications of most cave inverte-
brates are usually made through reference to
existing collections. Specific determinations
of fauna are often based on microscopic
morphological characteristics (1.e., legs,
antennae, mouth parts, reproductive organs).
Such identifications are usually well beyond
the capabilities of most investigators, and
taxonomic specialists need to be consulted.

Documenting cave fauna. Unfor-
unately, budget constraints significantly limit
the extent to which cave resources can be
studied. As a result, threatened or endangered
species usually receive priorities for study. This
limitation should not, however, restrict the
gathering of information to only those species.
If an ecosystem or habitat approach is fol-
lowed, all faunal elements in a cave protected
for listed species can be studied.

Noting species associations and
ecologically related information. The identifi-
cation of individual elements of a cave’s fauna
provides insight into the entire ecosystem.
Often the occurrence of a particular species can
be anticipated by the presence of another
species.

Identifying future study areas. Inven-
tories of biological resources are important in
identifying caves where more detailed studies
are needed. Priorities can then be set since a
detailed study of each cave is impossible in
terms of time, labor, and money.

Developing recommendations. Cave
resources are an integral part of our natural re-
sources, but responsible management or
enhancement of any resource cannot be
accomplished without first identifying its
elements. Cave resources require very special
management.

THE CAVE ENVIRONMENT

The cave environment affects the behavior,
development, and evolution of the organisms
living there. The absence of light, near-constant
temperatures, and the amount of humidity all

450 Illinois Natural History Survey Bulletin

influence the animals found in caves and their
positions within the cave relative to the
entrance. Cave climates vary little compared to
surface climates. The cave environment is cool
and humidity is usually high; evaporation rates,
therefore, are very low. Air currents in caves
(cave breathing) are normal events in response
to surface barometric pressure and can mark-
edly affect temperature and humidity within a
cave.

Caves can be divided into zones based on
the amount of light and the degree of changes
in temperature and humidity.

Twilight zone (cave entrance). The
twilight zone extends into the cave as far as
unaided human vision is possible. This zone is
usually damp and cool, but temperature and
humidity fluctuations are close to those found
outside the cave. Some green plants may
invade the entrance area, and this zone contains
the largest and most diverse fauna in the cave.
Animals found in the twilight zone include
surface species of birds, mammals, snakes,
frogs, and many different species of inverte-
brates that are commonly associated with the
surface.

Middle zone. This zone lies just beyond
the twilight zone and is characterized by total
darkness. Temperature and humidity vary
somewhat with seasonal changes at the surface.
Animals found in this zone include bats,
crickets, millipeds, and surface species of
amphipods and isopods.

Zone of total darkness and nearly
constant temperature. This zone, like the
middle zone, is devoid of light; however,
temperatures fluctuate only slightly from the
average annual mean temperature of the
ground, approximately 13 to 15°C (54 to 58°F)
in Illinois. The humidity remains nearly
constant, usually near 100%. Animals inhabit-
ing this zone are usually obligative cave-
dwelling species such as blind, unpigmented
amphipods, isopods, cave fishes, pseudo-
scorpions, and springtails.

THE CAVE ECOSYSTEM

A cave ecosystem can be defined as all of the
living organisms within a given cave bound
together by interrelationships and interacting
with the physical environment of the cave.
Cave animals can be classified by their

Vol. 34 Art. 4

interaction with the cave environment or by the
role they play in the cave ecosystem—their
ecological classification (Barr 1963). Some
organisms possess highly specialized adapta-
tions that allow them to live in a world of total
darkness, extremely low food availability, and
relatively constant temperature. The organisms
that inhabit caves are divided into two catego-
ries: epigean or surface-dwelling organisms and
hypogean or subsurface organisms.

Epigean (surface) organisms. These
animals usually must complete their entire life
cycle on the surface. When found in a cave
environment, they are classified as accidentals.
Epigeans that wander, fall, or get washed into a
cave will either escape or eventually perish
there.

Hypogean (subsurface) organisms.
These animals normally live below the surface
in caves, in subterranean water courses, or in
interstitial environments (i.e., between soil
particles). The three commonly recognized
classes of hypogeans are troglobites, troglo-
philes, and trogloxenes. The ecological term
endogean, or edaphobite, is used to classify
species that normally live in soil (e.g., earth-
worms). Additionally, phreatobite is a term
used to describe animals that inhabit the upper
layers of groundwater (Holsinger 1969); it is
considered synonymous with troglobite.
Troglobitic species account for only 20 to 30%
of the faunal assemblages of most North
American caves. The largest percentages of
cave fauna are troglophiles and trogloxenes.

Troglobites, as the derivation of their
name suggests (from the Greek for hole and to
live), live exclusively in caves, springs, or
subterranean water systems; they cannot
survive outside these environments. Troglobites
are perhaps the most fascinating of all cave
species because they possess marked morpho-
logical adaptations to subterranean environ-
ments. Illinois contains a diversity of troglo-
bitic invertebrates. Peck and Lewis (1977)
reported 18 troglobitic invertebrates from
Illinois, 14 of which are considered endemic
(found nowhere else on earth). However, no
populations of troglobitic vertebrates (i.e., true
cavefishes and salamanders) are known from
the state.

Troglobites possess morphological,
physiological, and behavioral adaptations that
make them unique. Compared to their surface

April 1991

relatives, troglobites have reduced metabolic
rates. Their sensory capabilities are modified,
including reduced or absent vision, increased
vibration (hearing) reception, increased
olfaction (smell or chemo-reception), and
increased tactile sensitivity. Their appendages
are longer and more slender, and their move-
ments are slower, more deliberate. Their bodies
also tend to be more slender. Reproduction
periods are acutely tuned to the seasonal
availability of food, and fewer and larger eggs
are generally laid.

Troglophiles (cave loving) commonly
inhabit caves and can complete their entire life
cycle there; however, they are also found in
cavelike microclimates on the surface (i.e.,
deep down in surface leaf debris, in crawl
spaces beneath buildings, or inside wet, rotting
logs). Examples of troglophiles in Illinois are
the cave salamander (Eurycea lucifuga) and
species of isopods and beetles.

Trogloxenes (cave visitors) frequent
caves for shelter and favorable microclimates
but must return to the surface to complete some
portion of their life cycle (i.e., feeding and re-
production). Bats are classified as trogloxenes
as are raccoons, birds that nest in the entrance
of caves, and certain species of snakes.

THE NEED FOR CONSERVATION

Bretz and Harris (1961) published descriptions
and locations of more than 60 caves throughout
Illinois. Their section on basic cave formation
(speleogenesis) and cave types is complete and
educational. Unfortunately, the publication of
the exact locations of these caves opened the
way for vandalizing the larger, more popular
ones. Enticed by descriptions of passageways
and the beautiful formations they contained,
novice, adventure-seeking explorers trampled
through the caves, defacing and destroying
some of the finest cave resources of Illinois.
Relatively few caves have been protected,
and many are in dire need of protection. In
response to this need, the Illinois legislature
passed the Cave Protection Act in 1985. Drafted
by the Illinois Department of Conservation
(J. D. Garner, pers. comm.), the act established
measures for the protection of the natural and
cultural resources of Illinois caves. An inven-
tory of the natural resources of over 80 Illinois
caves was conducted by the Illinois State

Symposium Proceedings: Our Living Heritage 451

Museum (Oliver and Graham 1988). Addition-
ally, the Illinois Department of Conservation
and the Illinois Natural History Survey conduct
investigations of biological cave resources;
emphasis is given to endangered bats.

Recent protection measures for Illinois
caves were perhaps precipitated by the recogni-
tion (White 1973) and classification (White
1978) of these resources during the Illinois
Natural Areas Inventory. As a result of that
study and the ongoing efforts of the Illinois
Department of Conservation, several caves
have been identified as having significant
natural resource features and are included in the
Illinois Natural Areas Inventory. Other caves
have been designated as Illinois Natural
Heritage Landmarks in order to protect their
valuable resources. One Illinois cave, with at
least 12 miles of passageway, was purchased in
1987 and dedicated on August 31, 1989, as an
Illinois Nature Preserve to protect a hibernating
population of the federally endangered Indiana
bat (Myotis sodalis). Another cave, Illinois
Caverns, was purchased in 1986 and classified
as an Illinois Natural Area. Six miles of
passageway in Illinois Caverns are open to the
public for exploration through a permit system
designed to protect the cave.

More studies are needed to identify and
understand the unique biological resources of
Illinois caves. The delicate and intricate natural
communities of our caves cannot be protected
unless we identify their elements. However,
biological collections in caves should never be
done without first consulting competent
authorities. Over-collecting and improper
collecting methods have been extremely
harmful to some populations of cave species.
The admonition, “Take only pictures and leave
only footprints” should have special signifi-
cance to every Illinoisan if we are to ensure that
our unique cave resource is secured for future
generations.

LITERATURE CITED

Barr, T.C., JR. 1963. Ecological classification of
cavernicoles. Cave Notes 5(2):9-12.

Barr, T.C., JR. 1968. Cave ecology and the
evolution of troglobites. Pages 35—102 in T.
Dobzhansky, M.K. Hecht, and W.C. Steere, eds.
Evolutionary Biology Vol. 2. Appleton-Century
Crofts, New York.

452

Bretz, J.H. 1938. Caves in the Galena Formation.
Journai of Geology 46:825—841.

Bretz, J.H., AND S.E. Harris, Jr. 1961. Caves of
Illinois. Illinois State Geological Survey Report of
Investigations 215. 87 p.

CaumarrIN, V. 1963. Review of the microbiology of
underground environments. Bulletin of the National
Speleological Society 25:1—14.

Forses, S.A. 1881. A rare fish in Illinois. American
Naturalist 15(3):232—233.

Forses, S.A. 1882. The blind cave fishes and their
allies. American Naturalist 16(1):1—S.

GarpneR, J.E. 1984. Missouri Department of
Conservation cooperative cave inventory project.
Final report, Missouri Department of Conservation,
Jefferson City. 125 p.

GARDNER, J.E. 1986. Invertebrate fauna from
Missouri caves and springs. Natural History Series 3,
Missouri Department of Conservation, Jefferson
City. 72 p.

GUNNING, G.E., AND W.M. Lewis. 1955. The fish
population of a spring-fed swamp in the Mississippi
bottoms of southern Illinois. Ecology 36(4):
552-558.

Harris, S.K., JR., AND B.D. ALLEN. 1952. Caves of
the Kincaid Limestone near Cobden, Illinois. Trans-
actions of the Illinois State Academy of Science
45:196-207.

HOoLsinceR, J.R. 1969. Biogeography of the fresh-
water amphipod crustaceans of the central and
southern Appalachians. Pages 19—50 in P. C. Holt,
ed. The distributional history of the biota:of the
southern Appalachians. Part I. The invertebrates.
Virginia Polytechnic Institute, Resource Division
Monograph I.

Howarth, F.G. 1981. The conservation of cave
invertebrates. Pages 57—64 in J.E. Mylroie, ed. First
International Cave Management Symposium, Pro-
ceedings. Murray State University Press, Murray, KY.

Layne, J.N., AND D.H. THompson. 1952. Recent
collections of the amblyopsid fish Chologaster
papillifera in Illinois. Copeia 3:39—40.

Lewis, J.J.. AND T.E. BowMAN. 1981. The subterra-
nean asellids (Caecidotea) of Illinois. (Crustacea:
Isopoda: Asellidae). Smithsonian Contributions to
Zoology 335. 66 p.

LIANG, C. 1970. The soil microfungi of Burton Cave,
Adams County, Illinois. M.S. thesis, Western Illinois
University, Macomb. 45 p.

Illinois Natural History Survey Bulletin

Vol. 34 An. 4

LipMAN, R.P. 1965. Speleologists role in conserva-
tion. Journal of Soil and Water Conservation
20:197-198.

Morr, C.E., AND T.L. PouLson. 1966. The life of
the cave. McGraw-Hill, New York. 232 p.

O.iver, J.S., AND R.W. GRAHAM. 1988. Preliminary
inventory of natural resources in select caves in
Illinois. Final report, Illinois Department of Energy
and Natural Resources, Illinois State Museum,
Springfield. 155 p.

Peck, S.B., AND J.J. Lewis. 1977. Zoogeography and
evolution of the subterranean invertebrate faunas of
Illinois and southeastern Missouri. National
Speleological Society Bulletin 40:39-63.

PouLson, T.L. 1972. Bat guano ecosystems.
National Speleological Society Bulletin 34:55—60.

PouLson, T.L. 1975. Management of biological
resources in caves. Pages 46—52 in National Cave
Management Symposium Proceedings. Speleobooks,
Albuquerque, NM.

PouLson, T.L., AND T.C. KANE. 1976. Ecological
diversity and stability: principles and management.
Pages 18-21 in T. Aley and D. Rhodes, eds.
National Cave Management Symposium Proceed-
ings. Speleobooks, Albuquerque, NM.

Poucson, T.L., AND W.B. Wuite. 1969. The cave
environment. Science 165:971—981.

Situ, P.W., AND N.M. WELCH. 1978. A summary
of the life history and distribution of the spring
cavefish, Chologaster agassizi, Putnam, with
population estimates for the species in southem
Illinois. Illinois Natural History Survey Biological
Notes 104. 8 p.

STEEVES, H.R., II], AND SEIDENBERG, A.J. 1971. A
new species of troglobitic asellid from Illinois.
American Midland Naturalist 85:23 1—234.

WEIsE, J.G. 1957. The spring cave-fish, Chologaster
papilliferus, in Illinois. Ecology 38:195—204.

Wuite, J. 1973. Preservation of caves in Illinois.
Report for the Illinois Nature Preserves Commission,
Rockford, IL. 11 p.

Wuire, J. 1978. Illinois natural areas inventory
technical report. Vol. I. Survey methods and results.
Illinois Natural Areas Inventory, Urbana, IL. 426 p.

YEATMAN, H.C. 1964. New cavernicolous cyclopoid
copepod from Tennessee and Illinois. Journal of the
Tennessee Academy Science 39:95-98.

Session Five: Agro-Urban Ecology

The time has long since passed when a citizen can function responsibly without a broad understanding of the

living landscape of which he is a part—Paul B. Sears

Agricultural and urban development practices
that take into account the conservation of the
remarkable biodiversity of Illinois must be
initiated and encouraged. A balance between
economic development and the preservation of
natural resources must soon be struck, for it
seems that “economics” continues to win and
very soon there will be little left to preserve.
With the conversion of the landscape to inten-
sive row cropping has come the realization that
perhaps our system could operate at a somewhat
less intense level. With 99.93% of the landscape
of Illinois reflecting some degree of develop-
ment, the point of no return seems imminent.

Although the production of food is of
course beneficial and necessary, the mainte-
nance and restoration of our natural heritage—
the landscapes that reflect presettlement
conditions complete with the organisms they
support—also represents a desirable and
perhaps even essential course of action. Com-
mon ground must be found between these two
opposing courses if the requirements of both are
to be met. Perhaps agro-ecology will provide
that common ground. In retrospect, we seem to
have been moving toward agro-ecology for
some time. Consider, for example, the interest
shown in organic gardening and low-input and
sustainable agriculture. Agro-ecology, however,
moves a step closer by requiring a balance
between the requirements of agriculture and the
obligation to preserve our natural heritage.
While our agricultural system presently requires
vast biological deserts populated by a single
species, the same principles need not be applied
to the surrounding landscape. Fields do not have
to be cultivated to the very edges of rivers and
streams; fencerows and windbreaks do not have
to be removed to squeeze in a few more rows of
com; railroad rights-of-way that support
corridors of native vegetation do not have to be
destroyed; streams do not have to be channel-
ized; and species of organisms need not be

driven to extinction in the name of short-term
economic development.

The next generation of agriculturalists
must farm from an ecological perspective and
the time has come when all Illinoisans, farmers
and city-dwellers alike, must adopt a conserva-
tion ethic. To quote Francis Moore Lappe,
“Individual well-being is impossible outside of
the well-being of others.” Ultimately, we can
maintain our well-being only if “others”
include all species of organisms, not merely
Homo sapiens.

Papers read at this session introduced
long-range perspectives (for example, the
movement of biota between natural and
managed ecosystems) as well as more immedi-
ate ones (for example, the management of
urban deer populations). The closing remarks,
both disturbing and challenging, concluded
this session and the symposium.

The Land Use Controversy: Maintaining and Increasing
Biotic Diversity in the Agricultural Landscape of Illinois

Michael E. Irwin, Illinois Natural History Survey

Approximately one hundred years ago our state
underwent a rapid and extensive agricultural
transformation that converted the rich, fertile
soils and relatively flat terrain underlying its
prairies and forests into vast tracts of field
crops, primarily corn and soybean. With the
exception of Iowa, a state with a history similar
to that of Illinois, the Great Plains, with its vast
expanses of wheat, and a few large tropical
countries like Brazil and Indonesia, which have
exploited their lands by putting in broad
stretches of such perennial crops as rubber and
African oil palm, there is perhaps no extensive
area on earth that is so heavily cultivated in so
few plant species as the state of Illinois. This
agricultural transformation has taken, and
continues to take, a heavy toll on native biota.
With only 11% of our land now left in natural
vegetation and over 53% of our woody plant
taxa found in cultivated areas, we have cause
for concern. Can this trend be reversed? If so,
at what price?

Two viewpoints seem in genuine conflict.
On the one hand, we have the argument that
agricultural production must be sustained to
meet our food needs and to offset the nation’s
balance of payment deficits through expanded
exports. Those holding this short-term view
make a powerful case that meeting these needs
benefits humanity and our citizens in nutri-
tional and economic ways. On the other hand,
the persistent exploitation of our natural areas
continues to deplete the biotic richness of our
lands, diminishing the legacy for future
generations and restricting our access to diverse
genes for future manipulation. An equally
powerful argument, this long-term perspective
recognizes that what is exterminated can never
be restored.

I propose that these seemingly opposing
positions might be resolved in a manner that
satisfies both factions. Aspects of natural

454

systems may enhance agriculture; similarly,
aspects of managed landscapes may safeguard
natural systems and provide a formula for
recovering biotic richness in pillaged habitats.

COMPONENTS FOSTERING
SYNERGISM

The components that are responsible for
fostering potential synergism must be deter-
mined, and the interactions among those
components examined. All systems could then
be managed with a view towards optimizing
selected synergistic interactions. Three ele-
ments seem of particular importance: refugia,
biological diversity, and genetic richness.

Refugia. Parcels of land that for one
reason or another retain unique biota during
times when that biota would otherwise not be
present are referred to as refugia. How agricul-
tural oases and other biotically favorable.
artificial environments sustain species locally
through times of natural emigration or diapause
and how this ability to sustain biota affects both
natural systems and managed landscapes are of
considerable consequence.

Irrigation in agricultural settings,
particularly in semitropical areas that undergo a
season of prolonged drought, can provide
habitats favorable for the atypical overseason-
ing of some biota. As a result, these organisms
need not emigrate or aestivate. Irrigation could
alter the customary overseasoning habits of a
variety of organisms, including insects and
their natural enemies, especially in dry tropical
forest habitats. Irrigation could also alter the
time of year during which certain biota invade
natural systems from agricultural settings, with
a conceivably enormous impact on both
systems. Such invasions already occur regu-
larly in Illinois through the introduction of
plants grown in greenhouses for propagation in
orchards and home gardens.

April 1991

Just as agricultural systems provide
niches for noncrop-related organisms, so do
natural areas harbor both pests and beneficial
organisms that either plague or safeguard
agricultural crops. The role of refugia in
sustaining these complex interactions is
relatively unknown; the repercussions, how-
ever, are undeniably profound.

Biological diversity. A portion of
the diversity of life in one system will inevita-
bly invade nearby systems; how this invasion
affects a recipient system is of considerable
interest to conservationists and agriculturalists
alike. If agriculture is considered an invasive
system that receives much of its noncrop biotic
diversity in the form of colonists from sur-
rounding systems, the long-term monitoring of
colonization might help us to formulate models
of invasion rates and types of colonists through
time.

Similarly, areas where agroforestry and
agriculture are practiced could greatly influence
the biological integrity of adjoining natural
systems. Scott Robinson (page 382, this
volume) provided an example at this sympo-
sium when he talked about how habitat
fragmentation increased nest parasitism among
some of our song birds. Another example is the
introduction of the honeybee, which has
probably had a great, although unmeasured,
impact on natural pollinators in some areas.
Monitoring herbivorous insects and their
natural enemies might help us develop models
of biotic interchange—a third case in point.

Genetic richness. Any biological
species consists of a number of populations.
Each population includes a number of individu-
als, each with a slightly different genetic
makeup or genotype. The genetic richness
within a population purportedly equips that
population to withstand environmental disrup-
tion, although the process itself is not well
understood. When a population from one
system invades another, a very restricted
portion of the invading population may manage
to pass successfully from its resident system
and colonize the other. Successful invasions of
this nature are sometimes referred to as genetic
bottlenecking. The result of colonization and
the accompanying extinctions has enormous
consequences on the sustainability of a given
population, especially one in the area being

Symposium Proceedings: Our Living Heritage

invaded. The genetic richness of invading
populations might well be influenced by the
proximity and relative sizes and shapes of the
systems in question. Such concepts as habitat
fragmentation and patch dynamics are very
much a part of this process. Natural systems
harbor genetically adaptable populations of
harmful and beneficial species that continually
invade agricultural systems. Similarly, agricul-
tural landscapes probably contain genetically
adaptable populations that continually invade
natural systems. Understanding the nature of
genetic richness and how that richness affects
invasion is important in designing sustainable
agricultural and forestry systems.

BIOTIC LINKS

An inevitable exchange of biota occurs
wherever two ecosystems come into contact.
The zone of interchange, called an ecotone, is
in a sense a battleground for genetic and biotic
dominance and compatibility. When a natural
system is ravaged by deforestation or by the
introduction of agriculture, the system usually
transforms in stages—for example, from
pristine forests to high-input row-crop agricul-
ture. An ecotone is established along the
spatiotemporal border of this shift and could
well govern the rates and types of biotic
interchanges between natural and managed
systems. The role of a shifting ecotone in the
ecological and economic balance of biota in
natural and managed systems remains a
mystery and demands investigation.

Refugia, biological diversity, and genetic
richness are each affected by successful
movement of biota across ecotones. The spatial
and temporal links between natural systems and
agricultural landscapes can influence the nature
and, perhaps more importantly, the rate at
which these interactions occur. The size and
configuration of areas of land where agro-
forestry and agriculture are practiced in relation
to the size and configuration of the remaining
natural area, for instance, could be decisive in
determining how managed expanses interact
with natural systems.

The movement of biota between natural
and managed ecosystems can have dramatic
effects on both types of systems. As stewards
of this earth, we must manage the effects so

456 Illinois Natural History Survey Bulletin

that a balance is achieved between short-term
and longer-term goals. The sobering realization
is that we know so little about these inter-
changes and how they affect both types of
systems.

Our ability to sustain high-input agricul-
ture has a limited horizon. Time is running out
for earth’s rich natural ecosystems. We must
set a course that will uncover the biotic
relationships between these systems so that
they can be wisely managed in the future. I
urge a strong, timely research and education
agenda that critically addresses this issue.

Vol. 34 Art. 4

Farm Programs, Agricultural Technologies, and Upland

Wildlife Habitat

Richard E. Warner, Illinois Natural History Survey

Since the late 1800s, the grassland habitat of
upland wildlife in Illinois has been modified in
one way or another by agriculture. Although
the prairie was gone by the early 1900s, much
of the farmland in Illinois through the 1950s
contained various grasses, including small
grains, forage crops (cool-season grasses and
forage legumes), and uncultivated areas. These
farmland mosaics sustained most small
vertebrates that had once been common on the
prairie, even though pasturing and haying
caused significant mortality. After World War
II, however, farm programs and agricultural
technologies began to change, gradually
leading to greater chemical and mechanical
disturbances of farmland and the loss of
grassland as row-crop farming expanded. By
the late 1970s, even the most common upland
wildlife—ring-necked pheasant, cottontail,
bobwhite, and ground-nesting sparrows—had
registered dramatic declines. During the 1980s,
the intensive cultivation of corn and soybeans
moderated, and grassland was more widely
planted, primarily as part of annual set-aside
programs that diverted cropland from produc-
tion. The response of upland wildlife to the
reestablishment of grassland has been minimal,
presumably because farm programs require or
encourage management practices on set-aside
fields that are not conducive to the reproduction
and survival of most small animals using
grassland in Illinois. Moreover, grasslands on
farm landscapes now tend to be small, linear
patches unattractive to “interior” species. Such
highly fragmented tracts also typically sustain
high densities of opportunistic mammalian
predators. Further, the intensive chemical and
tillage disturbances on cropland have limited
the availability of insects and plant seeds, the
critical food resources of wildlife.

457

Evaluating Alternatives for Urban Deer Management

James H. Witham, Illinois Natural History Survey

Deer management in metropolitan areas is
complicated by the conflicting values of
publics with special interests. Those in charge
of developing programs that address site-
specific needs are well advised to consider
various alternatives during the planning stage.
Failure to review management options can
result in uninformed or biased decisions, which
in turn contribute to further controversy and
reduce the credibility of those in charge of the
program. Published reviews of deer manage-
ment alternatives generally point out the
limitations and advantages associated with
various control methods and include an
assessment of the usefulness of each method.
Relying on such evaluations can be helpful, but
making judgments too early, for example at the
stage when potential options are being listed,
can result in less efficient methods being
censored or eliminated prematurely. Early
elimination may be detrimental because less
efficient methods often have desirable attrib-
utes that can be combined with more efficient
management techniques. Relying on a combi-
nation of methods for the management of deer
in urban areas is appealing because it creates a
basis for compromise among diverse interest
groups.

In large metropolitan areas, such as
Chicago, where deer are abundant and adverse
interactions with people are widespread and
frequent, the state wildlife agency can facilitate
local decision making by maintaining a
computerized data base of deer management
alternatives. Three categories are useful: an
unrestricted list of deer management options,
potential strategies that rely on a combination
of options or suggest how options can be
combined, and field-tested management
programs and research that document which
methods have worked and which have failed
and why. Such an information base is one

458

product of the Urban Deer Study conducted by
the Illinois Natural History Survey, and we
anticipate that it will be used by the Illinois
Department of Conservation and the many
airports, arboretums, forest preserves, and
municipalities in the Chicago Metropolitan
Area that manage local deer populations.

Illinois Railbanking Study

Richard Pietruszka, Greenway Coordinator, Illinois Department of Conservation

The Illinois Railbanking Study was initiated by
the Illinois Department of Conservation in
1989 in response to the growing recognition
within the state and nation that abandoned
railroad corridors should be preserved for
multiple public uses. Among the objectives of
the study are the exploration and evaluation of
the natural and outdoor recreational resources
associated with the acquisition and develop-
ment of greenways and their management.

Detailed analyses of the following issues
related to the conversion of abandoned railroad
corridors into multipurpose public resources
were conducted:

The concerns of local governments and
landowners adjacent to adandoned
railroad corridors.

The identification and evaluation of
strategies that might allay local concerns
and resolve conflict.

The evaluation of the economic impact,
including the impact on local taxes, of the
conversion of abandoned railroad
corridors to multipurpose public re-
sources.

The identification of the potential users
of converted corridors.

The principle purpose of the Illinois
Railbanking Study, concluded in August of
1990, is to assist the Illinois Department of
Conservation with the formulation of policies
and planning strategies for a statewide trail
system.

459

Closing Remarks

Brian D. Anderson, Director, Illinois Nature Preserves Commission

I was very pleased to be invited to offer the
concluding remarks for this symposium. The
Illinois Natural History Survey has developed
through the years a world-renowned reputation
as a center of scientific inquiry. I’ve found the
presentations of the last two days extremely
informative, but also disturbing. It is important,
I believe, to look at the information provided
on various species groups and community types
within the context of the landscape on which
they occur. Illinois has led the nation in
developing institutions like the Natural History
Survey, the Endangered Species Protection
Board, the Nature Preserves Commission, and
the Division of Natural Heritage of the Illinois
Department of Conservation—all dedicated to
the identification and preservation of the bio-
diversity of the state. Unfortunately, the
founding of these institutions was not by
coincidence. No place in the hemisphere has
been more drastically altered by the hand of
humankind. I might also mention that the
statistics I’m about to present were also largely
compiled by the Natural History Survey. Over
80% of Illinois is currently committed to
agriculture, and another approximately 5% of
its surface acreage is urbanized. That leaves
approximately 15% of Illinois as undeveloped
land. Of that, only 0.07 of 1% retains to some
degree its presettlement condition. The full
complement of native plants and animals has
been forced to survive on less than 100,000
acres of land. The impact to our biota has been
devastating; of the approximately 2,500 species
of vascular plants considered to be native to
Illinois, 356 (about 14%) are considered to be
threatened or endangered. Our vertebrate fauna
has been even more severely affected: of 649
native vertebrates, 93 (14%) are listed as
endangered or threatened, not to mention the 30
or so species that have already been extirpated
from our state.

460

And the carnage continues, but not
through spectacular catastrophic events. We
can’t point to an Exxon Valdez or a Chernobyl.
The greatest threat to the native biodiversity of
Illinois isn’t apocalyptic; it is simply diminu-
tion, the slow but steady erosion of our
biological heritage—a road here, a 404 permit
there, individual by individual, population by
population, species by species.

I spent Earth Day in Springfield, and
sprinkled among the rally speakers was the
reading of a contest-winning essay. The topic
was “What Earth Day Means To Me.” It caused
me to reflect, and I realized I had only hopes
for the meaning of Earth Day. And foremost
among these was one. I hoped that Earth Day
1990 was the last day I had to listen to the
terms environmental trade-off and environ-
mental compromise. We have to put a word
back into our vocabulary—a little word, an
important word, the word no. Where natural
areas or habitats of endangered species are
involved, we must “just say NO!” If it’s a road,
take another one. If it’s a condo complex, put it
somewhere else. If it’s am ORV? Well, if it’s an
ORY, send it back to Japan.

I also listened that day to many speeches
heralding our achievements since Earth Day
1970, always with special mention of passage
of the Clean Air Act, the Clean Water Act, and
the Endangered Species Act. All of these were
worthy achievements. They were also all
passed in the first decade after the first Earth
Day. And the reauthorization of each was
challenged by the Federal Administration in the
second decade after the first Earth Day
Perhaps I am confused, but I thought it was
pretty obvious that on Earth Day 1990 we were
celebrating the end of a decade of environ-
mental backsliding. It is my hope that Earth
Day 1990 was the day that 100 million citizens
of the world let the leaders of the western world

April 1991

know that environmental compromise had no
place on any political agenda, conservative or
liberal. Planetary survival is, in and of itself, a
conservative concept.

A couple of years ago I sat with a
conservative acquaintance listening to a
presentation on the decline of the natural
character of our national parks. He commented
that the fellow hadn’t learned that the gloom
and doom message of radical environmentalism
had lost its credibility. The world hadn’t ended,
and no one wanted to hear that message
anymore. I guess he’d missed the news of Love
Canal, Three-mile Island, Chernobyl, Bhopal,
the donut hole in the ozone layer, and global
drought perhaps due to global warming.

Well, I just want to assure him that’s not
my message. I don’t intend to sound morose;
however, we have wasted a critical decade.
Given our technological sophistication, we
should be much farther along in solving our
environmental problems, including the bio-
diversity crisis. So don’t worry. We no longer
have time for hand wringing. I don’t intend to
depress you, I intend to press you; press you on
every front where we possess the technology to
improve the environment.

So what is the job before us? First, where
the preservation of significant extant resources
is involved, we must be uncompromising. We
can afford to lose no more. Natural areas,
habitats of endangered species, and wetlands
are just plain off limits from here on. The
developers and planners must hear this message
from scientists, conservationists, environmen-
talists, and politicians. And if the latter are
raising their voices in the wrong chorus, they
should be sent to look for new jobs.

As for specifics: We must pass legislation
to extend the consultation provisions of the
Illinois Endangered Species Protection Act to
natural areas this session. That legislation was
recently introduced as House Bill 3991.
(Postscript: it never left committee. )

Second, we must pass strong legislation
to protect our remaining wetlands. You can
help do that by supporting HB 3712 and SB
1907. (Postscript: neither was brought to the
floor of the House of Representatives for a
vote.)

Third, we can no longer tolerate the
narrow interpretation of the definition of public
waters employed by the Division of Water
Resources of the Illinois Department of

Symposium Proceedings: Our Living Heritage 461

Transportation. Governor Thompson should
force the division to accept the Attorney
General’s opinion, which would extend the
division's jurisdiction to most of our streams.
If the Division of Water Resources hasn’t
assumed that responsibility by this time next
year, we should have those jurisdictions
removed entirely from the Illinois Department
of Transportation. (Postscript: a compromise
measure was drafted but not introduced.)

Fourth, we should hold every one of our
elected representatives responsible for seeing
that the first of these three objectives is
achieved. (Postscript: none was achieved.)

Even if we were to lose nothing else, we
probably could not ensure the long-term
survival of the biodiversity of our state. We
must also restore Illinois.

The Illinois Nature Preserves System
preserves remnants of high-quality natural
communities. Most of these, however, are too
small to protect wide-ranging or area-sensitive
species. We must begin to establish biotic
reserves, which are very large preserves having
a high-quality core surrounded by degraded but
restorable lands. Using the knowledge we will
gain in establishing biotic reserves, we must
then, through restoration management, begin to
restore our open spaces to native natural
communities.

I had a dream a couple months ago. I
dreamt I was in a village where everyone, small
children to the elderly, were preparing for a
wedding. Some were scouring the countryside
for rocks and metals; more skilled hands were
shaping gemstones and cutting jewels; still
others were crafting chains of silver and gold.
Finally, the bride appeared; she wasn’t a young
woman. She was tall—a little wide in the
middle—and bore the scars of nurturing several
generations of offspring. But when she was
draped in that cloak of jewels and gems linked
by golden and silver chains, she was trans-
formed into an unparalleled beauty. I see some
of those hands in our audience; I’ve seen them
in our nature preserves; I have seen them
building conservation areas, restoring railroad
prairies, and protecting river corridors. We
must do a lot more of all of these things, but we
must also integrate our efforts.

I would, therefore, call for the establish-
ment of an Institute of Land Use Studies. The
objective of this entity would be to apply the
most current computer and satellite technology

462 Illinois Natural History Survey Bulletin

available to the identification, protection,
preservation, and restoration of our native
landscapes—and thereby our biodiversity. This
institute would also allow Illinois to lead the
nation as the center for land use planning
technology. The federal government has
abdicated its traditional role as a leader in this
area. We should, therefore, help ourselves and
at the same time develop the tools to preserve
other important centers of biodiversity, for
example, those in the tropics.

Secondly, we should begin immediately
using the Geographic Information System of
the Natural History Survey to integrate state-
wide natural resource planning efforts. State-
wide rails-to-trails conversions, watershed
planning, nature preserve and biotic reserve
establishment, river corridor preservation,
wetland protection, and prairie and savanna
restoration should all be coordinated through a
statewide protection planning committee hosted
and chaired by the Department of Energy and
Natural Resources.

Thirdly, we should press immediately for
sustained funding for natural history survey
work. For far too long the Illinois Natural
History Survey has been dependent on con-
tracts from private, profit-motivated interests in
order to monitor what is happening in Illinois
landscapes. For example, although a great
effort is underway to complete basic survey
work on the state’s streams, we are desperate
for recent faunistic surveys of habitats of high
endemism such as caves, seeps, and springs.

We must also begin to look carefully at
invertebrates, including Illinois arthropods.
You will notice I didn’t even mention the
percentage of currently listed invertebrates.
Only well-known groups of invertebrates, like
mussels and crayfish, have been addressed, and
we are not even sure of the total numbers of
species in other groups of arthropods. A beetle
found in only one cave in Illinois, one cave in
the whole world, is a treasure; one that I am not
prepared to write off.

While we were all pleased that a portion
of the real estate transfer tax was dedicated last
legislative session to the acquisition of natural
areas, there are important natural areas that will
not survive the five years required for phasing
in the program. We only get 20% of $4 million
over the five-year period, 20%, 40%, 60%,
80%, and 100%, respectively. We desperately
need a stopgap appropriation or bond issue of

Vol. 34 Art. 4

about $15 million to acquire such areas before
they are lost. Otherwise, as we look forward to
achieving the ability to acquire outstanding
natural areas, we may have to watch some of
our most important natural areas slip between
our fingers.

Finally, we must ensure that resources
once acquired or protected are adequately
managed. I propose that a dollar be added to
the license fee for motor vehicles and that the
proceeds be dedicated to maintenance and
management of natural lands, thereby helping
to compensate for the slaughter of wildlife on
our highways. Now I’ve been told everybody
and their brother has tried to get a piece of that
action, but the very obvious cause-effect
relationship between transportation develop-
ment and loss of wildlife through habitat
conversion and habitat fragmentation, not to
mention direct wildlife mortality, is so obvious
that I believe the public would embrace the
surcharge if given the chance.

Thank you for your attention. Thank you
for coming, and I look forward to working with
all of you in these efforts in the future. Remem-
ber, we have an obligation to be objective, to
treat all development interests fairly, that is,
equally, but we must refrain from compromise.
We've already lost too much.

Appendix One: Native Illinois Species and Related Bibliography

Susan L. Post, Illinois Natural History Survey

The assemblage of living forms native to Illinois
...are held together as a definitely organized,
living whole. —Stephen A. Forbes, 1889

The Illinois State Agricultural Society was
formed in 1853 and brought zoologists and
botanists together in an organized natural
history society. In the first transactions of the
Agricultural Society, three Illinois species lists
were published: The Birds of Southern Illinois
by H. Pratten (1855), The Mollusca of Southern
Illinois by H.A. Ulffers (1855), and The Animals
of Cook County by R. Kennicott (1855). These
were the first attempts to list the species of
Illinois.

By the turn of the century, biologists from
the State Laboratory of Natural History, later to
become the Illinois Natural History Survey,
were systematically sampling the state. These
early field investigations formed the basis for
understanding our ecosystems and the natural
histories of the organisms found in them.
Because of these early records, comparisons can
be made between conditions that exist today and
those that existed a century ago. From its first
publication in 1876, Stephen A. Forbes’ List of
Illinois Crustacea, to its most recent, the Survey
has concerned itself not only with cataloging
organisms and their distributions in the state but
also with the relationships of these organisms to
their environments. The Survey's long existence
has allowed continuity. Field studies have been
and continue to be repeated at intervals, and
long-term changes in populations and natural
habitats have thereby been documented.

E.O. Wilson (1988) notes in his recent
discussion of biological diversity that we do not
know the true number of species on Earth,
possibly even to the nearest order of magnitude.
The same is true for Illinois. We are fairly
certain of the numbers of our more visible fauna
in the Phylum Cordata—the reptiles, amphibi-
ans, fishes, birds, and mammals. In other phyla,
however, we are less certain. Research on many
of these groups is at an early stage, and new

species are frequently found. Even though we
list approximately 17,000 insects, this number
is only an approximation. The nematodes,
which may outnumber even the insects, are an
even more difficult group to estimate. The vast
majority of the species in Illinois remain
unmonitored. Like the dead in Gray’s Elegy
Written in a Country Churchyard, they may
pass from the Earth unnoticed and unknown.

The list of species native to Illinois that
follows was not generated by a single biologi-
cal survey but is the result of a search of the
literature and a query of systematists familiar
with the organisms of Illinois. Sources are
listed in the bibliography and in the acknowl-
edgments. The list is divided into five king-
doms: Monera, Protista, Fungi, Plantae, and
Animalia (Whittaker 1959). Classification of
the invertebrates follows Brusca and Brusca
(1990), and plant nomenclature follows
Mohlenbrock (1986).

The numbers of certain groups were
impossible to estimate and are listed as
unknown—the bacteria, nematodes, and
protozoa. According to the Bacteriological
Code (1958), bacteria cannot be described as
simply as other organisms. Every individual is
treated as belonging to a number of categories
of consecutive rank. Only the individual is
considered “real.” Until the taxonomic prob-
lems have been solved, no list of species for
Illinois can be constructed. Although the
protozoa are divided into seven phyla (Levine
et al. 1980), we have left them as the generic
“protozoa.” Much of protozoan systematics is
still in the alpha stage, with thousands of
species yet to be discovered and classified
(Lee et al. 1985). Few invertebrate groups
illustrate the diversity in form, habitat, and
behavior found in the nematodes. An examina-
tion of virtually any organic substrate com-
monly yields nematode specimens represent-
ing undescribed species. The systematics of
this group is in an embryonic stage.

463

464 Illinois Natural History Survey Bulletin

Although the class Insecta is very large
and new species are continually being de-
scribed, an estimate was made by consulting
specialists for each group. The species number
for Coleoptera (J. Bouseman, pers. comm.),
Hymenoptera (W. LaBerge, pers. comm.), and
Diptera (D. Webb, pers. comm.) are only
estimates. The number of Diptera was deter-
mined by randomly choosing 1,000 species
from A Catalog of the Diptera of America
North of Mexico (Stone et al. 1965) and
determining how many of those occur in
Illinois. This process was replicated three times
and a homogeneity chi square was used to de-
termine if the three samples could be lumped.
A nonsignificant x° indicated that the three
samples could be combined and the mean
determined. The percent of species found to
occur in Illinois was multiplied by 17,000
(number of species of Diptera in North Amer-
ica) to estimate the number in Illinois.

Only a small fraction of the Illinois fungi
are known, but estimates suggested that Illinois
has at least 20,000 species (L. Crane, pers.
comm.). The number of species of mites in the
order Acari was estimated based on the number
of mite species in Canada and the assumption
that the total number of mites in Illinois would
equal half the number of insect species in the
state (J. Kethley, pers. comm.). In the class
Aves, the number of species includes native
breeding species and migrants.

Determining the numbers of species that
are extirpated from the state or extinct is
difficult. With the exception of the showiest
birds, mammals, and flowering plants, biolo-
gists are reluctant to say with finality that a
species has come to its end. The possibility
always exists that a few individuals or a
population will be discovered in some remote
habitat. As with species numbers, we know
with near certainty that some of the more con-
spicuous fauna have been extirpated; we are
less certain about other species.

Species thought to no longer exist in
Illinois are listed in Table 1A along with the
source from which the determination was
made. The plant list was compiled using
Sheviak (1978), Paulson and Schwegman
(1976), Paulson et al. (1976), and Bowles et al.
(1991), and was reviewed by M.L. Bowles, J.E.
Ebinger, D.M. Ketzner, G. Kruse, S. Lauzon,
L.R. Phillippe, K.R. Robertson, J. Schwegman,
M.K. Solecki, and J.B. Taft. The final list was
reviewed by K.R. Robertson.

Vol. 34 Art. 4

Included in Table 1A are species listed in
the 1990 Illinois Endangered Species Protec-
tion Board’s Checklist of Endangered and
Threatened Animals and Plants of Illinois but
now considered extirpated. Not included are
three species of birds, two species of mammals,
and one plant species that disappeared from the
state and were successfully reintroduced—
peregrine falcon, ruffed grouse, wild turkey,
white-tailed deer, beaver, and lakeside daisy.
Species that no longer occur in the United
States are indicated.

The bibliography that concludes this
appendix lists all publications that were used to
create the list of native Illinois species and the
table of extirpated species.

ACKNOWLEDGMENTS

I would like to thank the following people from
the Illinois Natural History Survey: Lawrence
Page and Michael Jeffords for their advice and
comments, Kenneth Robertson for his help with
the extirpated plant list, Monica Lusk for her
library assistance, and Kathryn McGiffen and
Kathleen Methven for their help with the insect
collection of the Survey. The following people
gave invaluable species information: John
Bouseman, J. Leland Crane, Kevin Cummings.
George Godfrey, Joyce Hofmann, Wallace
LaBerge, David Ketzner, Marcos Kogan,
Joseph Maddox, Patti Malmborg, Philip Nixon,
Loy R. Phillippe, John Taft, David Voegtlin,
Donald Webb, and Mark Wetzel—all of the
Illinois Natural History Survey: Merrill Foster,
Bradley University; John Ebinger, Eastern
Illinois University; Helen Pigage, Elmhurst
College: John Kethley, Field Museum of
Natural History; Kenneth Christiansen,
Grinnell College; Burt Shepard, Harza Engi-
neering Company; Glen Kruse, Susan Lauzon,
and John Schwegman, Illinois Department of
Conservation; Mary Kay Solecki, Illinois
Nature Preserves Commission; Everett Cashatt,
Illinois State Museum; Edward Mockford,
Illinois State University; Bill McKnight,
Indiana State Museum; Clyde Robbins, Loyola
University; Marlin Bowles, Morton Arboretum;
Max Hutchison, Natural Land Institute; Joseph
Beatty and George Garoian, Southern Illinois
University at Carbondale: Robert Allen,
University of Arkansas; Ellis Macleod and
James Sternburg, University of Illinois at
Urbana-Champaign; and Michael Morris,
Western Illinois University.

April 1991 Symposium Proceedings: Our Living Heritage 465

LIST OF NATIVE ILLINOIS TAXA (AND NUMBERS OF SPECIES)

Kingdom Monera (1 12° species)
Division Schizophyta: bacteria (number of species unknown)
Division Cyanophyta: blue-green algae (112 species)

Kingdom Protista (1,406° species)
Division Protozoa: (number of species unknown)
Division Euglenophyta: euglenoids (30 species)
Division Chrysophyta: diatoms and golden brown algae (440 species)
Division Pyrrophyta: fire algae (20 species)
Division Chlorophyta: green algae (507 species)
Division Phaeophyta: brown algae (0 species)
Division Rhodophyta: red algae (5 species)
Division Myxomycota: plasmodial slime molds (400 species)
Division Acrasiomycota: cellular slime molds (2 or 3 species)
Division Plasmodiophoromycota: (1 species)

Kingdom Fungi (~ 20,000 species)
Division Chytridiomycota: chytrids (~ 300 species)
Division Oomycota: water molds (~ 300 species)
Division Zygomycota: bread molds (~ 400 species)
Division Ascomycota: sac fungi (~ 9,000 species including 500 species of lichens)
Division Basidiomycota: club fungi (~ 5,000 species)
Division Deuteromycota: fungi imperfecti (~ 5,000 species)

Kingdom Plantae (2,574 species)
Division Bryophyta
Class Anthocerota: hornworts (3 species)
Class Hepaticae: liverworts (118 species)
Class Musci: mosses (385 species including 2 extirpated species)
Division Lycodiophyta: club mosses, quillworts, and spike mosses (12 species including 3 endangered
species of clubmosses and | extirpated species of quillwort)
Division Equisetophyta: horsetails (12 species including 3 endangered and 1 extirpated species)
Division Filicophyta: ferns (75 species including 11 endangered, 3 threatened, and 2 extirpated species)
Division Coniferophyta: conifers (14 species, including 4 endangered and 3 threatened species)
Division Anthophyta: monocots and dicots (1,955 species including 275 endangered, 54 threatened, 53
extirpated, 1 extinct, and | extirpated but reintroduced species)

Kingdom Animalia (29,662° species)
Phylum Porifera: sponges (14 species)
Phylum Cnidaria: polyps and jellyfish
Class Hydrozoa: hydra and freshwater jellyfish (<10 species of hydra and | species of freshwater jellyfish)
Phylum Platyhelminthes: flatworms (400 species)
Phylum Nemertea: ribbon worms (1 species)
Phylum Nematoda: nematodes (number of species unknown)
Phylum Nematomorpha: horsehair worms (2 species)
Phylum Acanthocephala: spiny-headed worms (27 species including | species found in the endangered
greater prairie-chicken)
Phylum Gastrotricha (60 species)
Phylum Rotifera: rotifers (150-175 species)
Phylum Entoprocta (1 species)
Phylum Annelida: segmented worms
Class Oligochaeta: “earthworms” (20 terrestrial and 83 aquatic species)
Class Hirudinea: leeches (32 species)
Class Aphanoneura (3 species)
Class Branchiobdeilida: crayfish worms (9 species)

466

Illinois Natural History Survey Bulletin

Phylum Arthropoda
Class Chelicerata (10,598* species)

Subclass Arachnida
Order Scorpiones: scorpions (1 species)
Order Araneae: spiders (530 species)
Order Pseudoscorpionida: pseudoscorpions (28 species)
Order Opiliones: daddy long-legs (19 species)
Order Acari: mites and ticks (20 species of ticks and ~10,000 species of mites)

Class Myriapoda (74 species)

Subclass Diplopoda: millipedes (29 species)
Subclass Pauropoda: pauropods (5 species)

Subclass Chilopoda: centipedes (37 species)
Subclass Symphyla: symphylans (3 species)

Class Insecta (~17,000 species)

Subclass Myrientomata
Order Proturans: proturans (6 species)
Subclass Oligoentomata
Order Collembola: springtails (73 species)
Subclass Diplurata
Order Diplura: diplurans (6—10 species)
Subclass Zygoentomata
Order Thysanura: silverfish (6* species)
Subclass Pterygota
Order Ephemeroptera: mayflies (126 species)
Order Odonata: dragonflies (98 species) and damselflies (44 species)
Order Blattodea: cockroaches (9 species)
Order Mantodea: mantids (1 species)
Order Isoptera: termites (5 species)
Order Plecoptera: stoneflies (57 species)
Order Orthoptera: grasshoppers, crickets, and katydids (157 species)
Order Dermaptera: earwigs (3 species)
Order Phasmida: walking sticks (5 species)
Order Zoraptera: zorapterans (1 species)
Order Psocoptera: book and bark lice (91 species)
Order Hemiptera: true bugs (910 species)
Order Thysanoptera: thrips (200 species)

Vol. 34 Art. 4

Order Anoplura: sucking lice (18 native and 19 nonnative [from domestic animals and man] species)
Order Mallophaga: biting lice (280 species including | extinct species that occurred on the passenger

pigeon)
Order Homoptera: plant bugs (1,485 species)
Order Strepsiptera: twisted-wing insects (15—20 species)
Order Coleoptera: beetles (5,000 species)

Order Neuroptera: lacewings, antlions, alderflies (45 species including | extirpated species)

Order Hymenoptera: bees, ants, wasps (2,000* species)
Order Mecoptera: scorpionflies (18 species)

Order Siphonaptera: fleas (33 species including | species that occurs on the endangered Eastern

wood rat)
Order Diptera: true flies, mosquitoes, and gnats (4,100 species)
Order Trichoptera: caddisflies (184 species)

Order Lepidoptera: butterflies and moths (2,000 species including | endangered, 2 threatened. and

5 extirpated species)

Subphylum Crustacea

Class Branchiopoda (52 species)
Order Anostraca: fairy shrimp (4 species)
Order Cladocera: water fleas (~43 species)
Order Conchostraca: clam shrimp (5 species)
Class Maxillopoda (84 species)
Subclass Ostracoda: seed shrimp (53 species)
Subclass Copepoda (21 species)
Subclass Branchiura: fish lice (10 species)

April 1991

Class Malacostraca (71 species)

Symposium Proceedings: Our Living Heritage

467

Order Decapoda: crayfish (23 species including 4 endangered and 2 extirpated species)
Order Isopoda: pillbugs (28 species including 1 endangered species)
Order Amphipoda: scuds (19 species including 5 endangered and | threatened species)

Order Musida: opossum shrimp (1 species)

Phylum Pentastomida: tongue worms (no species found in native fauna)

Phylum Tardigrada: water bears (13 species)
Phylum Mollusca

Class Gastropoda: snails (170 species including | endangered species)
Class Bivalvia: mussels and clams (104 species including 29 endangered, 4 threatened, 16 extirpated, and

4 extinct species)
Phylum Ectoprocta (9 species)
Phylum Chordata

Subphylum Vertebrata

Class Agnatha: lampreys and jawless fish (6 species including 1 endangered and | threatened species)
Class Osteichthyes: boney fishes (181 species including 12 endangered, 14 threatened, and 12 extir-

pated species)

Class Amphibia: amphibians (39 species including 2 endangered, | threatened, and | presumed

extirpated species)

Class Reptilia: reptiles (59 species including 5 endangered, 4 threatened, and | presumed extirpated

species)

Class Aves: birds (297 native breeding and migrant species including 37 endangered, 6 threatened,
8 extirpated, 4 extinct, and 3 extirpated but reintroduced species)
Class Mammalia: mammals (67 species including 7 endangered, 3 threatened, 9 extirpated, and

2 extirpated but reintroduced species)

Total number of species: 53,754+
Total number of extirpated species: 115

Total number of threatened and endangered species: 497

Table 1A. Native Illinois species presumed extirpated.

Scientific name

Common name

KINGDOM PLANTAE

Division Bryophyta
Brachylema subulatum (P. Beauvois)
Schimper ex Cardot
Neckera pennata Hedwig

Division Lycodiophyta
Isoetes engelmannii A. Braun

Division Equisetophyta
Equisetum palustre L.

Division Filicophyta
Asplenium ruta-muraria L.
Woodwardia virginica (L.) J.E. Smith

Division Anthophyta
Apios priceana Robinson
Arabis drummondii Gray
Arethusa bulbosa L.
Bacopa acuminata (Walter) B.L. Robinson
Baptisia tinctoria (L.) R. Brown
Carex cumulata (Bailey) Fernald
Carex plantaginea Lamarck
Cinna latifolia (Treviranus) Grisebach
Cirsium pitcheri (Torrey & Eaton) Torrey & Gray
Clintonia borealis (Aiton) Rafinesque
Corallorhiza trifida Chatelain
Daucus pusillus Michaux

Moss
Moss

Englemann’s quillwort
Marsh horsetail

Wall-rue spleenwort
Chain fern

Price’s groundnut
Rock cress

Dragon’s mouth
Purple hedge-hyssop
Yellow wild indigo
Sedge

Sedge

Drooping wood reed
Dune thistle
Bluebead lily

Pale coral root orchid
Small wild carrot

Source

McKnight pers. comm.
McKnight pers. comm.

Mohlenbrock 1967
Bowles et al. 1991

Mohlenbrock 1967
Bowles et al. 1991

Schwegman pers. comm.
Swink & Wilhelm 1979
Sheviak 1974

Bowles et al. 1991
Bowles et al. 1991
Bowles et al. 1991
Bowles et al. 1991
Bowles et al. 1991
Bowles pers. comm.
Swink 1988

Sheviak 1974

Bowles et al. 1991

468

Illinois Natural History Survey Bulletin

Vol. 34 An. 4

Scientific name

Delphinium carolinianum Walter

var. penardii (Huth) Warnock
Elatine brachysperma Gray
Eleocharis caribaea (Rottboell) Blake
Eleocharis equisetoides (Elliott) Torrey
Epigaea repens L. var. glabrifolia Fernald
Erianthus brevibarbis Michaux
Fuirena scirpoides Michaux
Gaillardia aestivalis (Walter) Rock
Geum rivale L.
Glyceria canadensis (Michaux) Trinius
Gnaphalium macountt Greene
Gratiola aurea Muhlenberg
Hippuris vulgaris L.
Hypericum ellipticum Hooker
Linnaea borealis L. ssp. americana (Forbes) Hulten
Malaxis monophylla (L.) Swartz
Malaxis unifolia Michaux
Nemopanthus mucronata (L.) Trelease
Oryzopsis asperifolia Michaux
Oryzopsis pungens (Torrey) Hitchcock
Paspalum lentiferum Lamarck
Plantago heterophylla Nuttall
Platanthera (Habenaria) dilatata (Pursh) Hooker
Platanthera (Habenaria) hookeri Torrey
Platanthera (Habenaria) orbiculata (Pursh) Torrey
Polygala paucifolia Willdenow
Potamogeton epihydrus Rafinesque
Potamogeton vaseyi J.W. Robbins
Ranunculus ambigens S. Watson
Ranunculus gmelinii DC.

var. hookeri (D. Don) L. Benson
Schedonnardus paniculatus (Nuttall) Trelease
Scheuchzeria palustris L. var. americana Fernald
Scirpus microcarpus Presl
Scirpus pedicellatus Fernald
Scirpus subterminalis Torrey
Sparganium minimum (Hartman) Fries
Thismia americana N.E. Pfeiffer!
Trautvetteria caroliniensis (Walter) Vail
Trifolium stoloniferum Eaton
Trillium cernuum L.
Valerianella patellaria (Sullivant) Wood

KINGDOM ANIMALIA
Phylum Arthropoda
Class Insecta
Columbicola extinctus Malcomson

Hesperia dacotae (Skinner)
Notodonta simplaria Graef
Pieris napi oleracea (Harris)
Schinia indiana (J.B. Smith)
Speyeria diana (Cramer)
Sympherobius occidentalis Fitch
Class Malacostraca
Cambarus robustus Girard
Macrobrachium ohione (Smith)

Common name

Prairie larkspur
Waterwort

Spike rush

Horsetail spike rush
Trailing arbutus
Brown plume grass
Umbrella grass
Blanket flower
Purple avens
Rattlesnake manna grass
Western cudweed
Goldenpert

Mare’s tail

St. John’s wort
Twinflower

Adder’s mouth orchid
Adder’s mouth orchid
Mountain holly

Rice grass

Rice grass

Bead grass

Small plantain

White orchis
Hooker’s orchid
Round-leaved orchid
Flowering wintergreen
Pondweed

Pondweed

Spearwort

Small yellow crowfoot
Tumble grass

Arrow grass

Bulrush

Bulrush

Bulrush

Least bur-reed
Thismia

False bugbane
Running buffalo grass
Nodding trillium
Corn salad

Chewing louse on
passenger pigeon
Dakota skipper
Simple promenant
Mustard white
Indiana schinia
Diana fritillary
Brown lacewing

Lusty crayfish
Ohio shrimp

Source

Mohlenbrock 1981
Mohlenbrock 1978
Mohlenbrock 1976
Bowles et al. 1991
Swink & Wilhelm 1979
Mohlenbrock 1973
Bowles et al. 1991
Mohlenbrock 1986
Bowles et al. 199
Bowles et al. 1991
Bowles et al. 1991
Swink & Wilhelm 1979
Swink & Wilhelm 1979
Mohlenbrock 1978
Swink & Wilhelm 1979
Sheviak 1978

Sheviak 1978
Mohlenbrock 1978
Mohlenbrock 1972
Mohlenbrock 1972
Bowles et al. 1991
Bowles et al. 1991
Sheviak 1974

Bowles et al. 1991
Sheviak 1974

Swink & Wilhelm 1979
Mohlenbrock 1970a
Bowles et al. 1991
Bowles et al. 1991

Swink & Wilhelm 1979
Mohlenbrock 1972
Bowles et al. 1991
Bowles et al. 1991
Bowles et al. 1991
Swink & Wilhelm 1979
Mohlenbrock 1970a
Mohlenbrock 1983
Mohlenbrock 1981
Schwegman 1989
Bowles et al. 1991
Sheviak 1978

Malcomson 1937
Sternburg pers. comm.
Godfrey pers. comm.
Irwin & Downy 1973
Godfrey pers. comm.
Irwin & Downy 1973
Macleod pers. comm.

Page 1985

April 1991

Scientific name

Phylum Mollusca

Class Bivalvia
Epioblasma flexuosa (Rafinesque)'
Epioblasma obliquata (Rafinesque)
Epioblasma personata (Say)'
Epioblasma propinqua (Lea)!
Epioblasma rangiana (Lea)
Epioblasma sampsonii (Lea)!
Epioblasma torulosa (Rafinesque)
Fusconaia subrotunda (Lea)
Hemistena lata (Rafinesque)
Lampsilis abrupta (Say)
Leptodea leptodon (Rafinesque)
Obovaria retusa (Lamarck)
Plethobasus cicatricosus (Say)
Pleurobema plenum (Lea)
Quadrula fragosa (Conrad)
Villosa fabalis (Lea)

Phylum Cordata

Class Osteichthyes
Atractosteus spatula (Lacépéde)
Coregonus nigripinnis (Gill)
Crystallaria asprella (Jordan)
Esox masquinongy Mitchill
Etheostoma histrio Jordan & Gilbert
Hybopsis amblops (Rafinesque)
Ichthyomyzon bdellium (Jordan)
Lythrurus ardens (Cope)
Noturus stigmosus Taylor
Percina evides (Jordan & Copeland)
Percina uranidea (Jordan & Gilbert)

Pteronotropis hubbsi (Bailey & Robison)

Class Amphibia

Cryptobranchus alleganiensis (Daudin)

Class Reptilia
Nerodia fasciata (Linnaeus)

Class Aves
Ajaia ajaja (Linnaeus)

Campephilus principalis (Linnaeus)!

Conuropsis carolinensis (Linnaeus)!
Corvus corax Linnaeus

Cygnus buccinator Richardson
Ectopistes migratorius (Linnaeus)!
Numenius borealis (Forster)!

Tympanuchus phasianellus (Linnaeus)

Class Mammalia
Bison bison (Linnaeus)
Canis lupus Linnaeus
Cervus elaphus Linnaeus
Erethizon dorsatum (Linnaeus)
Felis concolor Linnaeus
Martes americana (Turton)
Martes pennanti (Erxleben)
Peromyscus gossypinus (Le Conte)
Ursus americanus Pallas

'This species no longer occurs in the United States.

Symposium Proceedings: Our Living Heritage

Common name

Source

Leafshell

Catspaw

Round combshell
Tennessee riffleshell
Northern riffleshell
Wabash riffleshell
Tubercled blossom
Long-solid
Cracking pearlymussel
Pink mucket
Scaleshell

Ring pink

White wartyback
Rough pigtoe
Winged mapleleaf
Rayed bean

Alligator gar
Blackfin cisco
Crystal darter
Muskellunge
Harlequin darter
Bigeye chub
Ohio lamprey
Rosefin shiner
Northern madtom
Gilt darter
Stargazing darter
Bluehead shiner

Hellbender
Broad-banded watersnake

Roseate spoonbill
Ivory-billed woodpecker
Carolina parakeet
Common raven
Trumpeter swan
Passenger pigeon
Eskimo curlew
Sharp-tailed grouse

Bison

Gray wolf
Elk
Porcupine
Mountain lion
Marten
Fisher

Cotton mouse
Black bear

Cummings 1991
Cummings 1991
Cummings 1991
Cummings 1991
Cummings 1991
Cummings 199]
Cummings 1991
Cummings 1991
Cummings 1991
Cummings 199]
Cummings 1991
Cummings 199]
Cummings 1991
Cummings 1991
Cummings 199]
Cummings 1991

Burr 1991
Smith 1979
Smith 1979
Smith 1979
Burr 1991
Burr 1991
Smith 1979
Smith 1979
Burr 1991
Smith 1979
Smith 1979
Burr 1991

Morris pers. comm.

Morris pers. comm.

Bohlen 1989
Bohlen 1989
Bohlen 1989
Bohlen 1989
Bohlen 1989
Bohlen 1989
Bohlen 1989
Bohlen 1989

Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989
Hoffmeister 1989

469

470

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475

Appendix Two

County reference map. Readers who wish to
identify counties on maps shown in the text
will find this map a convenient reference.

476

Winnebago
1

Stephenson McHenry

Boone

DeKalb Kane
Whiteside
Kendall
Henry
Grundy
Mercer

Kankakee
Henderson Stark [saa
Waren Knox
an se

Vermilion
Logan Champaign

Macoupin

Calhoun

Bond

White

Hamilton

Johnson

ulaski :

Alexander P

Coffee break allowed Brooks Burr’s concern

time for speaker Joyce over threatened fish and
Hofmann to continue her dwindling aquatic habitat
advocacy on behalf of answered Thoreau’s
those troubled wetland query, “Who hears the
tenants, the swamp fishes when they cry?”
rabbit and rice rat. We do.

Louis Iverson’s use of satellite data piqued interest
in INHS Special Publication 11: Forest Resources
of Illinois with its 67 computer-generated maps.

Survey support staff set
up exhibits for the
symposium and rolled
posters for mailing. In

an economy drive, staff
collected the 450 paper
towel tubes used to mail
the posters!

James “Gene” Gardner’s research on caves intro-
duced us to the fragility and fascination of that dark
and silent habitat.

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NATURAL HISTGRY SUR‘
ILLINOIS
NATURAL
HISTORY MAY 10 1994
SURVEY
LIBRARY

matics of Leptosphaeria

Natural History Survey Bulletin
ume 34, Article 5

DEPOSITORY.
JUL 2 41992

UNIVERSITY OF ILLINOIS
AY URBANA-CHAMPAIGN

*)

ILLINOIS
NATURAL
HISTORY
SBURVEY

Systematics of Leptosphaeria
Species Found on the Rosaceae

Sabine M. Huhndorf

Illinois Natural History Survey

Illinois Natural History Survey Bulletin
Volume 34, Article 5
May 1992

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

A catalog of the publications of the Illinois Natural History Survey is available
without charge from the address below. A price list and an order blank are
included with the catalog.

Illinois Natural History Survey
Distribution Center

Natural Resources Building
607 East Peabody Drive
Champaign, Illinois 61820

Citation:

Huhndorf, S.M. 1992. Systematics of Leptosphaeria species found on the
Rosaceae. Illinois Natural History Survey Bulletin 34(5):479-534.

Editor: John P. Ballenot

Author’s current address: The New York Botanical Garden, Bronx, NY 10458.
US ISSN 0073-4918

Printed by Authority of the State of Illinois
(X0415-1,200-5-92)

Contents

Acknowledgments __ iv

Introduction 479
Materials and Methods 480
Discussion of Characters 480

Leptosphaeria Species Referable to the Pleosporales 482
Leptosphaeriaceae 482
Phaeosphaeriaceae 492
Lophiostomataceae 503

Leptosphaeria Species Referable to the Melanommatales 505

Leptosphaeria Species Referable to the Dothideales 508
Dothioraceae 508
Pseudosphaeriaceae 510

Leptosphaeria Species Referable to the Hymenoascomycetes 518
Clypeosphaeriaceae 518

Diaporthaceae 520

Amphisphaeriaceae 520

Species incertae sedis 525
Appendix: Leptosphaeria Species Described from the Rosaceae 526
Literature Cited 532

Index to Taxa 534

Acknowledgments

This research was supported by National Sci-
ence Foundation grant BSR 87-00065 to C.A.
Shearer, J.L. Crane, and D.L. Swofford and by
the Floyd Ingersoll Fellowship, Department of
Plant Pathology, University of Illinois, Fall
1989. The Friends of the Farlow Fellowship
supported research on specimens at the Farlow
Herbarium, and the H.H. Ross Memorial Grant
from the Illinois Natural History Survey sup-
ported research on specimens at the New York
Botanical Garden. I would also like to acknowl-
edge and thank the Morton Arboretum for the

Baker Fellowship, which helped in the
completion of the manuscript. I thank the
curators of the herbaria for the material made
available for study. I thank J.L. Crane, D.A.
Glawe, and ME. Barr Bigelow for reviewing
the manuscript.

This work was originally part of a dis-
sertation submitted to the Graduate College of
the University of Illinois at Urbana-
Champaign in partial fulfillment of the re-
quirements for the degree of Doctor of Phi-
losophy in plant pathology.

Introduction

The genus Leptosphaeria was established by
Cesati and de Notaris (1863) to include 26
species. The original description was superfi-
cial by modern standards and relied primarily
on ascospore characteristics to delimit the
genus. Cesati and de Notaris described
ascospores as oblong or fusoid, two- to many-
celled, and hyaline becoming yellow to dark
brown. Because other structural features were
poorly defined, a wide range of ascomycetes
has been included in this genus. The 1,689 taxa
described in Leptosphaeria (Crane and Shearer
1991) represent, according to current concepts
of ascomycete classification, a mixture of
Hymenoascomycetes and Loculoascomycetes.
Crane and Shearer (1991) and Miiller (1950)
provide good reviews of the historical back-
ground of the genus Leptosphaeria. Holm
(1957) and Shoemaker (1984a) consider a
limited number of species.

A large number of intergeneric transfers
of Leptosphaeria species have been made in the
past SO years. Five genera—Phaeosphaeria
Miyake, Paraphaeosphaeria Eriksson,
Nodulosphaeria Riess, Entodesmium Riess, and
Ophiobolus Riess—have become accepted
repositories for many Leptosphaeria species
(Holm 1957; Leuchtmann 1984; Shoemaker
1976, 1984a,b). Paraphaeosphaeria,
Nodulosphaeria, and Entodesmium are well
defined by several morphological features,
including ascocarp wall structure and ascospore
characteristics (Shoemaker 1984b, Shoemaker
and Babcock 1985). Phaeosphaeria is sepa-
rated from Leptosphaeria by several morpho-
logical characters and host specialization but
has itself become a very large and diverse
genus (Shoemaker and Babcock 1989b).
Ophiobolus species intergrade with long-spored
species of Leptosphaeria (Shoemaker 1976).
For a key to genera variously allied with
Leptosphaeria, see Shoemaker (1984a) or
Leuchtmann (1984).

Holm (1957) considered 62 species of
Leptosphaeria and restricted the genus to those
species most similar to the type of the genus,
Leptosphaeria doliolum (Pers.:Fr.) Ces. & de
Not. He emphasized the anatomy of the
ascocarp wall and found that in most
Leptosphaeria species the wall consisted of
thick-walled cells termed scleroplectenchyma.
He also emphasized the identity of the substrate
and the structure of the ascocarp relative to the
substrate in relegating a number of
Leptosphaeria species to other genera. Ex-
cluded species were distributed in
Nodulosphaeria, Phaeosphaeria, and
Entodesmium. He gave considerable weight to
the family of host plants in distinguishing these
genera. Holm’s (1957) scheme was accepted
by Shoemaker (1984a) in his treatment of
Canadian species of Leptosphaeria.

In recent years, Leptosphaeria has been
included in various orders in the Bitunicatae or
the Loculoascomycetes. Luttrell (1973) placed
Leptosphaeria in the family Pleosporaceae of
the Pleosporales. The single order Dothideales
was used by von Arx and Miiller (1975) for all
fungi classified as Loculoascomycetes; they
retained Leptosphaeria in the Pleosporaceae.
Eriksson and Hawksworth (1986) classified the
genus in the family Phaeosphaeriaceae of the
Dothideales. Barr (1987a) placed
Leptosphaeria in the family Leptosphaeriaceae
of the Pleosporales. Most recently, Eriksson
and Hawksworth (1990) accepted the family
Leptosphaeriaceae in the order Dothideales.
Bart’s classification of the Ascomycota (1983,
1987a) is employed in this dissertation because
of its relationship to previous classification
schemes (Luttrell 1973) and its detailed
explanation of the relationship of taxa. Barr’s
orders and families are based upon combina-
tions of developmental and morphological
characteristics that reflect the widespread
diversity within the group. Barr’s classification
provides a key to higher taxa and is practical

Illinois Natural History Survey Bulletin

for identification and for determining the
placement of organisms not yet included in the
scheme.

The present research reassesses the
taxonomic position of species of Leptosphaeria
described from plants in the family Rosaceae.
In doing so, this work emphasizes a concept of
Leptosphaeria based on its type species.
Because discrete groups of morphologically
similar species were found on related hosts
(Nodulosphaeria on Compositae, Phaeo-
sphaeria on Gramineae and Entodesmium on
Leguminosae), a goal of the study was to
determine whether a distinct group of related
species could be found on Rosaceous hosts.
Also, approaching the taxonomy of this large,
complex genus on the basis of host family is a
convenient way of dividing the large numbers
of Leptosphaeria species into smaller, work-
able groups.

Fifty-five species described from
Rosaceous hosts have been included in
Leptosphaeria (see appendix). Efforts were
made to locate and obtain type material for all
of the described species. The names of
herbaria providing material are abbreviated in
this dissertation according to Index
Herbariorum (Holmgren et al. 1990). When
possible, type specimens were studied, supple-
mented by other collections, and the species
were redescribed and illustrated from the type
material. Unfortunately, type material could
not be located for all of the species; in such
cases, descriptions were based on other
material only when the resulting concept of the
species was obviously in accordance with the
original description. From this work, it became
apparent that no discrete group of related
species is specialized on the Rosaceae. The
species studied were determined to represent a
mixture of Loculoascomycetes and
Hymenoascomycetes, and non-Leptosphaeria
species were reassigned to more appropriate
genera.

Materials and Methods

Several techniques were employed to assess
character states of specimens. Fungi were
observed microscopically in water mounts for
details of asci, ascospores, hamathecium,
centrum, and ascocarp wall surface, using
bright-field and Nomarski differential interfer-
ence contrast. India ink was used as a negative

Vol. 34 Art. 5

stain to reveal ascospore sheaths. Melzer’s
reagent (0.5 g iodine, 1.5 g KI, 20.0 g chloral
hydrate, 20.0 ml distilled water) was used to
observe amyloid reactions in unitunicate asci.
Semipermanent mounts were made in lacto-
phenol, and many of the photomicrographs
were made from these mounts. Micrographs
were made using Kodak T Max 100 and Kodak
Technical Pan film 2415 (Eastman Kodak
Company, Rochester, New York). For scan-
ning electron microscopy, dry ascocarps and
substrate were used direct from the herbarium
specimen, without any preparation. Specimens
were viewed with an Amray 1830 scanning
electron microscope.

The order of septation in ascospores is
given following Shoemaker (1984a). The
sequence of septation is recorded chronologi-
cally, with | being the first septum formed. A
series of numbers separated by colons indicates
the sequence in which the septa form. The first
number represents the septum nearest the
ascospore apex, and the last number is the
septum nearest the base. Unusual or infrequent
occurrences are recorded within parentheses
(Shoemaker 1984a).

To observe details of ascocarp anatomy,
semi-thin sections of the ascocarp were needed
for light microscopy. Fixation and embedding
techniques for plastic were developed to obtain
good, uniform sections from dried fungal
material in a relatively short period of time.
The techniques for fixation and embedding are
given in Huhndorf (1991).

Discussion of Characters

Semi-thin sections of ascocarps were used to
assess characteristics of the ascocarp walls.
Wall characteristics have proved useful in
taxonomic placement of species but may have
limitations in the routine identification of
specimens. It may be possible to observe this
feature with hand sections or frozen microtome
sections, as done by Shoemaker (1984a).
Ascocarp walls of cells radiating in surface
view (textura prismatica) and thick-walled cells
(scleroplectenchyma) are good indicators
within Leptosphaeria of alliance with L.
doliolum. Wall cells in surface view are seen
easily from crush mounts. Semi-thin sections
and scanning electron microscopy give a good
indication of the relationship of the ascocarp to
the substrate. This character has also proved

May 1992

important in delimiting Leptosphaeria; in-
cluded species have ascocarps that are superfi-
cial or that become superficial. Semi-thin
sections also show ostiole structure, a charac-
teristic that is important in related genera such
as Nodulosphaeria.

Ascospore characters, in combination
with other characters, are useful for placement
of species. Leptosphaeria species have more or
less fusiform, three- to multi-septate ascospores
with some pigmentation. These character states
are continuous rather than discrete, and in
practice, divisions are made in a somewhat
arbitrary manner. Also, ascospore pigmenta-
tion varies within a species and may reflect age,
environmental influences, and/or substrate.
Ascospore wall ornamentation often is difficult
to assess, especially when using type material,
which frequently is old and in poor condition.
In addition, type material is usually of uniform
developmental stage and frequently is sparse
and depleted by the activities of previous
researchers. Ascospore wall ornamentation is
difficult to assess with the light microscope
because of the limits of resolution, except in
cases for which there may be unusually
distinctive surface ornamentation. Using the
scanning electron microscope to resolve surface
features requires sufficient material of an
appropriate developmental stage, with the
ascospores released from the asci and with
surface detail not obscured by sheaths or
mucilaginous material. Unfortunately, this
situation does not usually exist in type material.
In recent collections with different develop-
mental stages it is easier to view ascospore wall
ornamentation. Although the taxonomic
significance of this character is unclear at this
time, largely because of the difficulties in-
volved in studying it, further studies would be
useful.

The importance of host specificity or
substrate preference in delimiting species or
genera needs further clarification. Whether
substrate preference does occur and can be used
as a taxonomic character can be demonstrated
only by extensive collection and comparison of
species from a variety of substrates, by com-
parative study of the morphology of pure
cultures of species obtained from a variety of
substrates and grown under identical environ-
mental conditions, and by determining experi-
mentally the range of substrates on which
isolates of species will grow and reproduce.

Leptosphaeria Species on the Rosaceae 481

The structure of the interascal filaments,
part of what Eriksson (1981) termed the
hamathecium, plays an important role in
distinguishing members of the Melanom-
matales from those in the Pleosporales accord-
ing to Barr’s (1987a) classification. Only one
member of the Melanommatales was found in
this group of species, and the diagnostically
important trabeculate pseudoparaphyses were
difficult to recognize. It was placed in that
group because of a combination of other
characters. The distinction between cellular
pseudoparaphyses (filaments that appear
septate at 450x magnification and branch and
anastomose) and trabeculate pseudoparaphyses
(thin filaments that do not appear septate at
450x magnification and branch and anasto-
mose) seems to be clear only in certain cases;
the pseudoparaphyses often appear as continu-
ous, rather than discrete, character states.
Trabeculate pseudoparaphyses can be mistaken
for cellular ones if septa appear regularly and
anastomoses are sparse.

Ascus morphology, emphasized by
Eriksson (1981), was not employed in this
study because of the difficulty in observing
modes of ascus dehiscence and details of ascus
wall layers in type and other herbarium
specimens. Also, in most cases, the use of
stains to enhance ascus wall layers gave
virtually no results.

Leptosphaeria Species Referable to the Pleosporales

All of the species in this chapter belong in the
Pleosporales (sensu Barr 1987a) and share a
combination of character states, any one of
which may deviate somewhat for a particular
taxon. These characters include cellular
pseudoparaphyses, asci in a basal layer, a
peridium that is usually pseudoparenchyma-
tous, and bipolarly asymmetrical ascospores.
The families included within the order are the
Leptosphaeriaceae, Phaeosphaeriaceae, and
Lophiostomataceae.

Leptosphaeriaceae

The Leptosphaeriaceae as defined by Barr
(1987a) include five genera united by the
characters of coelomycetous anamorphs, asci
that are narrower and thinner-walled than in the
Pleosporaceae, and ascocarp walls that consist
of relatively large, thick-walled or sclero-
plectenchymatous cells. The only genus in the
family treated here is Leptosphaeria.

Leptosphaeria V. Cesati & G. de Notaris,
Commentario della Societa Crittogamologica
Italiana, Milan 1:234. 1863. Nomen conser-
vandum.

Lectotype: Sphaeria doliolum C.H. Persoon:
E.M. Fries.

Ascocarps scattered or clustered, superficial or
immersed beneath epidermis, becoming
superficial as epidermis is shed, conic globose,
subglobose or depressed, glabrous, papillate.
Ascocarp wall often of radiating textura
prismatica in surface view; in longitudinal
section usually more than three cells thick,
composed of scleroplectenchyma, often with an
external crust. Papilla central, short, some-
times longer and beaklike, without hyaline
periphyses. Pseudoparaphyses broad or
narrow, septate, anastomosing, with or without
guttules, septal thickenings, and gelatinous

coating. Asci bitunicate, numerous in a broad
basal hymenium, cylindrical to clavate, short-
stalked, 8-spored in most, with spores often
biseriate or overlapping uniseriate. Ascospores
fusiform, clavate, 3 or more septate, primary
septum often median and often constricted,
some shade of yellow or brown, appendages or
sheath may or may not be present. Anamorphs
coelomycetous where known (see Crane and
Shearer 1991).

The concept of Leptosphaeria accepted
here is essentially that of Holm (1957), later
adopted by Shoemaker (1984a) with some
modification. The lectotypification of L.
doliolum (Shearer et al. 1990) represents a
basis for circumscribing the genus. More
emphasis is placed on scleroplectenchyma
found in the ascocarp walls than is considered
important by Shoemaker (1984a). As Shoe-
maker (1984a) noted, thin sections are essential
to show this thick-walled cell structure in
which the lumen of the cells is very small.
Emphasis is also placed on the wall tissue
arrangements in surface view, described using
the textura types of Korf (1958). Species
having a tissue arrangement similar to that of L.
doliolum, of cells radiating in surface view
(textura prismatica), are regarded as most
typical, but species without this character state
are not necessarily excluded. The position of
the ascocarp relative to the substrate is very
important, with included species having either
superficial ascocarps or ascocarps immersed
beneath the epidermis and becoming superficial
as the epidermis is shed. Because only type or
herbarium specimens were studied, anamorphs
were not considered because cultures could not
be made from this dead material.

Examination of 28 purported species of
Leptosphaeria from the Rosaceae revealed five
species that fit the adopted generic concept.
The excluded species are treated following
these five species.

May 1992

Leptosphaeria Species on the Rosaceae

Key to species of Leptosphaeria on the Rosaceae

483

AMBANS COCALP WALCO USAC ALC re. cons racetcevcrondecnessccscethoecessnrseisnacynseacvastcusverstesevaisescectss 2
HbeAscocarp)wall'cellsiandomly Oriented::.....-..sccecce.sacncoosesescevascaccecacstovsersarencesucences 4
2a. Ascocarp superficial or immersed becoming
superficial, ascospores mature inside CeMtruM............:.:cesesceeeeeeseeseeseeeeeeeees 3
2b. Ascocarp immersed beneath host cuticle,
ascospores mostly maturing on substrate surface.............:cceseeeee L. umbrosa
3a. Neck papillate, short conic, ascospores
narrowly fusiform, 22-28 x 4—5 LM.............sscessesersessseesseeseees L. doliolum
3b. Neck cylindrical, beaklike, ascospores
broadly fusiform, 17—20 x 5-8 LM... eeeeeeseeteeeteeeeee L. praetermissa

4a. Ascocarp immersed-subepidermal, wall pseudoparenchyma,
or partly scleroplectenchyma, on leaves of Cercocarpus
(provisionally retained within Leptosphaerid)..........ccscceseseeeseeseeeees L. cercocarpi

4b. Ascocarp erumpent to superficial, wall
scleroplectenchyma, on Dryds ............4..

eLeptosphaeria doliolum (C.H. Persoon:E.M.
Fries) V. Cesati & G. de Notaris, Commentario
della Societa Crittogamologica Italiana, Milan
1:234-235. 1863. See Figure 1.
= Sphaeria doliolum C.H. Persoon:E.M.
Fries, Icones et Descriptiones Fungorum
Minus Cognitorum, p. 39. 1800; E.M.
Fries, Systema Mycologicum Sistens
Fungorum 2:509. 1823.
= Leptosphaeria rustica P.A. Karsten, Fungi
Fenniae Exsiccati, Century 10, No. 964.
Anno 1870.
= Metasphaeria rustica (P.A. Karsten)
P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:157. 1883.

Ascocarps scattered, sparse, immersed-
subcuticular to erumpent, papillate, glabrous to
tomentose at base, globose, flattened at base,
300-325 um diameter, 250-300 um high.
Ascocarp wall of textura prismatica in surface
view, composed of cells radiating from apex
outward; in longitudinal section 45—55 um
thick at sides, 20-27 um thick at base, com-
posed of 12-15 layers of hyaline, polygonal,
isodiametric, scleroplectenchymatic cells (3.6—
6.5 um diameter), outer 2-3 layers somewhat
brown-melanized, with an external brown-
melanized crust. Papilla broadly rounded, not
fully formed, 130-135 tm high, 70-75 um
wide; wall 40-50 um thick, composed of 15—20
layers of small, brown-melanized, thick-walled,
polygonal, isodiametric cells (2.5—6.5 um
diameter), outer layers heavily brown-

L. dryadophila

melanized, inner layers hyaline, surrounding a
20-25 tum wide immature circular ostiole
formed from small hyaline cells, without
periphyses. Pseudoparaphyses 1|.0—2.0 um
wide, 150-175 tm long (height of the ascocarp
cavity), numerous, narrowly cellular, with
guttule-like thickenings at septa, without
gelatinous coating. Asci 90-125 x 5-9 um,
numerous, basal, cylindrical-clavate, with 8
overlapping uniseriate ascospores. Ascospores
22-28 x 4-5 um, narrowly fusiform, with acute
end cell shape, second cell slightly enlarged,
straight to slightly curved, 3-septate, septa
evenly distributed, order of septation 2:1:2,
primary septum median and slightly con-
stricted; light brown, guttulate, without
appendages, surrounded by gelatinous material.

Lectotype: As Sphaeria doliolum Pers.,
910.270-650 (L).

Exsiccatae: FINLAND: Tammela, on Spireae
ulmaria L. (=Filipendula ulmaria Maxim.),
Sept., Fungi Fenniae Exsiccati, Century 10, No.
964 (isotype of L. rustica, H, FH).

Comments: The collections of L. rustica, on
stems of Spiraea ulmaria L. in both of the
exsiccatae sets, were immature. No ascospores
were found, and asci were only beginning to
form. Karsten (1873) describes the ascospores
as elongate-fusoid, subhyaline, uniseptate with
several guttules in each cell, and 30—36 x S—6
lim in size. He probably described immature
spores. This information—along with the

Illinois Natural History Survey Bulletin

Vol. 34 Art. 5

Figure 1. Leptosphaeria doliolum. a. SEM of erumpent ascocarp, x125. b. longitudinal median section
through ascocarp, x150. c. longitudinal, median section through ascocarp wall, x950. d. ascus, x525.
e. ascocarp wall surface with radiating cells, xS5SO0. All from isotype of L. rustica.

radiating wall cells in surface view, the
scleroplectenchymatic wall cells in section, and
the location of the ascocarp relative to the
substrate—is consistent with the character of L.
doliolum, the type of the genus. Leptosphaeria
doliolum is found on many dead herbaceous
stems, although not previously described from
Spiraea.

*Leptosphaeria dryadophila S.M. Huhndorf

nom. nov. See Figure 2.

Etymology: Gr. philos = having affinity for.
= Melanomma dryadis C.J. Johanson in
Rabenhorst, Fungi europaei no. 3659
(1890). (Basionym).

= Leptosphaeria dryadis E. Rostrup,
Botanisk Tidsskrift 25:305. 1903.

Ascocarps clustered, erumpent to superficial,
papillate, glabrous to slightly tomentose toward
base, tomentum of dark brown hyphae, conic-
globose, 150-250 um diameter, 180-275 um
high. Ascocarp wall of textura angularis in
surface view; in longitudinal section uniformly
12—20 um thick (up to 27 lum thick near apex),
composed of 4—5 layers of polygonal, isodia-
metric to slightly elongate, scleroplecten-
chymatic cells, outer 2—3 layers composed of
brown-melanized cells (3.54.3 x 5.5-6.8 Lim),
inner 2—3 layers composed of hyaline, com-
pressed cells (2.1—3.5 x 10.0—-12.3 um).
Papilla conical, 60-65 um high, 25-50 um
wide at the apex, 90-110 uum wide at the base,
composed of 7—9 layers of isodiametric cells
(2.8—3.6 um diameter), wall 10-12 um thick
surrounding a 35-45 Lim wide ostiole without

May 1992 Leptosphaeria Species on the Rosaceae 485

~ }
'g
Figure 2. Leptosphaeria dryadophila. a. SEM of erumpent ascocarps, x50. b. ascocarp wall surface, x550.
c. longitudinal median section through ascocarp, x250. d. cellular pseudoparaphyses, x2,000. e. longitudinal
median section through ascocarp wall, x950. f. ascus, xSO0. g. ascospore, x1,150. a,c, e, and f from
holotype of L. dryadis; b, d, and g from isotype of L. dryadophila.

Illinois Natural History Survey Bulletin

criphyses. Pseudoparaphyses 0.7—3.0 um
wide, | 10-175 um long (height of the ascocarp
cavity), numerous, narrowly cellular, with
gelatinous coating. Asci (72—)90-118 x 10.8—
13.6 um, numerous, basal to slightly lateral,
cylindrical, thin-walled but with a thickened
rounded apex, short-stalked, with 8 biseriate
ascospores. Ascospores 20.1—24.5 x 5.7—7.2
um, fusiform to slightly clavate, with acute
end-cell shape, slightly curved; 3(-4-5)-septate,
septa evenly distributed, order of septation
2:1:2(:3:4), without constrictions; pale brown-
ish yellow, with ornamented wall, without
sheath or appendages.

Isotype: SWEDEN: Jamtland, Renfjallet, c. 900m,
in fructibus et calycibus emortuis Dryadis
octopetalae L. 13, Juli 1884, leg. C.J.
Johanson, Rabenhorst, Fungi europaei no. 3659
(1890) (FH, NY).

Exsiccatae: SWEDEN: Jamtland, Renfjallet, 13,
Juli 1884, leg. C.J. Johanson, Vgr., Micr. rar.
sel. 105 (FH).

Other material examined: ICELAND: Sponsgerdi,
on Dryas octopetala, Juli 12, 1901, O.
Davidson (type of Leptosphaeria dryadis, C).

Comments: This fungus was originally de-
scribed in Melanomma and was retained with
some reservation in that genus by Holm (1957,
1979), who stated, “The species is hardly a
Melanomma but its true affinities are doubtful
to me.” Melanomma dryadis differs from the
current concept of Melanomma (Barr 1987a) in
not having asci that arise peripherally within
the centrum and not having trabeculate
pseudoparaphyses. Barr (1982) erected the
genus Bricookea for a biologically similar
species, Bricookea sepalorum (Vleugal) Barr,
which is found on the inflorescences of Juncus
species. Melanomma dryadis differs from this
fungus in several respects: cells of the ascocarp
wall are scleroplectenchymatous, not pseudo-
parenchymatous; the ascocarp apex opens by a
pore, not by a slit; and the ascospores are not
hyaline. Melanomma dryadis, occurring in and
on the dead flowers and fruits of Dryas
octopetala, is a true Leptosphaeria, having the
characteristics of an erumpent to superficial
ascocarp with a wall of scleroplectenchymatous
cells and 3-septate yellow-brown ascospores. It
does not, however, show the radiating arrange-

Vol. 34 Art. 5

ment of cells of the ascocarp surface character-
istic of L. doliolum. Because the epithet
dryadis is pre-empted in Leptosphaeria by L.
dryadis Rostrup, a new name, Leptosphaeria
dryadophila is proposed for Melanomma
dryadis Johanson.

eLeptosphaeria praetermissa (P.A. Karsten)
P.A. Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:26. 1883. See Figure 3.
= Sphaeria praetermissa P.A. Karsten,
Fungi Fenniae Exsiccati, Century 9, No.
852. Anno 1869.

Ascocarps clustered, sparse, immersed-
subepidermal becoming erumpent, papillate,
glabrous to tomentose at base, globose,
flattened at base, 350-625 um diameter, 300—
500 um high. Ascocarp wall of textura
prismatica in surface view, composed of cells
radiating from apex outward; in longitudinal
section uniformly 32—58 um thick at sides and
base, composed of 7-8 layers of hyaline,
polygonal, isodiametric, scleroplectenchymatic
cells (5.7—13.6 x 3.6—7.2 um), inner 2—3 layers
somewhat elongate and compressed, with an
external brown-melanized crust. Papilla
beaklike, cylindrical, erumpent, 120-180 um
high, 75-135 um wide; wall 25-36 um thick,
composed of 5-6 layers of small, brown-
melanized, thick-walled, polygonal, isodiamet-
ric cells (4.3—7.2 um diameter), surrounding a
35-80 um wide circular ostiole formed from
small hyaline cells, without periphyses.
Pseudoparaphyses |.0—2.0 um wide, 180—260
tum long (height of the ascocarp cavity),
numerous, narrowly cellular, without gelati-
nous coating. Asei 100—130(—150) x 7-13 um,
numerous, basal, cylindrical, thin-walled, short-
stalked, rounded apex with apical chamber
present, with 8 biseriate ascospores. Ascos-
pores 17—20(—25) x 5.5—-8.0 um, broadly
fusiform, end cells acute to rounded, straight to
slightly curved; 3-septate, septa evenly distrib-
uted, order of septation 2:1:3 or 2:1:2, primary
septum median, with constrictions at all septa,
second cell occasionally enlarged (wider);
brownish-yellow, smooth, without sheath or
appendages.

Isotype: FINLAND: In caulibus emortuis Rubi
odorati et R. idaei in par. Tammela sat fre-

May 1992 Leptosphaeria Species on the Rosaceae 487

Figure 3. Leptosphaeria praetermissa. a. SEM of erumpent ascocarps, x65. b. ascocarp wall surface with
radiating cells, x550. c. longitudinal median section through ascocarp, x150. d. cellular pseudoparaphyses,
x2,000. e. longitudinal median section through ascocarp wall, x950. f. ascus, x525. g, h. ascospores, x1,150.
All from isotype of L. praetermissa.

Illinois Natural History Survey Bulletin

quenter per annum obvia. Fungi Fenniae
Exsiccati, Century 9, No. 852 (H, FH).

Other material examined: CANADA: Quebec:
Gatineau Provincial Park, Chemin Ridge Road,
on Rubus odoratus, 20 Jun. 1987, C.T.
Rogerson (NY); SwebeN: Jemtland: Five
collections, all on Rubus ideaus, Leg. A.G.
Eliasson, 20 May 1930, 27 Jun. 1930, 17 Jun.
1931, 19 Jun. 1931, 28 Jul. 1931 (S); Umea,
Apr. 1908, J. Vleugel (S).

Comments: Leptosphaeria praetermissa
warrants inclusion in the genus because of its
large erumpent ascocarp with a wall composed
of scleroplectenchymatous cells. As in L.
doliolum, the wall has an external crust, and the
cells form a radiating pattern in surface view.

It differs from L. doliolum in having a rather
long, cylindrical neck that grows through the
host epidermis before the ascocarp becomes
erumpent. This species seems to be found
predominantly in Europe but is probably not
common there. It was not encountered in any
collections of Rubus sp. in the United States but
was found in one collection from Canada.

eLeptosphaeria umbrosa G. Niessl in G.L.
Rabenhorst, Fungi Europaei Exsiccati,
Klotschii Herbarii vivi Mycologici Continuatio,
Edition 3 (Edita Nova), Series 2, Century 20,
No. 1934. Anno 1875; Just’s Botanisch
Jahresberichte 3:262. 1887. See Figure 4.
= Massaria umbrosa (G. Niessl) H. Rehm
in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 9:761. 1891.

Ascocarps scattered, immersed-subcuticular,
papillate, glabrous, globose, flattened at base,
150-300 um diameter, 90-200 Lm high.
Ascocarp wall of textura prismatica in surface
view, composed of cells radiating from apex
outward; in longitudinal section 4—6 um thick
at the base, composed of thin, hyaline, com-
pressed cells, 10.8—22 tm thick at the sides,
composed of 5—6 layers of polygonal,
scleroplectenchymatic cells, outer 2-3 layers of
isodiametric to slightly elongate, brown-
melanized cells (3.5 x 7-10 um) surrounded by
a brown-melanized crust, inner 2—3 layers of
hyaline, elongate-compressed cells (2-3 x 10-
18 um). Papilla bluntly conical, (18—)25-54
um high, 36-56 [tm wide at the apex, 60-70

Vol. 34 Art. 5

tum wide at the base, composed of 5-6 layers of
small, isodiametric, heavily brown-melanized,
thick-walled cells (2—3.6 um diameter), wall
14.422 um thick surrounding a 25-45 um
wide ostiole without periphyses. Pseudo-
paraphyses 0.7—2.0 um wide, 80-110 um long
(height of the ascocarp cavity), numerous,
narrowly cellular, with guttule-like thickenings
at septa, without gelatinous coating. Asci (72—)
80—100(—117) x 10.8—15.0 um, numerous,
basal, cylindrical-clavate, thick-walled,
rounded apex with apical chamber, short-
stalked, with 8 tri- to biseriate ascospores, most
commonly immature. Ascospores 27.3—33.0 x
4.3—7.2 um, hyaline to subhyaline when
immature, 28—37.4 x 5.7—9.3 um, pale brown
when mature, narrowly fusiform, with acute
end-cell shape, slightly curved; 4-septate, septa
unevenly distributed, order of septation 2:1:2:3,
primary septum supramedian and slightly
constricted, second cell slightly enlarged at
maturity, slightly roughened wall, continuous
sheath (0.7—1.5 um thick) surrounding imma-
ture spore, without appendages.

Holotype: Austria: An Spiraea Aruncus
[=Aruncus dioicus (Walt.) Fern.] bei Voitsberg
in Steiermark. Septbr. 1874 (M).

Exsiccatae: Austria: Voitsberg, Steiermark,
Septbr., G. v. Niessl, Rabenhorst, F. europaei
exs. 1934 (FH, NY isotypes); Voitsberg,
Steiermark, Aug. 1882, G. v. Niessl, Rehm,
Ascomyceten 690 (FH, NY authentic material):
Voitsberg, Steiermark, G. v. Niessl, Weese,
Eumycetes sel. exs. 638 (FH).

Other material examined: GERMANY:
Frauenfeld, on Solidago, October (NY).

Comments: Leptosphaeria umbrosa is interest-
ing because initially it seems to resemble
members of the Phaeosphaeriaceae having
small, pseudoparenchymatic-walled,
subcuticular ascocarps that lift the host cuticle
on maturity. Upon closer examination, it
reveals the characteristic features of
Leptosphaeria such as ascocarp walls com-
posed of scleroplectenchyma surrounded by an
external brown crust and wall cells that radiate
in surface view. Other distinctive characteris-
tics that may aid in the identification of this
species are the wide pseudoparaphyses with
prominant thickenings at the septa and mature

May 1992 Leptosphaeria Species on the Rosaceae 489

Figure 4. Leptosphaeria umbrosa. a. SEM of immersed ascocarps, x150. b. ascocarp wall surface with
radiating cells, x550. c. longitudinal median section through ascocarp, x200. d. cellular pseudoparaphyses
with thickenings at septa, x2,000. e. longitudinal median section through ascocarp wall, x950. f. ascus with
immature ascospores, x525. g. ascus with mature ascospores, x525. h. immature ascospore, x1,150.

i. mature ascospore from substrate surface, x1,150. All from holotype of L. umbrosa.

Illinois Natural History Survey Bulletin

brown ascospores that are often seen only on
the surface of the substrate surrounding the
ascocarp. Also diagnostic are the thin ascocarp
bases often left behind on the substrate when
ascocarps are removed from the substrate
surface.

*Leptosphaeria cercocarpi H. Sydow & P.
Sydow, Annales Mycologici 5:339. 1907. See
Figure 5.

Ascocarps clustered, abundant, immersed-
subepidermal, with surrounding stroma
extending into substrate, papillate, glabrous,
globose, 175-240 um diameter, 175-240 um
high. Ascocarp wall of textura angularis-
globulosa in surface view; in longitudinal
section 22—31 tm thick at sides and base, up to
50 um thick near apex, composed of 6—8 layers
of polygonal, isodiametric-elongate, pseudo-
parenchymatic cells [3.6—7.2 x 5.4-12.6(—16)
uum], outer 24 layers of brown-melanized cells
at sides and base, inner 4—5 layers of hyaline-
compressed cells at base. Papilla very short,
erumpent, rounded-conical, 31—45 um high,
62-93 tum wide, wall 25-30 um thick, com-
posed of 8—10 layers of small, isodiametric,
scleroplectenchymatic cells (2.0—7.2 ium
diameter), outer 5—6 layers brown-melanized,
inner 34 layers hyaline, surrounding a 30-40
lim wide circular ostiole without periphyses.
Pseudoparaphyses |.0—2.0 tum wide, 125-200
tum long (height of the ascocarp cavity),
numerous, narrowly cellular, with gelatinous
coating. Asci 95—120 x 17-21 um, numerous,
basal, cylindrical-oblong, thick-walled, short-
stalked, rounded apex, with 8 biseriate asco-
spores. Ascospores (25—)27—31(—33) x 8.5—
11.5 um, broadly fusiform to slightly clavate,
end cells rounded, straight to slightly curved,
3-septate, septa evenly distributed, order of
septation 2:1:2, primary septum median, slight
constrictions at all septa, brown, ornamented
walls, without sheath or appendages.

Holotype: UNirep States: In foliis emortuis
Cercocarpi ledifolii, Mill Creek Canyon, Salt
Lake Co., Utah Americae bor. leg. A.O. Garrett
no. 677 (S).

Other material examined: Unirep States:
Utan: Box Elder Co., One Mile Creek, north
side of Raft River Mts., Aug. 25, 1986; Juab

Vol. 34 Art. 5

Co., 5.2 mi up Granite Creek Canyon, east side
of Deep Creek Mts, Aug. 26, 1986; Rich Co.,
Sunrise Campground, Cache National Forest,
Route 89, west of Garden City, July 13, 1985;
Weber Co., Malans Peak, Aug. 11, 1972:
Weber Co., south slope of Malans Peak,
Wasatch Mts., east of Ogden, July 1, 1976;
Weber Co., mouth of Taylor’s Canyon,
Wasatch Mts., east of Ogden, May 18, 1972,
Apr. 29, 1981, May 6, 1982, all on
Cercocarpus ledifolius Torrey in Torrey &
Gray var. intermontanus (Brittonia 39:424.
1987), all C.T. Rogerson (all NY).

Comments: Leptosphaeria cercocarpi is an
easily recognized species on Cercocarpus
ledifolius. The ascocarps usually entirely cover
both surfaces of dead leaves. Retaining this
species in Leptosphaeria is not totally satisfac-
tory, but placing it elsewhere is also problem-
atic. Certain characteristics resemble those
found in the Phaeosphaeriaceae (sensu Barr
1987a). The ascocarps are small-medium sized
and immersed in the substrate, and there
appears to be some kind of hyphal growth or
stroma surrounding the ascocarps and extend-
ing into the substrate. Other characteristics do
not coincide with inclusion in that group.
Although the lateral walls of the ascocarp of L.
cercocarpi consist of pseudoparenchymatous
cells, they are not thin and soft as is character-
istic of the Phaeosphaeriaceae. In fact, the walls
in the upper regions of the ascocarp consist of
thick-walled scleroplectenchymatous cells. In
Barr’s (1987a) key to this family, the only two
genera in which this fungus could be placed are
Phaeosphaeria and Kalmusia. The ascocarps
are not small and delicate enough for inclusion
within Phaeosphaeria, and the current concept
includes only species on monocots (Shoemaker
and Babcock 1989b) (but see also the discus-
sions for P. pomona and P. lucilla herein). The
current concept of Ka/musia, which includes K.
clivensis (see discussion herein), does not
adequately accommodate L. cercocarpi.
Therefore, unlike other leaf-inhabiting species
of Leptosphaeria belonging elsewhere (i.e.,
Phaeosphaeriaceae), this species is retained
within Leptosphaeria because of the larger,
robust nature of the ascocarp and wall.

May 1992 Leptosphaeria Species on the Rosaceae 49]

U

Figure 5. Leptosphaeria cercocarpi. a. SEM of immersed ascocarps, x50. b. ascocarp wall surface, x550.
c. longitudinal median section through ascocarp, x225. d. section through ascocarp wall neck region, x950.
e. longitudinal median section through ascocarp wall, x950. f. ascus, x525. g, h. cellular pseudoparaphyses,
x2,000. 1. ascospore, x1,150. All from holotype of L. cercocarpi.

Illinois Natural History Survey Bulletin

Phaeosphaeriaceae

The species considered in this section all
possess characteristics of the Phaeo-
sphaeriaceae (Barr 1987a). Species in the
Phaeosphaeriaceae are distinguished from those
in the Leptosphaeriaceae by the smaller
ascomatal size, the thin, soft ascocarp wall
composed of pseudoparenchymatous cells, and
the rather sparse pseudoparaphyses (Barr
1987a). The genera represented within the
family are Paraphaeosphaeria Eriksson,
Phaeosphaeria Miyake, and Kalmusia Niessl.

*Paraphaeosphaeria concentrica (J.B. Ellis &
B.M. Everhart) S.M. Huhndorf comb. nov. See
Figure. 6.
= Leptosphaeria concentrica J.B. Ellis &
B.M. Everhart, The North American
Pyrenomycetes. A Contribution to
Mycologic Botany, p. 354. 1892.
(Basionym).

Ascocarps scattered, sparse, immersed-
subepidermal, globose, papillate, glabrous, 90—
100 um diameter, 100-125 um high. Ascocarp
wall of textura angularis in surface view; in
longitudinal section 7.2—10.8 [1m thick at sides
and base, composed of 2-3 layers of elongate-
compressed, hyaline, pseudoparenchymatic
cells (2.8-3.6 x 12—14.5 um), wall up to 13 um
thick near apex, composed of 34 layers of
isodiametric-elongate, slightly brown-
melanized cells (3.6—5.6 x 7.2—12.2 um).
Papilla very short, erumpent, bluntly conical,
15-20 um high, 25-30 um wide, 10-13 Lum
thick near base of papilla, papilla apex wall
cells hyaline, thin-walled surrounding a circular
ostiole without periphyses. Pseudoparaphyses
1.4—2.0 um wide, 70-90 Lim long (height of the
ascocarp cavity), numerous, narrowly cellular,
without gelatinous coating. Asci (43—)54—66 x
10.8—12.2 um, numerous, basal, cylindrical-
clavate, thick-walled, short-stalked, rounded
apex, with apical chamber, with 8 biseriate
ascospores. Ascospores (14.5—)16.5—18.7 x
(3.6—) 4.3-5.7(-6.5) um, cylindrical, with
rounded end cells, end cells longer than central
cell, straight; 2-septate, septa unevenly distrib-
uted, order of septation 2:1, primary septum
submedian and constricted; brown, wall
roughened, thin cellular sheath entirely
surrounding spore, without appendages.

Vol. 34 Art. 5

Holotype: Unitep States: On apple leaves
(Malus sp.), Columbia, Missouri (H. Dorsett),
and Louisiana (Langlois) (NY).

Comments: Paraphaeosphaeria species are
distinguished by ascocarps that form below or
within the host epidermis, an ascocarp wall
consisting of a few layers of pseudoparen-
chyma, and cylindrical, usually echinulate
ascospores with the primary septum forming in
the lower half of the spore.

Although Paraphaeosphaeria species are
commonly found on monocotyledonous plants,
species on dicot plants have also been placed in
the genus (Hedjaroude 1969, Shoemaker and
Babcock 1985). Such is the case here with
Paraphaeosphaeria concentrica described from
apple leaves. The description of P. concentrica
closely resembles the description of the type of
the genus, P. michotii (Westendorp) Eriksson,
with the only difference being the dicotyledon-
ous host plant. Paraphaeosphaeria michotii
and its synonyms have so far been found
exclusively on monocot hosts (Shoemaker and
Eriksson 1967, Shoemaker and Babcock 1985).
Overlooking substratum preference, P.
concentrica would be synonymous with P.
michotii. Because of the historical value placed
on this character and the need for experimental
evidence to demonstrate the range of substrata
on which species will grow and the total lack of
collections of P. michotii on dicot hosts, P.
concentrica is retained as a separate species.

*Phaeosphaeria pomona (P.A. Saccardo) S.M.
Huhndorf comb. nov. See Figure 7.
= Leptosphaeria (Leptosphaerella)
pomona P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della
Societa Botanica Italiana 8:176. 1876.
(Basionym).

Ascocarps scattered, sparse, immersed-
subcuticular, globose, papillate, glabrous to
slightly tomentose near base, with loose, dark
hyphal growth on host cuticle surrounding
ascocarps, (55—)90—125 tm diameter, 75—115
tim high. Ascocarp wall of textura angularis-
globulosa in surface view; in longitudinal
section uniformly 6.1—7.7 um thick, composed
of 2—3 layers of brown-melanized, compressed,
isodiametric-to-elongate pseudoparenchyma-
tous cells (1.5—2.0 x 8-12 um). Papilla very
short, erumpent, bluntly conical, 15—21 uum

May 1992 Leptosphaeria Species on the Rosaceae 493

Figure 6. Paraphaeosphaeria concentrica. a. longitudinal median section through ascocarp, x375. b. cellular
pseudoparaphyses, x2,000. c. longitudinal median section through ascocarp wall, x950. d, e. asci, x1,050.
f, g. ascospores, x1,150. All from holotype of P. concentrica.

high, 31-37 um wide, composed of 34 layers ostiole without periphyses. Pseudoparaphyses
of small, brown-melanized, isodiametric cells 1.0—-1.5 um wide, (46—)77—93 [um long (height
(2.64.6 um diameter), (9—)12—17 um thick of the ascocarp cavity), numerous, narrowly
near base of papilla, apex of papilla wall cells cellular, with guttule-like thickenings at septa,
hyaline, thin-walled surrounding a circular without gelatinous coating. Asci 55.5—-65 x

Illinois Natural History Survey Bulletin

a i as
Figure 7. Phaeosphaeria pomona. a. longitudinal median section through ascocarp, x375. b. ascocarp wall

surface, x550. c. longitudinal median section through ascocarp wall, x950. d. ascus, x1,050. e. ascospore,
x1,150. All from holotype of P. pomona.

8.3-11.5 um, numerous, basal, cylindrical,
thick-walled, short-stalked, rounded apex, with
apical chamber, with 8 biseriate ascospores.
Ascospores 20.1—26.5 x (2.8—)3.6—5.0 Lum,
narrowly f._.form, with acute end cells, second
cell from the top occasionally enlarged,
straight; 5-septate, septa unevenly distributed,
without constrictions, order of septation
unknown, pale brownish yellow, guttules
present in ascospore cells, smooth, thin cellular
sheath entirely surrounding spore (0.7—2.0 um
thick).

Vol. 34 Art. 5

Holotype: Iracy: In pag. super. folior. Pyri
Mali, socia Vermicularia Pomona, a selva
(Treviso), Sept. 1875, raro (PAD).

Comments: See under Phaeosphaeria lucilla.

¢Phaeosphaeria lucilla (P.A. Saccardo) S.M.
Huhndorf comb. nov. See Figure 8.
= Leptosphaeria lucilla P.A. Saccardo,
Nuovo Giornale Botanico Italiano e
Bolletino della Societa Botanica Italiana
7:310-311. 1875. (Basionym).

May 1992 Leptosphaeria Species on the Rosaceae 495

Figure 8. Phaeosphaeria lucilla. a. longitudinal median section through ascocarp, x375. b. cellular
pseudoparaphyses, x2,000. c. longitudinal median section through ascocarp wall, x950. d. asci, x1,050.
e. ascocarp wall surface, x550. f, g. ascospores, x1,150. All from holotype of P. lucilla.

Ascocarps scattered, sparse, immersed- brown-melanized, compressed, isodiametric-to-
subcuticular, papillate, glabrous, depressed- elongate, pseudoparenchymatous cells (2.3—3.8
globose, 110-130 um diameter, 90-115 um x 6.9-10.0 um). Papilla very short, erumpent,
high. Ascocarp wall of textura angularis in conical, 20-25 um high, 30-40 um wide,

surface view; in longitudinal section uniformly | composed of 4—S layers of brown-melanized,
6.0—7.7 um thick, composed of 34 layers of isodiametric cells (3.0-4.6 ium diameter), 10.7—

Illinois Natural History Survey Bulletin

0 tim thick near base of papilla, apex of
papilla wall cells hyaline, thin-walled surround-
ing a circular ostiole without periphyses.
Pseudoparaphyses 1.5—2.0 um wide, 62-71
um long (height of the ascocarp cavity),
numerous, narrowly cellular, without gelati-
nous coating. Asci 40—51(—56) x 8.4-10(-11.5)
im, numerous, basal, in a broad hymenium,
cylindrical, thin-walled, sessile, rounded apex,
with 8 bi- to triseriate ascospores. Ascospores
15.3-18.4(—20) x 3.0-4.6 um, fusiform, with
acute end cells, second cell from apex slightly
enlarged, straight to slightly curved; 3-septate,
septa evenly distributed, order of septation
2:1:2, primary septum median and slightly
constricted; pale yellowish brown, wall smooth,
without sheath or appendages.

Holotype: Iraty: In foliis languidis Pyri
communis in agro Tarvisino et Patavino
(PAD).

Comments: Fungi in the genus Phaeosphaeria
are characterized by immersed ascocarps with
thin walls of pseudoparenchyma, fusiform 3- to
multiseptate ascospores and monocotyledonous
hosts. Nonetheless, Leuchtmann’s (1984)
treatment of Phaeosphaeria includes a group of
species found on the leaves and stems of plants
in the dicot family Caryophyllaceae. The
placement of dicotyledonous Leptosphaeria
species, morphologically similar to
Phaeosphaeria, into the genus Phaeosphaeria
is contrary to Shoemaker and Babcock’s
(1989b) concept of the genus. These authors
do not, however, suggest an alternative genus
for these species. They do place certain species
on dicots in Leptosphaeria, citing the presence
of thick-walled cells in the ascocarp wall.
Phaeosphaeria pomona and P. lucilla are
morphologically similar to species of
Phaeosphaeria found on monocot hosts. They
do not have the ascocarp and wall characteris-
tics of Leptosphaeria.

When P. pomona and P. lucilla were
described, Saccardo (1875, 1876) placed them
in Leptosphaeria subgenus Leptosphaerella
Sacc., which included fungi from leaves of
dicotyledonous plants. Leptosphaeria subgenus
Leptosphaerella was subsequently placed in
synonymy with Phaeosphaeria (Leuchtmann
1984). At this time, there is no other genus
suitable for the placement of these “dicot

Vol. 34 Art. 5

Phaeosphaeria” species. Erecting a new genus
for these species based solely on substrate
preference, with no morphological differences
from Phaeosphaeria seems unwise. With proof
of strict substrate preference requiring experi-
mental evidence, there seems to be some
justification for their placement within
Phaeosphaeria until such work is done.

¢Phaeosphaeria thomasiana (P.A. Saccardo &
C. Roumeguére) S.M. Huhndorf comb. nov.
See Figure 9.
= Leptosphaeria thomasiana P.A.
Saccardo & C. Roumeguére, Revue
Mycologique 5:236. 1883. (Basionym).

Ascocarps scattered, numerous, immersed-
subcuticular, globose, flattened at base,
papillate, glabrous, 150-175 um diameter,
110-125 um high. Ascocarp wall of textura
angularis-globulosa in surface view; in longitu-
dinal section uniformly 9.3—12.2 um thick,
composed of 4—5 layers of polygonal, pseudo-
parenchymatous cells, outer 2—3 layers of
polygonal-to-elongate, brown-melanized cells
(2.2-5.0 x 6.5—8.6 um), inner 1—2 layers of
elongate-compressed, hyaline cells (0.7—1.5 x
8.0-10.8 um). Papilla very short, erumpent,
bluntly conical, 12-30 um high, 10-36 um
wide, composed of 7—8 layers of small, brown-
melanized, isodiametric cells (2.64.6 um
diameter), 14-18 um thick near base of papilla,
apex of papilla wall cells hyaline, thin-walled
surrounding a 12—18 um wide circular ostiole
without periphyses. Pseudoparaphyses 1.0—
1.5 um wide, 80-85 um long (height of the
ascocarp cavity), numerous, narrowly cellular,
without gelatinous coating. Asci 55.5—65.5 x
8.0-10.8 um, numerous, basal, cylindrical-
clavate, thick-walled, short-stalked, rounded
apex, with apical chamber, with 8 biseriate
ascospores. Ascospores 15.0—18 x 3.64.5 um,
fusiform, with acute end cells, straight or
slightly curved; 3-septate, septa evenly distrib-
uted, order of septation 2:1:2, primary septum
median, without constrictions; subhyaline to
pale brownish yellow, guttules absent in
ascospore cells, smooth, without sheath or
appendages.

Holotype: France: In sarmentis Rubi emortuis
(305-Reliquiae Libertianae) n.v.

May 1992 Leptosphaeria Species on the Rosaceae 497

Figure 9. Phaeosphaeria thomasiana. a. longitudinal median section through ascocarp, x375. b. cellular
pseudoparaphyses, x2,000. c. longitudinal median section through ascocarp wall, x950. d. asci, x1,050.
e. ascocarp wall surface, x550. f, g. ascospores, x1,150. All from Roum. F. sel exs. 6039.

Illinois Natural History Survey Bulletin

scatae: FRANCE: Bois des Roches (Noidan),
May 1891, F. Fautrey, Roumeguere, F. sel exs.
6039 (NY).

Other material examined: UNiTED STATES:
OrEGON: Corvallis, on loganberry (Rubus
loganobaccus Bailey), Mar. 12, 1930, S.M.
Zeller; on loganberry, Mar. 15, 1916, A. Frank
(all NY).

Comments: Leptosphaeria thomasiana 1s
placed in Phaeosphaeria because of its small
ascocarps immersed beneath the host cuticle
and its ascocarp wall of pseudoparenchymatous
cells. I was unable to see the type specimen,
which is number 305 in the exsiccatae set
Reliquiae Libertianae. This is not a regular
exsiccatae set (Pfister 1985), and the set at FH
did not contain this specimen. The description
and plate herein were prepared from
Roumeguere, F. sel exs. 6039 specimen from
NY, which included few ascocarps. I did not
find any ascocarps of P. thomasiana in the
collections from Oregon, but the canes did have
large gray patches on them as Zeller (1927)
described. He also mentions that a cane blight
or “Loganberry gray bark disease” has been
ascribed to this organism in western Washing-
ton. I have seen no reports from other regions
of this organism as a pathogen of Rubus.

*Kalmusia clivensis (M.J. Berkeley & C.E.
Broome) M.E. Barr, Mycotaxon. An Interna-
tional Journal Designed to Expedite Publication
of Research on Taxonomy & Nomenclature of
Fungi & Lichens 29:504. 1987. See Figure 10.
= Sphaeria (Caulicolae) clivensis M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, Series 2,
9:379. 1852.
= Leptosphaeria clivensis (M.J. Berkeley
& C.E. Broome) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:16.
1883.
= Diapleella clivensis (M.J. Berkeley &
C.E. Broome) A. Munk, Dansk Botanisk
Arkiv 15(2):75. 1953.
= Leptosphaeria galiorum P.A. Saccardo
var. lapsanae P.A. Saccardo & P.A. Briard,
Revue Mycologique 7:209. 1885.
= Leptosphaeria steironematis J.B. Ellis &
B.M. Everhart, Proceedings of the Academy
of Natural Sciences of Philadelphia 1890:
237. (1890) 1891.

Vol. 34 Art. 5

= Leptosphaeria arunci S.M. Zeller,
Mycologia 19:134—135. 1927.

= Leptosphaeria longipedicellata J.H. Miller
& G. Burton, Mycologia 34:2-3. 1942.

Ascocarps scattered, sparse, immersed-
subepidermal, papilla erumpent, at times with a
surrounding clypeus, glabrous, depressed-
globose, 275-400 um diameter, 175-375 um
high. Ascocarp wall of textura prismatica in
surface view; in longitudinal section uniformly
15-20 um thick, composed of 5-6 layers of
parallel, elongate, prismatic, scleroplecten-
chymatic cells (0.7—1.5 x 15—20 um), outer 34
cell layers slightly brown-melanized, inner 2—3
layers hyaline; at the base cells are compressed,
flattened, hyaline. Papilla conical, short-
intermediate, (50—)90—100(—130) um high, 40—
80 um wide at the apex, 50-100 um wide at the
base, composed of 6-8 layers of small, light-
brown pigmented, isodiametric cells with no
external, melanized crust (2—5 im diameter),
10-13 um thick, surrounding a circular ostiole
18—20 um wide, composed of thin-walled,
hyaline, compressed cells, without periphyses.
Pseudoparaphyses 0.5—1.5 um wide, 200-225
um long (height of the ascocarp cavity),
numerous, narrowly cellular, without gelati-
nous coating. Asci 95—110(—150) x 11—15(-21)
iim, numerous, basal, clavate, thin-walled,
long-stalked (27-39 um long), rounded apex,
with 8 biseriate ascospores. Ascospores 19—
25(—32) x 5-8.5(-11) um, fusiform, with
rounded to acute end cells slightly longer than
central cells, straight to slightly curved; 3-
septate, septa slightly unevenly distributed,
order of septation 2:1:2, with slight constric-
tions at all septa; dark brown, smooth, without
sheath or appendages.

Holotype: Great Britain: King’s Cliffe, on
dead stems of Pastinaca sativa, Jul 1850, Herb.
Berk. 1879 (K).

Exsiccatae: CANADA: London, on Steironema
ciliatum, as Leptosphaeria steironematis, May
1890, Ell. & Ev., North American Fungi 2615
(NY); Sphaer. Brit. II] 60, (FH).

Other material examined: Canapa: London, on
Steironema ciliatum, May 1890, with 1640,
Dearness (Holotype of Leptosphaeria
steironematis, NY); 1640, London, as
Leptosphaeria steironematis, 19 Apr., 1890

May 1992 Leptosphaeria Species on the Rosaceae 499

Figure 10. Kalmusia clivensis. a. SEM of immersed ascocarps with erumpent papilla, x100. b. ascocarp wall
surface, x550. c. longitudinal median section through ascocarp, x125. d. cellular pseudoparaphyses, x2,000.
e. longitudinal median section through ascocarp wall, x950. f, g. asci, x525. h-j. ascospores, x1,150. a, c, d,
e, f, and h from holotype of Leptosphaeria arunci; b, g, and i from holotype of K. clivensis; j from lectotype
of L. longipedicellata.

Illinois Natural History Survey Bulletin

JY). Great Brirain: England, Chute Meadow,
Lambriggan, W. Cornwall, on Centaurea nigra,
May 9, 1942, F. Rilstone (NY-MEBB coll);
Scotland, Wester Ross, Rassal N.N. Reserve,
on ? Senecio jacoboea, June 2, 1982, P.F.
Cannon (NY). ITAty: Briard no. 8, on Lapsana
communis, June 12, 1885 (holotype of
Leptosphaeria galiorum var. lapsanae, PAD).
Unirep States: CoLorapo: Larimer Co., 3 mi
W. Redfeather Lakes, 8100' Roosevelt National
Forest, on Rosa sp., Aug. 4, 1984, MEBB #
7008 (NY); GeoraiA: 7684, Clarke Co., Athens,
South Campus, University of Georgia, on
Daucus carota, Sept. 11, 1939, J.H. Miller
(Lectotype of Leptosphaeria longipedicellata,
GAM); MIicuiGAN: Emmet Co., Gill and Elder
Roads, Carp Lake, on ? Acer, Sept. 11, 1969,
MEBB #5527 (NY); OreEGON: Multnomah Falls,
April, on dead stems of Aruncus silvester
(holotype of Leptosphaeria arunci in Zeller
Herb. 6811, NY).

Comments: Four of the putative Leptosphaeria
species that were examined are synonymous
with Kalmusia clivensis; one was found on a
Rosaceous host (L. arunci on Aruncus). The
most distinctive feature of this species is the
presence of long-stipitate asci with prominent
dark brown, 3-septate ascospores. The long
stipe, as well as the thin ascus wall, has led
previous workers to treat it as a unitunicate
fungus in the monotypic genus Diapleella
(Munk 1957, Dennis 1978). Shoemaker
(1984a) retained the genus Diapleella but
treated it as bitunicate. Barr (1987b) trans-
ferred D. clivensis to the genus Kalmusia Niessl
without much explanation. The original
description of Kalmusia (Niessl 1871) and the
illustration of the genus in Berlese (1890) show
a fungus with long-stipitate asci and dark
brown, 3-septate ascospores. Kalmusia
clivensis appears to be well-placed within this
genus.

Shoemaker (1984a) looked at one
collection of Leptosphaeria longipedicellata on
Solidago caesia L., which is cited in the
original description of the species. He men-
tions that “the original description of L.
longipedicellata is strongly suggestive of
Diapleella clivensis.” This collection was in
fact L. macrospora (Fuckel) Thiimen, and
Shoemaker did not resolve the placement of L.
longipedicellata. The collection on Daucus
carota L., cited in the description and marked

Vol. 34 Art. 5

as type on the herbarium packet, matches the
original description exactly and is the same as
Kalmusia clivensis except that the ascospores
and asci are slightly larger than those in the
type collection of K. clivensis. The ascocarp
and wall in section appear exactly the same. I
believe the size differences are probably due to
environmental differences or other individual
variation and that L. longipedicellata is
synonymous with K. clivensis. Collection
7684, Clarke Co., Athens, South Campus,
University of Georgia, on Daucus carota, Sept.
11, 1939, J.H. Miller (GAM) is chosen as the
lectotype of Leptosphaeria longipedicellata.

¢Kalmusia coniothyrium (L. Fuckel) S.M.
Huhndorf comb. nov. See Figure 11.

= Sphaeria coniothyrium L. Fuckel,
Symbolae Mycologicae, p. 115. 1870.
(Basionym).
= Leptosphaeria coniothyrium (L.
Fuckel) P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della
Societa Botanica Italiana 7:317. 1875.
= Melanomma coniothyrium (L. Fuckel)
L. Holm, Symbolae Botanicae
Upsalienses 14(3):56—-S7, 1957.
= Diapleella coniothyrium (L. Fuckel)
MLE. Barr in M.E. Barr, C.T. Rogerson,
S.J. Smith, and J.H. Haines, Bulletin of
the New York State Museum 459:30.
1986.

= Sphaeria (Obtectae) hendersonia J.B.

Ellis in M.C. Cooke and J.B. Ellis, Grevillea

6:14-15. 1877.
= Clypeosphaeria hendersoniae (J.B.
Ellis) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:91. 1883.
= Leptosphaeria (Clypeosphaeria)
hendersoniae (J.B. Ellis) M.C. Cooke,
Grevillea 17:91. 1889.

Ascocarps clustered, numerous, immersed-
subepidermal, depressed, globose, flattened at
top and base, sometimes beneath blackened
clypei, papillate, glabrous, 175-300 um
diameter, 175-200 um high. Ascocarp wall of
textura angularis-globulosa in surface view; in
longitudinal section 15—25 um thick at the
sides, 13-18 um at the base, composed of 8-12
layers of polygonal, pseudoparenchymatous
cells, outer 3—5 layers of isodiametric-to-
slightly-elongate, light brown cells (5.0-9.5 x

May 1992 Leptosphaeria Species on the Rosaceae 501

> \h i j g
Figure 11. Kalmusia coniothyrium. a. longitudinal median section through ascocarp, x250. b. ascocarp wall
surface, x550. c. longitudinal median section through ascocarp wall, x950. d, e. asci, x1,050. f, g. cellular
pseudoparaphyses, x2,000. h-j. ascospores, x1,150. a—d, f-i from holotype of K. coniothyrium; e, j from
neotype of Sphaeria hendersonia.

Illinois Natural History Survey Bulletin

3.6-5.0 ym), inner 5-7 layers of small,
isodiametric-elongate, compressed, hyaline
cells (3.64.3 x 2.0-3.6 um). Papilla short,
erumpent, bluntly conical, 45—55 um high, 25—
35 tum wide, 16-18 tm thick, composed of 5—
10 layers of small, hyaline, isodiametric cells
(1.4-2.8 um diameter), surrounding a 10-20
um wide, circular ostiole without periphyses.
Pseudoparaphyses 1.0—1.5 um wide, 75—100
um long (height of the ascocarp cavity),
numerous, narrowly cellular, with guttule-like
thickenings at septa, with gelatinous coating.
Asci 60-75 x 5.5—7.5 um, numerous, basal in a
broad hymenium, cylindrical, thin-walled,
short-stalked, rounded apex, with apical
chamber, with 8 overlapping, uniseriate
ascospores. Ascospores | 1.5—14.4(—15.8) x
3.64.5 um, fusiform to ellipsoidal, with acute
end cells, second cell somewhat enlarged,
straight or slightly curved; 3-septate, septa
slightly unevenly distributed, order of septation
2:1:2, primary septum median and constricted;
brownish yellow, guttules lacking, smooth,
without sheath or appendages.

Holotype: Austria: Auf durren Ranken von
Rubus fruticosus, selten, im Fruhling. An der
Heimbach bei Oestrich (G).

Exsiccatae: Rehm, Asc. 388, on Rubus
fruticosus (NY); Krieger, Fungi saxon. 18,
1120, 1121 (NY); Petrak Kryptogamae exsic.
2318 (NY); Sacc, Mycotheca Veneta 72, as
Sphaeria fuscella f. Ampelopsidis hederaceae,
(FH); Petrak, Fl. Boh. et Mor. exsic. II no. 4, as
Clypeosphaeria notarisii (FH); Ellis N.
American Fungi 581 as Sphaeria Hendersonia,
Mar. 1878 (ILL, FH, NY).

Other material examined: CANApDa: Ottawa, on
Sambucus racemosa, Mar. 10, 1897 (NY);

London, as Clypeosphaeria Hendersonia, 1883,

3 May 1892 (NY). ITALy: 198 as Sphaeria
clypeata, de Notaris (RO). Unirep States:
DeLAware: Faulkland, as Sphaeria

Hendersonia, Mar. 20, 1887 (NY); New JERSEY:

Newfield, on Rubus strigosus, J.B. Ellis 101,
May 29, 1880 (neotype of Sphaeria
Hendersonia, NY); on Rubus occidentalis, as
Sphaeria Hendersonia, Aug. 4 1879, Apr.
1880, J.B. Ellis (FH); as Clypeosphaeria
Hendersonia, Aug. 15, 1894, J.B. Ellis 773

Vol. 34 Art. 5

(NY); Massacuusetts: Andover, as Sphaeria
Hendersonia, Rev. J. Blake No. 79 (NY);
NortH Dakota: Kulm, on Rubus strigosus,
Mar. 12, 1916, Brenkle, Fungi Dakotenses 384
(NY); OreGon: Troutdale, Feb. 10, 1929, M.J.
O’Connell (NY); Gresham, Apr. 30, 1934,
S.M. Zeller (NY); PENNSYLVANIA: Avestrud Co.,
on Rubus sp., July 10, 1944 (NY); Wisconsin:
Sauk Co., Aldo Leopold Reserve, on Rubus sp.,
9 Apr. 1988, 30 Sept. 1988, S.M. Huhndorf
(ILLS).

Comments: This fungus is placed in Kalmusia
because of its immersed, clypeate ascocarps
with pseudoparenchymatous walls composed of
compressed cells. Kalmusia coniothyrium
lacks the characters distinctive of
Leptosphaeria, including erumpent to superfi-
cial ascocarps with a wall of scleroplec-
tenchymatous cells. Holm (1957) placed the
species in Melanomma, but the fungus does not
fit the current concept of that genus (Barr
1987a) because it lacks erumpent ascocarps
with walls composed of small thick-walled
cells, asci formed peripherally within the
centrum, and trabeculate pseudoparaphyses.
However, the ascospores of this fungus do
strongly resemble those of some Melanomma
species. The ascus shape of K. coniothyrium
differs from that of K. clivensis in being
cylindrical and short-stalked, and the asco-
spores are brownish yellow rather than dark
reddish brown. But in Kalmusia ebuli Niessl,
the type of the genus, these characters are
variable; asci are clavate to cylindrical, short-
or long-stalked, and ascospores are lighter
brown than those of K. clivensis. There
appears to be sufficient range within the genus
to accommodate K. coniothyrium.

Sphaeria hendersonia is synonymous
with K. coniothyrium. The description for the
holotype specimen is at NY, but the actual
specimen is missing. Because no other
specimen is cited in the published description
and it is uncertain what was available to Ellis
when he described the organism, a neotype was
chosen from the Ellis collection at NY which
matches the description and is in good condi-
tion. The neotype of Sphaeria hendersonia is
J.B. Ellis 101, Newfield, New Jersey, on Rubus
strigosus, May 29, 1880.

May 1992

Lophiostomataceae

The following species possesses characteristics
of the Lophiostomataceae in the order
Pleosporales (Barr 1987a). Lophiostomataceae
sensu Holm and Holm (1988) is considered a
heterogeneous group whose overall common
characters, the flattened neck and slotlike
ostiole, are highly adaptive and unstable.
Consequently, the family in this sense may be
completely dissociated (Holm and Holm 1988).
Lophiostomataceae sensu Barr (1987a) contains
genera that are united by wall characteristics,
pseudoparaphysis structure and ascospore
morphology. The compressed apical papilla is
considered to be of generic or specific impor-
tance. Taxa in the Lophiostomataceae may
have this type of papilla or may have a con-
spicuous, rounded apical papilla, or a short
papilla with or without grouped setae, or the
apex may open by a pore or slit.

*Lophiostoma subcutanea (M.C. Cooke & J.B.
Ellis) S.M. Huhndorf comb. nov. See Figure
12:
= Sphaeria (subtectae) subcutanea M.C.
Cooke & J.B. Ellis, Grevillea 7:41. 1878.
(Basionym).
= Leptosphaeria subcutanea (M.C.
Cooke & J.B. Ellis) J.B. Ellis in N.L.
Britton, Catalogue of Plants Found in
New Jersey. Geological Survey of New
Jersey, Final Report of the State Geolo-
gist, 2(1):525. 1889.
= Metasphaeria subcutanea (M.C. Cooke
& J.B. Ellis) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo
2:167. 1883.

Ascocarps clustered, sparse, superficial,
papillate, glabrous, conic, 150-170 um
diameter, 180-225 um high. Ascocarp wall of
textura angularis-prismatica in surface view; in
longitudinal section (15—)20—23 um thick at
sides and base, composed of 6—7 layers of
polygonal, isodiametric-to-elongate,
pseudoparenchymatic cells (2.0—3.6 x 5—7 um),
outer 3-4 layers of brown-melanized cells at
the sides, inner 1—3 layers of hyaline, com-
pressed cells; basal wall composed only of
hyaline, elongate-compressed cells. Papilla
elongate-conical, longitudinally compressed,

Leptosphaeria Species on the Rosaceae 503

75-85 um high, 55—75 um wide at the apex,
100-125 um wide at the base, composed of 6-7
layers of brown-melanized, thickened, isodia-
metric cells (2.5—4.0 um diameter); at apex,
cells coalescing to form a thickened dark brown
mass; wall 25—27 um thick, surrounding a 40—
50 um wide slotlike ostiole lined with
periphyses. Pseudoparaphyses 0.5—1.5 um
wide, 108—124 im long (height of the ascocarp
cavity), numerous, narrowly cellular, without
gelatinous coating. Asci 77—84(—92) x 5.3-7.6
lim, numerous, basal, cylindrical, thin-walled,
short-stalked, rounded apex, with 8 biseriate
ascospores. Ascospores (20—)25—29 x 3.0-3.5
lim, narrowly fusiform, with acuminate end
cells, second cell slightly enlarged, straight to
slightly curved; 4-6(-8)-septate, septa unevenly
distributed, order of septation unknown, with
slight constrictions at all septa; hyaline to
subhyaline, guttules present in ascospore cells,
wall smooth, possibly with thin cellular sheath
surrounding entire spore, without appendages.

Holotype: Unitep States: On decorticated
limbs of Pyrus communis L (NY, K).

Comments: Holm and Holm (1988) considered
the Swedish species of the Lophiostomataceae
and emphasized ascocarp shape and peridial
anatomy in distinguishing the genera
Lophiostoma Ces. & de Not. and Lophiotrema
Sacc. emend. L. Holm & K. Holm. The
presence of a flattened papilla or neck and a
slotlike ostiole have been used to distinguish
the Lophiostomataceae from other groups.
Although these features are highly variable in
some species, they are still useful for placing
this species in the Lophiostomataceae. How-
ever, generic placement within the family is
problematic. The conic or pyriform ascocarp
shape with a distinct flattened neck and the
fusiform ascospores that are multiseptate within
the ascus suggest Lophiostoma. The cylindric
asci suggest Lophiotrema. The ascocarp wall
anatomy in section resembles that of
Lophiotrema, with uniformly thick walls of
polygonal globose or angular cells. But in
surface view, the cells seem to form a textura
prismatica suggesting the long, parallel cells of
Lophiostoma. The fungus is placed in
Lophiostoma with some reservation because, at
this time, there is no other suitable place for it.

Illinois Natural History Survey Bulletin Vol. 34 Art. 5

>»

aS cae

. is
~~ w/ et f 3 “nh
= * zz + .
Figure 12. Lophiostoma subcutanea. a. SEM of ascocarp with apical crest, x175. b. longitudinal section of
irp neck with periphyses, x525. c. longitudinal median section through ascocarp, x325. d. ascocarp
550. e. longitudinal median section through ascocarp wall, x950. f. ascus, x525. g. ascospores
is, x1,150. h. cellular pseudoparaphyses, x2,000. All from holotype of L. subcutanea.

Leptosphaeria Species Referable to the Melanommatales

One species investigated had characteristics of
the Melanommataceae in the order Melanom-
matales (Barr 1987a). The order
Melanommatales is distinguished from the
Pleosporales by trabeculate pseudoparaphyses,
asci arranged peripherally within the centrum,
an ascocarp wall composed of small or com-
pressed cells, and ascospores with bipolar
symmetry. Any one of these features may
deviate in a particular taxon. Fungi in the
Melanommataceae are distinguished by
gregarious, erumpent ascocarps with a wall of
small, thick-walled cells.

*Melanomma pulvis-pyrius (C.H. Persoon:E.M.

Fries) L. Fuckel, Symbolae Mycologicae p.
159, 1870. See Figure 13.
= Sphaeria pulvis-pyrius C.H. Persoon,
Synopsis Methodica Fungorum p. 86,
1801; E.M. Fries, Systema Mycologicum
Sistens Fungorum 2:458. 1823.
= Cladosphaeria rimicola G.H. Otth,
Mitteilungen der Naturforschenden
Gesellschaft in Bern 1870:106. 1871. Nom
inval. Art. 43.1. [The genus Cladosphaeria
was validated in 1894.]
= Leptosphaeria rimicola (G.H. Otth)
P.A. Saccardo, Hedwigia, Dresden
35:XXIX. 1896; Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 11:XXIX. 1896.

Ascocarps clustered, sparse, superficial,
papillate, glabrous with tomentum of thick-
walled brown hyphae surrounding and between
ascocarps, conic-subglobose to irregular, 425—
550 um diameter, 475-600 um high. Asco-
carp wall of textura angularis-epidermoidea in
surface view; in longitudinal section 55-75 um
thick at sides, 80-100 tm thick at base,
composed of 21—35 layers of cells, outer 5-6
layers of polygonal, isodiametric, brown-
melanized, scleroplectenchymatic cells (3.6—
4.3 x 7-8 um) giving rise to the thick-walled

hyphae of the tomentum, middle 6—9 layers
composed of hyaline, polygonal-elongate,
scleroplectenchymatic cells (S—7 x 10-12 um),
inner 10—20 layers of small, polygonal, thick-
walled, hyaline cells (2.0—5.7 um diameter),
which become compressed and obscured
toward the interior of the cavity. Papilla
broadly rounded to dome-shaped, | 10—160 um
high, 160-250 tum wide at the apex, 220-300
lim wide at the base; wall 36—54 um thick,
composed of 12-15 layers of cells, outer layers
of brown-melanized, thickened, isodiametric
cells (3.6—5.7 um diameter), inner layers
hyaline, compressed, surrounding a 75-150 um
wide circular ostiole lined with periphyses.
Pseudoparaphyses 0.5—1.5 um wide, 250-375
tim long (height of the ascocarp cavity),
numerous, trabeculate with sparse branching,
with guttule-like thickenings at septa, without
gelatinous coating. Asci (95—)120—150 x 8.5—
12 um, numerous, basal and lateral, partially
lining the peripheral wall of the centrum,
cylindrical, thin-walled, short-stalked, rounded
apex, with 8 overlapping uniseriate ascospores.
Ascospores 17.2—22.3 x 5.7—8.0 um, fusiform,
with acute end cells, straight to slightly curved;
3-septate, septa evenly distributed, order of
septation 2:1:2, primary septum median and
slightly constricted, bipolarly asymmetrical
with a wider anterior and a narrower posterior
part, second cell slightly enlarged; pale brown
occasionally with slightly lighter colored end
cells, wall smooth, without sheath or append-
ages.

Exsiccatae: CZECHOSLOVAKIA: Betschwa-Ufer,
Dec. 27, 1911, M. Weisskirchen, Petrak, FI.
Boh et Mor. exsic. Lfg. 4 Nr. 196 (FH);
FINLAND: Bjork, Dec. 9, 1865, Karsten, Fungi
Fenniae 992 (FH); FINLAND: on Sambucus
racemosa, Jan. 1866, Karsten, Fungi Fenniae
995 (FH).

Illinois Natural History Survey Bulletin Vol. 34 Art. 5

g

Figure 13. Melanomma pulvis-pyrius. a. SEM of erumpent ascocarps, x35. b. ascocarp wall surface, x550.
c. longitudinal median section through ascocarp, x75. d, e. trabeculate pseudoparaphyses, x2,000.

f. longitudinal median section through ascocarp wall, x950. g. ascus, x525. h, i ascospores, x1,150. All from
holotype of Cladosphaeria rimicola.

May 1992 Leptosphaeria Species on the Rosaceae

Other material examined: SwiTZERLAND:
Bremengartenwald, on Prunus avium L.
(holotype of Cladosphaeria rimicola, BERN).
Unitep States: CALIFORNIA: Spruce Cove
Heads, Trinidad, Humbolt Co., on Rubus
parviflorus, 30 Jan. 1941, H.E. Parks 6513
(FH).

Comments: Leptosphaeria rimicola, described
from the fallen branches of Prunus avium L., is
synonymous with Melanomma pulvis-pyrius.
Leptosphaeria rimicola has the overall appear-
ance of M. pulvis-pyrius, with large, gregarious,
superficial ascocarps and ascospores with the
characteristic Melanomma shape: 3-septate
with the primary median septum dividing the
spore into a wider anterior and a narrower
posterior part. The measurements of the
ascospores of L. rimicola are somewhat larger
than were found by Chesters (1938) for M.
pulvis-pyrius, but they are not beyond the range
for M. pulvis-pyrius given by Saccardo (1878).
The asci peripherally lining the centrum about
halfway up the wall and the ascocarp wall
composed of small thickened cells correspond
to Barr’s (1987a) concept of the genus
Melanomma. The pseudoparaphyses are thin
and flexuous, but the branchings and anastomo-
ses are infrequent and the septa often show
thickenings, making them difficult to accurately
identify as trabeculate.

507

Leptosphaeria Species Referable to the Dothideales

The species in this chapter all belong in the
Dothideales (sensu Barr 1987a). Barr’s
concept of this order differs considerably from
the all-inclusive concept of von Arx and Miiller
(1975) or the broad concept of Eriksson and
Hawksworth (1985, 1986). The Dothideales
(Barr 1987a) are characterized by ascocarps
without a hamathecium (although interthecial
cells are often present) and by asci that tend to
be ovoid to saccate, arranged in a basal fascicle
or a basal layer. The families represented are
the Dothioraceae and the Pseudosphaeriaceae.

Dothioraceae

*Saccothecium sepincola (E.M. Fries:E.M.
Fries) E.M. Fries, Summa Vegetabilium
Scandinaviae, p. 398. 1849. See Figure 14.
= Sphaeria sepincola E.M. Fries,
Observationes Mycologicae. 1:181. 1815;
Systema Mycologicum Sistens Fungorum
2:498. 1823. [As saepincola.]
= Metasphaeria sepincola (E.M. Fries:
E.M. Fries) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo
2:164. 1883. [As Fr? Fuckel.]
= Leptosphaeria sepincola (E.M.
Fries:E.M. Fries) H.G. Winter, Dr. L.
Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der
Schweiz, Second edition, 1(2):473. 1885.
= Sphaerulina sepincola (E.M. Fries:
E.M. Fries) K. Starback, Botaniska
Notiser 1890:117. 1890; Botanisches
Zentralblatt, 46:261. 1891.
= Pringsheimia sepincola (E.M. Fries:
E.M. Fries) F. v. Hohnel, Annales
Mycologici 18:97. 1920.
= Pleosphaerulina sepincola (E.M. Fries:
E.M. Fries) H. Rehm in F. v. Héhnel,
Annales Mycologici 18:96. 1920.
= Sclerodothis sepincola (E.M. Fries:

E.M. Fries) F. Petrak, Annales
Mycologici 19:41. 1921.
For other synonyms see Barr (1972).

Ascocarps thickly scattered, immersed-
subepidermal, globose, glabrous, 180-200 um
diameter, 170-180 um high. Ascocarp wall of
textura angularis in surface view; in longitudi-
nal section 25—36 um thick at sides and base,
up to 45 um thick at apex, composed of 7-8
layers of pseudoparenchymatic, polygonal cells
(5—6 x 6-12 Lum), outer 2-3 layers of
isodiametric-to-elongate, brown-melanized
cells, inner 4—5 layers of hyaline-subhyaline,
thin-walled, elongate-compressed cells, at the
base giving rise to a central column (18-23 um
high, 27-36 um wide) of hyaline, isodiametric
cells (34 um diameter, but cell boundaries
becoming obscured in mass) on which the asci
are borne. Papilla broadly rounded, bluntly
conical, 45-55 um high, 55—70 um wide. Asci
40-60 x (13—)15—25 um, numerous, fasciculate,
borne on a basal column, oblong-ovate to
clavate, thick-walled, short-stalked, apex
rounded and thickened, with 8 bi- to triseriate
ascospores. Ascospores (17—)18.5—20.5(—22.5)
x 5.0—7.2 um, obovate, with acute end cells,
broad above and tapering to a narrower base,
straight to slightly curved; 4-5(-6)-septate,
septa unevenly distributed, septation order
3:2:1:2:4 or 4:2:1:3:5, primary septum
submedian, vertical septum occasionally
present in second or third cell from the top;
hyaline, wall smooth, without sheath or
appendages.

Lectotype: SwepEN: Ad ramos emortuos Rosae
aliorumque fruticum (UPS).

Exsiccatae: AusTtRIA: Nassau, on Rosa canina,
as Metasphaeria sepincola, 1894, Fuckel,
Herbier Barbey-Boissier 385 (FH); Nassau, on
Rubus fruticosus, as Sphaerulina intermixta
(Berk & Br) Sacc, 1894, Fuckel, Herbier

May 1992 Leptosphaeria Species on the Rosaceae 509

Figure 14. Saccothecium sepincola. a. SEM of immersed ascocarps, x125. b. longitudinal median section
through ascocarp wall, x950. c. longitudinal median section through ascocarp, x225. d. asci, x1,050.
e. ascocarp wall surface, x550. f. ascospores, x1,150. All from lectotype of S. sepincola.

Illinois Natural History Survey Bulletin

rbey-Boissier 501 (FH); CZECHOSLOVAKIA:
Weisskirchen, Ohrensdorf, as Sphaerulina
intermixta, 25 Jan. 1912, Petrak, Fl. Boh. et
Mor. exsic. Lfg. 4 Nr. 174. (FH); GERMANY:
Leihterfeld bei Berlin, on Philadelphus sp.,
1890, Sydow, Myc. March. 2934 (FH); Iracy:
Selva, as Sphaerulina intermixta, Sept. 1878,
Saccardo, Myc. Ven. 1367 (FH).

*Saccothecium sepincola var. abbreviata (M.C.
Cooke) S.M. Huhndorf comb. nov. See Figure
15.
= Sphaeria abbreviata M.C. Cooke,
Handbook of British Fungi, p. 893. 1871.
(Basionym).
= Leptosphaeria abbreviata (M.C.
Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:26.
1883.

Ascocarps | 17—144 um diameter, 81-99 ium
high; wall 14-21 tm thick at the sides and
apex, up to 27 um thick at the base, composed
of 7-8 layers of pseudoparenchymatic, poly-
gonal cells (5—6 x 6-12 tum). Papilla broadly
rounded. Asci (32—)41—61 x (13—)16.5—24.5
tum. Ascospores (12—)13.7—16.6 x 3.6—5.7 um,
4-5(-6)-septate.

Holotype: Great Britain: On dead stems of
bramble. Jan.—April (K).

Comments: Saccothecium sepincola is not
uncommon, but it has been confused by
mycologists over the years as exemplified by
the number of name changes based on the
fungus. Wehmeyer (1957) gave an account of
the history and nomenclatural confusion
surrounding this species, including its relation-
ships with other organisms and its relationship
to the later genus Pringsheimia Schulzer von
Miiggenburg. He also included a lectotypifi-
cation of Saccothecium Fr. Barr (1972) lists the
taxonomic synonyms of Saccothecium
sepincola. In some treatments, Pringsheimia
sepincola is still used as the name for this
species (Froidevaux 1973, von Arx and Miiller
1975, Sivanesan 1984). Holm (1975) argued
for the lectotypification of Saccothecium Fr.
1835 by S. sepincola (Fr.) Fr. 1849, which, as
mentioned above, apparently was already done
by Wehmeyer (1957) (see Dennis 1978). Holm
(1975) does not mention the lectotypification
by Wehmeyer, so it is unknown if this

Vol. 34 Art. 5

lectotypification was not accepted or if Holm
overlooked Wehmeyer’s work. In any case,
with lectotypification, Saccothecium is the
correct name and Pringsheimia becomes a
synonym.

Saccothecium sepincola is placed in the
family Dothioraceae by Barr (1987a) and is
characterized by sphaeroid ascomata with walls
of pseudoparenchymatous cells and oblong to
clavate asci with a thickened apex which arise
from a central basal column or mound of
hypothecial cells. The septate, hyaline, obovate
ascospores usually have a vertical septum
present in one or more of the central cells.
These characters suggest a similarity to some
Dothiora species with raised basal areas (Barr
1972).

Leptosphaeria abbreviata is regarded as a
variety of Saccothecium sepincola because
ascospore sizes differ. The ascospores of S.
sepincola are 17—22 x 5—7 um, whereas the
spores of S. sepincola var. abbreviata are 12—
16 x 3-6 um. The ascospores appear to be
mature in specimens of both species. The
ascocarps also differ in size between the two
species in the specimens seen.

Some discrepancy exists between
Cooke’s (1871) description of Sphaeria
abbreviata and the fungus that was present on
the type specimen. Cooke’s (1871) description
includes “perithecia minute, in short parallel
lines” and “asci very short and broad, elliptical,
pyriform or obovate,” which matches the
fungus in the type. But then he describes
spores that are “triseptate, slightly torulose and
pale brown,” which does not match this fungus,
although occasionally the spores when mature
may appear slightly pale brown. It is unclear
whether his description was simply inaccurate
or whether he was looking at two different
fungi. Only one fungus is present on the type
specimen and it was similar to S. sepincola.

Pseudosphaeriaceae

eLeptosphaerulina pulchra (H.G. Winter) M.E.
Barr, Contributions de I’Institut Botanique de
L’Université de Montréal 73:7. 1959. See
Figure 16.
= Sphaerella pulchra H.G. Winter,
Hedwigia 11:145—146. 1872.
= Leptosphaeria pulchra (H.G. Winter)
P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:53—54. 1883.

May 1992

Leptosphaeria Species on the Rosaceae 511

Figure 15. Saccothecium sepincola var. abbreviata. a. longitudinal median section through ascocarp, x375.
b. ascocarp wall surface, x550. c. longitudinal median section through ascocarp wall, x950. d. ascus, x1,050.
e. ascospores within an ascus, x1,150. All from holotype of S. sepincola var. abbreviata.

= Mycotodea pulchra (H.G. Winter) W.

Kirschstein, Annales Mycologici 34:201.

1936; Kryptogamenflora de Mark
Brandenburg und Angrenzender Gebiete
herausgegeben von den Botanischen
Verein der Provinz Brandenburg
7(3):433. 1938.
= Leptosphaeria oligotheca F. Petrak & H.
Sydow, Annales Mycologici 22:359. 1924.
[Described from type material of Laestadia
potentillae E. Rostrup, Botanisk Tidsskrift
25:300. 1903.]
For other synonyms see Barr (1959).

Ascocarps thickly scattered, immersed-
subcuticular, globose, papillate, glabrous, 5O—
75 um diameter, 55—75 um high. Ascocarp
wall of textura globulosa in surface view; in
longitudinal section uniformly 7.2—10.8(—12.7)
tum thick, composed of 2-3 layers of
pseudoparenchymatic, polygonal cells, outer 1—
2 layers of brown-melanized cells (3.6—5.6 x
5.6—-10.1 um) covered by a brown-pigmented
crust, inner 1—2 layers of hyaline, elongate-
compressed cells (2.1—3.6 x 8.6—10.1 um).
Papilla short, erumpent, bluntly conical, 10-20
tim high, 10—21 [tm wide at apex, 21-29 um

Illinois Natural History Survey Bulletin Vol. 34 Art. 5

Figure 16. Leptosphaerulina pulchra. a. SEM of ascocarp, x250. b. ascocarp wall surface, x550. c, d. longi-
tudinal median section through ascocarps, x650. e. longitudinal median sections through ascocarp wall, x950.
f. ascus, x1,050. g, h. ascospores, x1,150. a, b, d, and f from holotype of Leptosphaeria oligotheca: c. e, g.
and h from neotype of Leptosphaerulina pulchra.

May 1992

wide at base, neck wall 3.6—8.6 [1m thick,
composed of 2—3 layers of isodiametric-
elongate cells (3.64.3 x 3.6—6.5 [im) sur-
rounding a circular ostiole (9-12 um wide)
without periphyses. Remnants of interthecial
tissue remain between and above the asci. Asci
32-45 x 15-20 um, few, basal, fasciculate,
ovoid-saccate, thick-walled, sessile, rounded
apex, without apical chamber, with 8 tri- to
tetraseriate ascospores. Ascospores 1|7.2—20.8
x 5.7-7.2 um, clavate to obovoid, bipolarly
asymmetrical with broadly rounded apical end
cell and acutely rounded basal end cell,
straight; 4(-5)-septate, septa unevenly distrib-
uted, order of septation 3:1:2:4, primary septum
supramedian and constricted, a single vertical
septum occasionally present in the central cells;
subhyaline to pale brownish yellow, wall
smooth, without sheath or appendages.

Holotype: Austria: Ad caules foliaque aridos
Potentillae caulescentis, “am Martinstein bei
Seis in Tyrol” 1870 von v. Hausmann
gesammelt (B), (n.v.), presumed destroyed.

Neotype: Unitep States: Maine: Basin Pond,
Baxter St. Park, July 5, 1962, MEBB # 3316
(NY).

Other material examined: Canapa: Labrador,
Newfoundland, Blanc Sablon, July 19, 1957,
R.T. Wilce #161 (NY). ICELAND: on Potentilla
maculata, type material of Laestadia
potentillae Rostrup, July 14, 1884 (Holotype of
Leptosphaeria oligotheca, C). UNITED STATES:
Maine: Mt. Katahdin, Baxter St. Park, Table-
lands from Saddle Trail, ca. 4200', Aug. 3,
1962, MEBB # 3546 (NY); New HAMPSHIRE:
Mt. Washington, on Potentilla tridentata, June
9, 1894 (FH as Sphaerulina potentillae); Mt.
Monadnock, near Dublin, June 27, 1961,
MEBB # 2927 (NY).

Comments: Leptosphaerulina pulchra is placed
in the family Pseudosphaeriaceae because of
the minute, sphaeroid-globose ascocarps with a
very thin wall composed of pseudoparenchy-
matous cells and the interthecial tissues. The
species is recognized readily on the basis of its
saccate asci, ascospore shape, and septation.
Barr (1959) mentions the presence of a vertical
septum in the central cells, but in most of the
collections seen, vertical septa were lacking.

Leptosphaeria Species on the Rosaceae

513

Still, the obovate, asymmetrical ascospore
shape is distinctive.

The holotype of Leptosphaerulina
pulchra at B, not located, is presumed to have
been destroyed with the rest of the ascomycete
collection during World War II. Also unsuc-
cessful were attempts to locate authentic
material of Winter or of the collector, von
Hausmann. Because of this, a neotype for L.
pulchra was chosen (Maine: Basin Pond,
Baxter St. Park, July 5, 1962, MEBB # 3316)
from the M.E. Barr collection now at NY. The
neotype reflects both the original description
and the current, well-established concept of this
species.

Leptosphaeria oligotheca is synonymous
with Leptosphaerulina pulchra. When Petrak
and Sydow (1924) originally described
Leptosphaeria oligotheca from material on the
type of Laestadia potentillae, on stems of
Potentilla maculata, they did not segregate any
of the material as the holotype for L. oligotheca
and none could be found with their herbarium
specimens at W or S. The type specimen of
Laestadia potentillae at C contained abundant
material matching the description of
Leptosphaeria oligotheca, so a portion was
segregated as the holotype of that fungus.

Barr (1959) mentions several other
species whose descriptions suggest they may be
synonymous with Leptosphaerulina pulchra.
One of these was Sphaeria minima Duby in C.
Roumeguere, Fungi Selecti Gallici Exsiccati,
Century 7, No. 694. Anno 1880. Examining
specimens from the exsiccatae sets at G, FH,
NY, and ILL showed that they included many
ascocarps, but none contained asci or asco-
spores. Therefore, it was impossible to resolve
the placement of this species.

*Diadema obtusa R.A. Shoemaker & C.E.
Babcock, Canadian Journal of Botany 67:
1353-1354. 1989. See Figure 17.

Ascocarps thickly scattered, immersed-
subcuticular becoming erumpent, globose,
glabrous to slightly tomentose, with smooth
brown hairs, 100-150 ium diameter, 120-150
tum high. Ascocarp wall of textura angularis
in surface view; in longitudinal section (7.5—)
10-18 tm thick at sides and base, composed of
2-3 layers of brown-melanized, polygonal,
pseudoparenchymatic cells (1.84.3 x 7-11

Illinois Natural History Survey Bulletin Vol. 34 Ant. 5

= f

Figure 17. Diadema obtusa. a. longitudinal median section through ascocarp, x250. b. ascocarp wall
surface, x550. c. hyphae on substrate surface, x550. d. longitudinal median section through ascocarp wall,
x950. e. ascus, x525. f. ascospore, x1,050. All from collection 123871, on Potentilla agrophylla.

tim). Apex broadly rounded, 18—27 um thick, tissues present. Asci (100—)120—145 x (30-)

45—65 um diameter, composed of 2—3 outer 50-70 um, few, basal, fasciculate, ovoid-

layers of brown-melanized, isodiametric cells saccate to clavate, thick-walled, short-stalked,
(3.6—5.4 um diameter) with 6-8 layers of rounded apex, with apical chamber, with 8 tri-
hyaline pseudoparenchyma below center, to tetraseriate ascospores. Ascospores (41.5—)

Opening area not seen, possibly discoid, caplike | 45—5O0(—55) x (15—17—)19—23 um, broadly
opening lacking, ostiole not seen. Interthecial fusiform, acutely rounded end cells, second cell

May 1992

enlarged, straight to slightly curved, occasion-
ally flattened in one plane; 3-septate, septa
slightly unevenly distributed, order of septation
2:1:2, primary septum median and constricted,
slightly constricted at other septa; brown, wall
smooth, sharply delimited sheath entirely
surrounding spore (5—7.5 tm thick), without
appendages.

Holotype: INDIA: KASHMIR: 123831(b), on
Trisetum spicatum, Pensi La, Zaskar, 16,500 ft.
W. Koelz (5906), 23 July 1933, TYPE, ex
Herb. Wehmeyer, as Leptosphaeria hollosiana
nom. nov. (DAOM).

Other material examined: INpIA: 123871, on
Potentilla agrophylla Wall., Spiti Valley, Bara
Lacha Pass, 16,500 ft, Bhagwan Singh (5), 8
July 1932, ex Herb. Wehmeyer, as
Leptosphaeria hollosiana nom. nov. (DAOM).

Comments: Shoemaker and Babcock (1989a)
established the new genus Diadema for a group
of alpine fungi with relatively large, very dark
brown ascospores. They did not place the
genus within a family or order. Eriksson and
Hawksworth (1990) placed it in the
Dothideales but did not designate a family. In
the present report I treat it under the
Pseudosphaeriaceae because it resembles
Wettsteinina Hohnel; for example, some
species have a peculiar disclike opening of the
ascocarp and have interthecial tissue in the
centrum. Diadema is characterized by globose,
subcuticular ascocarps with a thin wall of
brown polygonal cells and broadly fusiform
ascospores that are dark brown and have a
prominent sheath.

Wehmeyer (1963) published the name
Leptosphaeria hollosiana for Leptosphaeria
maritima L. Hollos because the latter binomial
was predated by L. maritima (Cke. & Plowr.)
Sacce. He applied the new name to several
collections from India and Pakistan, including
two collections on Potentilla. Shoemaker and
Babcock (1989a) described two new species of
Diadema from these collections, including the
type of the genus, D. acuta, on Trisetum
spicatum (L.) Richt., and D. obtusa, also on that
host. The collections on Potentilla were also
determined to be D. obtusa. The collection on
Potentilla served as the basis for this descrip-
tion and plate because of the Rosaceous host
and because it had numerous ascocarps. The
other collections of D. obtusa were sparse.

Leptosphaeria Species on the Rosaceae S15

As Shoemaker and Babcock (1989a)
noted, the ascocarp opening mechanism in this
species is not clearly understood. It is not
distinctly caplike but in vertical section
becomes almost papillate (Figure 17a). The
entire upper section of the wall is subtended by
hyaline pseudoparenchyma. Interthecial
pseudoparenchymatic threads occurred between
the asci. Also, coarse, brown hyphae were seen
on the host surrounding several of the asco-
carps, and some of the ascocarps were not
glabrous but had tapered hairs at the top. I did
not see the ascospores flattened in one plane, as
did Shoemaker and Babcock (1989a). Shoe-
maker and Babcock (1989a) suggested that the
generic placement of this species is not ideal
because several characters were not identical to
those of the type species. The overall aspect of
this species, however, suggests an affinity with
the type species, and that it is adequately placed
in this genus. It appears that certain characters,
such as the ascocarp cap and interthecial
tissues, are variable among collections and may
depend on the maturity of the specimen, as was
also noted by Shoemaker and Babcock (1989a).
Species of Weftsteinina are similarly variable,
either showing a disclike cap or an ostiolar
opening to the ascocarp; species with either
character state are adequately retained within
the genus because of overall resemblances
(Shoemaker and Babcock 1987). Thus, such
variation does not appear sufficient to exclude
species from these genera.

*Diadema sieversiae (C.H. Peck) S.M.
Huhndorf comb. nov. See Figure 18.
= Lophiostoma sieversiae C.H. Peck,
New York State Museum Bulletin
167:44. 1913. (Basionym).
= Wettsteinina sieversiae (C.H. Peck)
M.E. Barr, Canadian Journal of Botany
45:1042. 1967.
= Massaria sieversiae F.E. Clements,
Cryptogamae Formationum Coloradensium,
Century II, No. 234. Anno 1906. Nom.
nud. Art. 36.1.
= Leptosphaeria sieversiae (F.E.
Clements) F. Petrak, Sydowia. Annales
Mycologici 6(1-4):6. 1952. [Combina-
tion is not valid because the basionym is
an invalidly published species. ]

Ascocarps thickly scattered, immersed-
subcuticular becoming erumpent, globose,

Illinois Natural History Survey Bulletin Vol. 34 Art. 5

Figure 18. Diadema sieversiae. a. SEM of erumpent ascocarps, x125. b. ascocarp wall surface, x550.

c. longitudinal median section through ascocarp, x250. d. hyphae on substrate surface, x200. e. elongated
ascus, X250. f. longitudinal median section through ascocarp wall, x950. g. ascus, x525. h. ascospore,
x1,050. a, b from holotype of D. sieversiae; ch from isotype of Massaria sieversiae.

May 1992

glabrous to slightly tomentose, with loose dark-
walled, longitudinal hyphae on the substrate,
surrounding and connecting the ascocarps,
130-190 um diameter, 120-175 um high.
Ascocarp wall of textura angularis in surface
view; in longitudinal section uniformly 12.6—20
uum thick, composed of 3-4 layers of brown-
melanized, pseudoparenchymatic, polygonal,
isodiametric-elongate cells (3.6—5.6 x 7.2-13
tim) with a brown-melanized crust, at base 3-4
inner layers of small, hyaline pseudoparen-
chyma. Apex broadly rounded, with inner 3-4
layers of hyaline, pseudoparenchymatic cells
(2-5 um diameter), no evidence of papillate or
caplike opening, ostiole not seen. Interascal
pseudoparenchyma present. Asci (110—)130—
170 x 40-70 um, few, basal, fasciculate, ovoid-
saccate, thick-walled, sessile, rounded apex,
with apical chamber, with 8 tri- to tetraseriate
ascospores. Ascospores (43.5—)50.6—56.0
(—57.9) x (16—)20—23.5(—25.5) um, broadly
fusiform, with acutely rounded end cells,
second cell enlarged, straight to slightly curved;
3-septate, septa slightly unevenly distributed,
order of septation 2:1:2, primary septum
slightly supramedian and constricted, occasion-
ally constricted at other septa; brown, wall
smooth, sharply delimited sheath entirely
surrounding spore [2.0—5.5(—10.5) um thick,
often constricted at midseptum, sheath occa-
sionally gelatinizing and becoming amorphous
(7-12 um thick)], without appendages.

Holotype: UNirep States: UTAH: 193727, on
Sieversia turbinata (Rydb.) Greene, Big
Cottonwood Canyon, Salt Lake Co., A.O.
Garrett 702, 3 July 1913 (NYS).

Exsiccatae: UniTep States: COLORADO: Bottom-
less Pit, July 13, 1906, Clements, Crypt. Form.
Colorad. 234 (isotype of Massaria sieversiae,
NY, FH).

Comments: Diadema sieversiae, found on
stems of Acomastylis turbinata, is characterized
by globose ascocarps with thin walls of brown
polygonal cells, dark-walled hyphae on the
substrate, and broadly fusiform, dark brown
ascospores with a prominant gelatinous sheath.
It bears a strong resemblance to D. obtusa, but
in that species the spores are slightly shorter
and the dark brown hyphae in the host are not
as prevalent. The ascocarp shapes differ
somewhat, but I also saw no indication of a cap

Leptosphaeria Species on the Rosaceae

517

or disclike opening mechanism in D.
sieversiae. Interthecial tissues were seen
between the asci.

Diadema sieversiae was described by
Peck (1913) as a species of Lophiostoma, but it
bears no compressed beak or ostiole that would
warrant inclusion in that genus. Barr (1967)
transferred it to Wertsteinina because of the
medium- to small-sized ascocarps immersed in
dead host tissues and the oblong to saccate asci
interspersed with interthecial tissue. Shoe-
maker and Babcock (1987) excluded it from
Wettsteinina because of the dark brown
ascospores with thick septa that are unlike the
usual spores for Wettsteinina. It was left in
Lophiostoma and later (Shoemaker and
Babcock 1989a) was not included in Diadema.
Clearly this fungus does not belong in
Lophiostoma, and it bears a much greater
resemblance to Diadema than to Wettsteinina.
Even if it ultimately does not stay in Diadema,
because of the lack of a disclike cap opening
(as suggested for D. obtusa by Shoemaker and
Babcock 1989a), it seems useful to place this
species in Diadema, in the hope that additional
collections may be discovered.

Massaria sieversiae, which was trans-
ferred to Leptosphaeria, is synonymous with
Diadema sieversiae.

Leptosphaeria Species Referable to the Hymenoascomycetes

The following Leptosphaeria species are all
referable to the class Hymenoascomycetes
(sensu Barr 1987a). The presence of perithecia
and unitunicate asci separates these fungi from
those in the Loculoascomycetes possessing
pseudothecia and bitunicate asci. The families
represented are Clypeosphaeriaceae,
Diaporthaceae, and Amphisphaeriaceae.

Clypeosphaeriaceae

*Clypeosphaeria mamillana (E.M. Fries:E.M.
Fries) J.B.E. Lambotte, Memoires de la Societe
Royale des Sciences de Liege, ser 2, 14:128.
1887. See Figure 19.
= Sphaeria mamillana E.M. Fries:E.M.
Fries, Systema Mycologicum 2:487.
1823.
= Sphaeria clypeiformis G. de Notaris,
Memorie della Accademie della Scienze di
Torino Series 2, No. 7, p. 113. 1853;
Micromycetes Italici Novi vel Minus
Cogniti 7, p. 113. 1845. Non Sphaeria
clypeiformis L.V. de Lacroix in G.L.
Rabenhorst.
= Clypeosphaeria notarisii L. Fuckel,
Symbolae Mycologicae, p. 117. 1870.
= Leptosphaeria (Clypeosphaeria)
notarisit M.C. Cooke, Grevillea 17:91.
1889.
See Barr (1989) for other synonyms.

Ascocarps scattered, separate or gregarious,
immersed-subepidermal beneath blackened
clypei, papilla erumpent, glabrous, conic-
globose, 300-400 um diameter, 400-450 um
high. Ascocarp wall of textura angularis in
surface view; in longitudinal section uniformly
18—25 tm thick, composed of 10-15 layers of
hyaline, elongate-compressed, scleroplecten-
chymatic cells (0.7—1.5 x 7-15 tum), outer 2-3
layers of hyaline, rounded, polygonal cells
(2.0-3.6 um diameter). Papilla conic, 130-160
um high, 50-75 um wide at the apex, 150-225

lim wide at the base; wall 15—30 um thick at
apex, 35-65 1m thick at base, composed of 10—
15 layers of small, brown-melanized, thick-
walled, rounded, polygonal cells at base (1.5—
3.6 um diam), cells converging and appearing
as elongate, setaelike structures at apex,
surrounding a 25—50 um wide circular ostiole
with periphyses. Paraphyses 0.5—1.5 um
wide, numerous. Asci 150-170 x 8-10 um,
unitunicate, numerous, basal to peripheral,
cylindrical, short-stalked, rounded apex, apical
ring J+, with 8 overlapping uniseriate ascos-
pores. Ascospores 21—26 x 5.7—8 um, ovoid-
oblong, straight to slightly curved; at times 1-
septate near base, large cell appearing 3-septate
with separation of cytoplasm; brown, smooth,
without sheath or appendages, germ slits not
seen.

Exsiccatae: Austria: On Rubus fruticosus,
Fuckel, F. rhen. 1823 (FH); Mappen, on
Epilobium angustifolii, as Sordaria
clypeiformis f. Epilobii, Fuckel, F. rhen. 2036
(FH); Herb. Barbey-Boissier 146 = F. rhen.
1823, as C. notarisii (NY); GREAT BRITAIN:
Lynn, on Epilobium hirsutum, as Sphaeria
clypeata Nees., Jan. 1877, Plowright, Sphaer.
Brit. 57 (FH); Ivacy: on Rubus fruticosus, as
Sphaeria clypeiformis, Aug. 1857, leg. Caldesi,
Rabenhorst, Herb. myc. 645 (FH); as Sphaeria
clypeiformis, 1857, leg. Caldesi, de Notaris,
Erb. Critt. Ital. 47 (FH); La Spezia, as C.
notaristi, Aug. 1880, Roumeguere, F. Gall. exs.
2095 (NY): Vittorio (Treviso), on Rubus

fruticosus, as C. notarisii, Sept. 1897,

Saccardo, Mycotheca ital. 101 (NY):
Conegliano, as C. notarisii, Aug. 1877,
Mycotheca veneta 1159 (NY); St. Romani-ad-
Vigennam, as S. clypeiformis de Lerx., 1859,
leg. T. de Lacroix, Rabenhorst, F. europ. 331
(RO).

Other material examined: ITALy: Capraria, on
Rubus, as Sphaeria sepincola ruborum, 1837,

May 1992

:
‘

Leptosphaeria Species on the Rosaceae

a.
Wh:

319

Figure 19. Clypeosphaeria mamillana. a. longitudinal median section through ascocarp, x100. b, c. asco-
spores, x1,150. d. longitudinal median section through ascocarp wall, x950. e. ascus, x525. All from

lectotype of C. notarisii.

de Notaris (Lectotype of Sphaeria
clypeiformis); two collections of Sphaeria
clypeata, 1840; Spezia, Aug. 1857, Herb. N.A.
Pedicino, 1883; on Rubus, as S. clypeiformis,
Mar. 17, 1842, de Notaris (all RO).

Comments: Clypeosphaeria notarisii, a
synonym of Clypeosphaeria mamillana, is
characterized by cylindrical asci with an
elongate, J+ apical ring and curved, brown
ascospores appearing 3-septate with separation
of the cytoplasm. Barr (1989) gives the

synonymy and a short history of
Clypeosphaeria and argues for the utilization of
the family Clypeosphaeriaceae.

Clypeosphaeria was established by
Fuckel (1870), who included C. notarisii as a
new name for Sphaeria clypeiformis de Not. A
specimen of Sphaeria sepincola ruborum was
mentioned in the description of Sphaeria
clypeiformis, and a collection of this at RO is
here designated as the lectotype of S.
clypeiformis (ITALy: Capraria, on Rubus, 1837,
de Notaris, RO).

Illinois Natural History Survey Bulletin

Diaporthaceae

*Diaporthe eres T.R.J. Nitschke, Pyrenomy-
cetes germanici, p. 245. 1867. See Figure 20.
= Sphaeria controversa J. Desmazierés (pro
parte), Annales des Sciences Naturelles,
Paris, Botanique, Series 2, 17:102. 1842.
= Leptosphaeria controversa (J.
Desmazierés) V. Cesati & G. de Notaris,
Commentario della Societa
Crittogamologica Italiana 1:235. 1863.
= Diaporthe controversa (J.
Desmazierés) T.R.J. Nitschke in L.
Fuckel, Symbolae Mycologicae Nachtrag
1:319. 1871.
See Wehmeyer (1933) for other synonyms.

Ascocarps scattered, separate or clustered,
immersed-subepidermal often beneath black-
ened stromatic tissue, papilla erumpent singly
or in loose clusters, glabrous, globose, flattened
at top and base, 300-400 um diameter, 250—
350 um high. Ascocarp wall of textura
angularis in surface view; in longitudinal
section 25—35 um thick at the sides, 15—20 um
thick at the base, composed of 7-8 layers of
hyaline, elongate-compressed, scleroplecten-
chymatic cells (0.7—1.5 x 10-15 tm), wall near
apex 45—50 um thick, composed of 10-12
layers of brown-melanized, polygonal, isodia-
metric cells (2-3 um diam). Papilla long
cylindrical, central or eccentric, 125-180 um
high, 50-75 um wide; wall 25—35 um thick,
composed of small, brown-melanized, thick-
walled, rounded, polygonal cells (1.5—3.0 um
diameter), surrounding a 25-40 um wide
circular ostiole with periphyses. Paraphyses
0.5—1.5 [tm wide, numerous, septate, un-
branched. Asci 38—55 x 6—8.5 tim, unitunicate,
numerous, basal to peripheral, dehiscent,
clavate, short-stalked, rounded apex, with 8
biseriate ascospores. Ascospores 10.5—13.5 x
2.0—3.6 |im, fusiform, straight to slightly
curved; |-septate, constricted; hyaline, with
four distinct oil droplets, smooth, without
sheath, occasionally with small, cellular bipolar
appendages.

Isotype: FRANCE: Habitat in caulibus plantarum,
in ramis exsiccatis Fraxini, Rubi, Aceris
Negundinis, etc. Desm. Pl. Crypt. Fr. exs.
XXVI 1255 (PC, FH, BPI).

Comments: Sphaeria controversa (pro parte) is
synonymous with Diaporthe eres (Wehmeyer

Vol. 34 Art. 5

1933) and is characterized by ascocarps
immersed beneath blackened stromata, with
central or eccentric, long cylindrical necks with
periphyses. As they mature, the numerous asci
loosen and become free within the centrum.
Ascospores are hyaline and 1-septate and have
four distinct oil droplets.

The exsiccati collections all consisted of
several different twigs of unknown identity,
none of which appeared to be Rubus, most with
varying amounts of the fungus on them.

Amphisphaeriaceae

*Discostroma fuscella (M.J. Berkeley & C.E.
Broome) S.M. Huhndorf comb. nov. See
Figure 21.
= Sphaeria (Obtectae) fuscella M.J.
Berkeley & C.E. Broome, Annals and
Magazine of Natural History, London,
Series 2, 9:325. 1852. (Basionym).
= Leptosphaeria fuscella (M.J. Berkeley
& C.E. Broome) V. Cesati & G. de
Notaris, Commentario della Societa
Crittogamologica Italiana 1:236. 1863.
= Sphaeria corticola L. Fuckel, Symbolae
Mycologicae, p. 114, 1870.
= Leptosphaeria corticola (L. Fuckel)
P.A. Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 1:342. 1878; Fungi Italici
autographice delineati (additis nonnullis
extra-italicis asterisco notatis), Patavii,
table 288. 1878.
= Metasphaeria corticola (L. Fuckel)
P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit
P.A. Saccardo 2:166. 1883.
= Griphosphaeria corticola (L. Fuckel)
F. v. Hohnel, Annales Mycologici 16:87.
1918.
= Clathridium corticola (L. Fuckel) R.A.
Shoemaker & E. Miiller, Canadian
Journal of Botany 42:404. 1964.
= Discostroma corticola (L. Fuckel) I.
Brockmann, Sydowia 28:313. 1975.
= Sphaeria (Leptosphaeria) leiostega J.B.
Ellis, Bulletin of the Torrey Botanical Club
(and Torreya) 8:91. 1881.
= Leptosphaeria leiostega (J.B. Ellis) J.B.
Ellis, Catalogue of Plants Found in New
Jersey. Geological Survey of New Jersey,
Final Report of the State Geologist
2(1):525. 1889.

May 1992 Leptosphaeria Species on the Rosaceae 521

Figure 20. Diaporthe eres. a. longitudinal median section through ascocarp, x150. b. section through
ascocarp neck showing periphyses, x400. c. longitudinal median section through ascocarp wall, x950.
d. ascocarp wall surface, x550. e. ascus, x1,050. f. ascospore, x1,150. All from type of Leptosphaeria
controversa.

= Leptosphaeria lejostega (J.B. Ellis) F. Ascocarps scattered, separate or clustered,
Hazslinszky, Matematikai és immersed-subepidermal beneath blackened
Természettudomanyi K6zlemenyek clypei, glabrous, depressed globose, flattened
Vonatkozolag a Hazai Viszonyokra top and bottom, with papilla protruding through
25(2):148. 1892. surface, 300-500 um diameter, 150-250 um

= Metasphaeria lejostega (J.B. Ellis) P.A. high. Ascocarp wall of textura prismatica in
Saccardo, Sylloge Fungorum Omnium surface view; in longitudinal section 8-18 um
Hucusque Cognitorum Digessit P.A. thick at base, 18—27 um thick at sides, com-
Saccardo 2:164. 1883. posed of 6-8 layers of brown, elongate-

For other synonyms see Brockmann (1975). compressed, scleroplectenchymatic cells (12-

Illinois Natural History Survey Bulletin Vol. 34 Art. 5

a
k
Figure 21. Discostroma fuscella. a. longitudinal median section through ascocarp, x150. b. ascocarp wall
urface, XxS50. c. section through ascocarp wall neck region, x550. d. longitudinal median section through
), X150. e. longitudinal median section through ascocarp wall, x950. f-h. asci, x525. i-k. ascospores,

¢, e, f, andi from holotype of D. fuscella; d, g, and j from lectotype of Sphaeria leiostega;
pe of S. corticola

May 1992

25 x 2-3 um); near apex, wall 21—36 um thick,
composed of 6—7 outer layers of brown,
polygonal-isodiametric cells (7—11.5 x 2-3;
2.1—3.6 um diameter) and 6—7 inner layers of
compressed-elongate, scleroplectenchymatic
cells. Papilla conical, 50-75 um high, 20—30
tim wide at the apex, 35-50 um wide at the
base; wall 7-10 tm thick, composed of 7—10

layers of small, brown-melanized, thick-walled,

rounded, polygonal cells, surrounding a 10-20
uum wide circular ostiole with periphyses.
Paraphyses 0.5—1.5 um wide, numerous,
septate, unbranched. Asci (101—)115—132
(-137) x 7-12(—14) um, unitunicate, numerous,
basal in a broad hymenium, cylindrical, short-
stalked, rounded apex, apical ring J+, with 8
overlapping uniseriate ascospores. Ascospores
15-21 x S—9.5 um, ellipsoidal, end cells
rounded; 3-septate, septa often close together
and end cells longer, seldom with 4 or more
septa (up to 7), occasionally with vertical septa
in central cells only or in every cell, order of
septation variable, often 2:1:2, primary septum
median and occasionally constricted; hyaline,
smooth, without sheath or appendages.

Holotype: Great Britain: Easton, Norths., on
dead twigs of rose (Rosa sp.), March 9, 1850
(K).

Exsiccatae: Austria: Fuckel, Fungi Rhenani
911 (Isotype of Sphaeria corticola, G, FH);
CZECHOSLOVAKIA: Welka, on Prunus spinosa, as
Metasphaeria corticola f. Pruni, Oct. 3, 1912,
Petrak, Fl. Boh et Mor. exsic. 61 (FH); Welka,
on Rosa canina, as Metasphaeria corticola f.
Rosae, Apr. 17, 1912, Petrak, Fl. Boh et Mor.
exsic. 62 (FH); Ohrensdorf, on Rubus
thyrsoideus, as Metasphaeria corticola f. Rubi,
Sept. 25, 1911, Petrak, Fl. Boh et Mor. exsic.
63 (FH); Swepen: Lockerud, Wenenborg, on
Rubus fruticosus, as Leptosphaeria lejostega
(Ellis) Eliasson, Aug. 1892, Rehm, Asc. 1189
(FH); Unirep States: New Jersey: Newfield, as
Sphaeria (Lept.) leiostega, Ellis, N. American
Fungi 888 (NY).

Other material examined: Austria: Nassau, on
Prunus domestica and P. spinosa, Fuckel
(Isotype, as Metasphaeria corticola (Fuckel)
Sacc., FH); Unitep States: Kansas: Rockport,
Rooks Co., on Cornus asperifolia, as
Metasphaeria leiostega, Dec. 24, 1894, E.
Bartholomew (FH); Rooks Co., on Ribes

Leptosphaeria Species on the Rosaceae 523

aureum, as Clypeosphaeria hendersonia 2555,
Mar. 15, 1899, E. Bartholomew (NY); Micui-
GAN: 573, Ann Arbor, on Ribes frondium, as
Metasphaeria leiostega, Apr. 1, 1893, L.N.
Johnson (NY); New Jersey: Newfield, on wild
rose (Rosa lucida ?), May 6, 1879, J.B. Ellis
(lectotype of Sphaeria leiostega, NY);
Newfield, on Viburnum lentago, as Sphaeria
leiostega, Dec. 1878, J.B. Ellis (NY);
Newfield, Gloucester Co., on Hickory, as
Sphaeria leiostega (NY); on Staphylea trifolia,
as Metasphaeria leiostega, Dec. 23, 1891
(NY); on Viburnum, as Metasphaeria leiostega
(NY); 1273, on wild currant, as Metasphaeria
leiostega, Jan. 4, 1890 (NY); on Comptonia, as
Metasphaeria leiostega Mar. 1890, J.B. Ellis
(NY); Vineland, on Vaccinium corymbosum, as
Sphaeria leiostega, Jan. 22, 1878; Jan. 10, 1880
(NY); New York: Washington Co., Vaughans,
J. Vaughan’s Copse, near Devines Woods,
north of Hudson Falls, on Ribes americanum
(floridum), as Metasphaeria leiostega, Apr. 24,
1917, P.P. 25255, S.H. Burham (FH).

Comments: Discostroma fuscella is character-
ized by subepidermal perithecia with short
protruding papillae, cylindrical asci with a J+
apical ring and hyaline, 3-septate, elliptical
ascospores with broadly rounded ends. Certain
collections have wider spores and vertical
septa, most often in the central cells, but often
also in the end cells. Most notably, vertical
septa are found in the lectotype of Sphaeria
leiostega. All other characters are identical to
those of D. fuscella.

Discostroma corticola, the most common
name of this fungus, is identical to the earlier
named Sphaeria fuscella. Previous interpreta-
tions of Sphaeria fuscella have caused some
confusion in the literature. The type material
contains a fungus resembling D. corticola, with
hyaline ascospores. However, it has been
considered to be a 3-septate, brown-spored
fungus because of Berkeley and Broome’s
(1852) description of ascospores as “pale
brown, oblong-elliptic, obtuse, triseptate,
sometimes slightly curved.” In the accompany-
ing Latin description they do not mention the
color of the ascospores. On the type material
(K), labeled “Sphaeria fuscella B & B. In Rosa,
Easton, March 9, 1850,” are three rose twigs,
one of which is labeled (with a dotted line and
a figure) as Sphaeria (Subtectae) intermixta B
& B. The other two twigs contain abundant

Illinois Natural History Survey Bulletin

material of the hyaline-spored fungus resem-
bling D. corticola. Also on this material is a
figure of Sphaeria fuscella showing an ascus
and spores with “hyaline” written next to the
spores. Wehmeyer (1957) suggested that a
mixture of fungi in this and another collection
of S. fuscella resulted in a confused description.
He felt the name should be a nomen dubium
and not used as a basis for later names. I
believe, from the description and the fungus
found on the type material, and from the figure
on the type material, that Berkeley and
Broome’s confused description was not based
on a mixture of fungi but was a miscommunica-
tion of the actual information. I believe they
were aware that S. fuscella had hyaline, not
brown spores; thus, it is suitable for use as a
basis for later names.

Vol. 34 Art. 5

Species incertae sedis

The following species were examined but could
not be placed in any taxa with certainty because
the type specimens contained no ascocarps and

no other collections were available on which to
base the descriptions.

¢Leptosphaeria eriobotryae H. Sydow, P.
Sydow & E. Butler, Annales Mycologici 9:409.
1911.

Holotype: Hab. in foliis Eriobotrya japonicae,
Saharanpur, 27. 6. 1906, leg. Inayat (E.J. Butler
no. 1365). (S) Isotype: (W, HCIO)

eLeptosphaeria minima (J.E. Duby) ex P.A.
Saccardo, Michelia Commentarium
Mycologicum Fungos in Primis Italicos
Illustrans 2:320. 1881.
= Sphaeria minima J.E. Duby in C.
Roumegueére, Fungi Selecti Gallici
Exsiccati, Century 7, No. 694. Anno
1880. Nom. nud.
= Mycotodea minima (J.E. Duby ex P.A.
Saccardo) W. Kirschstein in O.C.
Schmidt, Kryptogamenflora der Mark
Brandenburg und Angrenzender Gebiete
herausgegeben von den Botanishen
Verein der Provinz Brandenburg
7(3):432. 1938.

Isotype: Sur les feuilles du Potentilla argentea
L., Mende, Prost. (G, FH, ILL, NY)

*Leptosphaeria sorbi A. Jaczewski, Annales
Mycologici 1:30. 1903.

Holotype: in foliis vivis Sorbi aucupariae, IX.
18 1914 , Gjatsk. Smolensk. Gouv. Jaczewski.
(LE)

Appendix: Leptosphaeria Species Described from the

Rosaceae

The following is a list of all Leptosphaeria species
described from the Rosaceae and the species’
nomenclatural synomyms. This information is taken
from Crane and Shearer (1991). The taxonomic
synonyms applied herein and the herbaria contacted
have been added. The fungal specimens cited in this
dissertation were studied at or on loan from various
herbaria. Requests were made to various herbaria
for type material for all of the described species.
Unfortunately, type material could not be located for
all of the species because either the material was
destroyed, lost, or not deposited at a certain
institution, or the institutions involved did not reply
or do not lend specimens. Abbreviations follow
Holmgren et al. (1981).

L. abbreviata (M.C. Cooke) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:26. 1883. =Sphaeria
abbreviata M.C. Cooke, Handbook of British
Fungi, p. 893. 1871. On dead stems of Rubus
sp., Rosaceae. Great Britain.
=Saccothecium sepincola (E.M. Fries:E.M.
Fries) E.M. Fries var. abbreviata S.M.
Huhndorf, comb. nov.

L. aculeorum G. Passerini, Atti della R. Accademia
dei Lincei Memoriae, Rome, Series 4, 6:458—
459. (1889) 1890. On stems of Rosa canina
L., Rosaceae. Italy. (non vidi). Type
material was not found at PARMA, PISA,
PAD.

L. arunci S.M. Zeller, Mycologia, Lancaster,
Pennsylvania 19:134—135. 1927. On dead
stems of Aruncus silvestris Kostel. (=Aruncus
dioicus (Walter) Fernald), Rosaceae. United
States (Oregon).
= Kalmusia clivensis (M.J. Berkeley & C.E.
Broome) M.E. Barr.

L. cercocarpi H. Sydow & P. Sydow, Annales
Mycologici, Berlin 5:339. 1907. On dead
leaves of Cercocarpus ledifolius Nutt. ex Torr.
& Gray, Rosaceae. United States.

L. concentrica J.B. Ellis & B.M. Everhart, The North
American Pyrenomycetes. A Contribution to
Mycologic Botany, p. 354. 1892. On leaves
of Malus sp., Rosaceae. United States.
=Paraphaeosphaeria concentrica (J.B. Ellis
& B.M. Everhart) S.M. Huhndorf, comb. nov.

L. coniothyrium (L. Fuckel) P.A. Saccardo,Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 7:317.
1875. =Sphaeria coniothyrium L. Fuckel,
Symbolae Mycologicae, p. 115. 1870.
=Melanomma coniothyrium (L. Fuckel) L.
Holm, Symbolae Botanicae Upsalienses,
Uppsala 14(3):56. 1957. On stems of Alnus
glutinosa (L.) Gaertner, Broussonetia sp.,
Chimonanthus sp., Kerria sp., Menispermum
canadense L., Rosa sp., Rubus fruticosus L.,
Salix vitellina L., Sambucus nigra L.,
Betulaceae, Moraceae, Calycanthaceae,
Rosaceae, Menispermaceae, Salicaceae,
Caprifoliaceae. Austria, France, Germany,
Italy, Portugal.
=Kalmusia coniothyrium (L. Fuckel) S.M.
Huhndorf, comb. nov.

L. controversa (J. Desmazierés) V. Cesati & G. de
Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:235. 1863.
=Sphaeria controversa J. Desmazierés,
Annales des Sciences Naturelles, Paris,
Botanique, Series 2, 17:102. 1842.
=Diaporthe controversa (J. Desmazierés)
T.R.J. Nitschke in L. Fuckel, Symbolae
Mycologicae Nachtrag 1:319. 1871. On plant
stems, on dry branches of Acer sp., Fraxinus
sp., Rubus sp., Aceraceae, Oleaceae,
Rosaceae. France.
=Diaporthe eres T.R.J. Nitschke, fide L.E.
Wehmeyer, The Genus Diaporthe Nitschke
and its Segregates, p. 63. 1933.

L. corticola (L. Fuckel) P.A. Saccardo, Michelia
Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 1:342. 1878; Fungi
Italici autographice delineati (additis nonnullis
extra-italicis asterisco notatis), Patavii, table
288. 1878. =Sphaeria corticola L. Fuckel,
Symbolae Mycologicae, p. 114. 1870.
=Metasphaeria corticola (L. Fuckel) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:166. 1883. =Griphosphaeria corticola (L.
Fuckel) F. v. Hoehnel, Annales Mycologici,
Berlin 16:87. 1918. =Clathridium corticola
(L. Fuckel) R.A. Shoemaker & E. Mueller,
Canadian Journal of Botany, Ottawa 42:404.

May 1992

1964. =Discostroma corticola (L. Fuckel) I.
Brockmann, Sydowia 28:313. 1975. On
cortex of branchlets, on dry stems of Cornus
sanguinea L., Lonicera caprifolium L.,
Prunus baccata Borkh., Prunus domestica L.,
Prunus spinosa L., Rosa sp., Caprifoliaceae,
Rosaceae. Austria, Canada, Czechoslovakia,
Germany, Hungary, Italy, Switzerland.
=Discostroma fuscella (M.J. Berkeley & C.E.
Broome) S.M. Huhndorf, comb. nov.

L. dryadis E. Rostrup, Botanisk Tidsskrift,
Kjobenhavn 25:305. 1903. On decorticated
wood stems and fruits of Dryas octopetala L.,
Rosaceae. Iceland.
=Leptosphaeria dryadophila S.M. Huhndorf,
nom. nov.

L. dryadis E. Rostrup, Norske Ascomyceter i
Christiania Universitetets Botaniske Museum.
Christiania [Oslo] (I. Kommission Hos Jacob
Dybwad), p. 24. 1904. [Issued in Skrifter
Udgivne af Videnskabselskabet i Christiania.
Mathematisk-naturvidenskabelig klasse.
Christiania (Oslo) No. 274.] Non vidi. On
upper surface of dried leaves of Dryas
octopetala L., Rosaceae. Norway. (n. v.)

L. eriobotryae H. Sydow, P. Sydow & E. Butler,
Annales Mycologici, Berlin 9:409. 1911. On
leaves of Eriobotrya japonica (Thunb.)
Lindley, Rosaceae. India.

Species incertae sedis.

L. fuscella (M.J. Berkeley & C.E. Broome) V. Cesati
& G. de Notaris, Commentario della Societa
Crittogamologica Italiana, Milan 1:236. 1863.
=Sphaeria (Obtectae) fuscella M.J. Berkeley
& C.E. Broome, Annals and Magazine of
Natural History, London, Series 2, 9:325.
1882. On dead twigs of Rosa sp., Rosaceae.
Great Britain.
=Discostroma fuscella (M.J. Berkeley & C.E.
Broome) S.M. Huhndorf, comb. nov.

L. hazslinszkii P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.

Saccardo 2:28. 1883. [Based on Ditopella sp.,

published by F.A. Hazslinszky,
Verhandlungen der Zoologisch-Botanischen
Gesellschaft in Wien 20:217. 1870.] On dead
branches of Rosa sp., Rosaceae. Rumania.

(n. v.). Type material was not found at B, BP,
F, Fl, L, NBG, UPS, W, WAG. No reply
from IBF, TNP. It is not known whether type
material is at PAD because this institution has
stopped lending specimens until further
notice.

L. (Clypeosphaeria) hendersoniae (J.B. Ellis) M.C.
Cooke, Grevillea, London 17:91. 1889.
=Sphaeria (Obtectae) hendersonia J.B. Ellis
in M.C. Cooke and J.B. Ellis, Grevillea,
London 6:14—15. 1877. =Clypeosphaeria
hendersoniae (J.B. Ellis) P.A. Saccardo,

Leptosphaeria Species on the Rosaceae

an
i)
~

Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:91.
1883. On canes of Rubus sp., Rosaceae.
United States.

=Kalmusia coniothyrium (L. Fuckel)
S.M. Huhndorf, comb. nov.

L. hollosiana L.E. Wehmeyer, Mycologia,
Lancaster, Pennsylvania 55:319. 1963.
=Leptosphaeria maritima L. Hollos, Annales
Historico-Naturales Musei Nationalis
Hungarici 5:46. 1907. Non L. maritima Ellis
& Everhart, Non L. maritima Saccardo. On
stems of Biebersteinia emodii Jaub. & Spach.,
Potentilla argyrophylla Wallich ex Lehm.,
Potentilla fragarioides L., Triglochin
maritima L., Trisetum spicatum (L.) K.
Richter, Geraniaceae, Gramineae, Juncaceae,
Rosaceae. India. (n. v.). Type material was
not found at BP, PAV, PRE.

L. leiostega (J.B. Ellis) J.B. Ellis, Catalogue of
Plants Found in New Jersey. Geological
Survey of New Jersey, Final Report of the
State Geologist, Trenton 2(1):525. 1889.
=Sphaeria (Leptosphaeria) leiostega J.B.
Ellis, Bulletin of the Torrey Botanical Club
(and Torreya), New York 8:91. 1881. On
various dead twigs and limbs of Carya sp.,
Rosa sp., Vaccinium sp., Juglandaceae,
Rosaceae, Ericaceae. United States (New
Jersey).
=Discostroma fuscella (M.J. Berkeley & C.E.
Broome) S.M. Huhndorf, comb. nov.

L. lejostega (J.B. Ellis) F. Hazslinszky, Matematikai
és Természettudomanyi K6zlemenyek
Vonatkozoélag a Hazai Viszonyokra, Budapest
25(2):148. 1892. =Sphaeria (Leptosphaeria)
leiostega J.B. Ellis, Bulletin of the Torrey
Botanical Club (and Torreya), New York
8:91. 1881. =Metasphaeria lejostega (J.B.
Ellis) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:164. 1883. On branches of Carya
sp., Rosa sp., Vaccinium sp., Juglandaceae,
Rosaceae, Ericaceae. United States (New
Jersey).

L. leucoplaca (K.F. Wallroth) E.A. Vainio, Acta
Societatis pro Fauna et Flora Fennica,
Helsingforsiae 49(2):143—144. 1921.
=Verrucaria leucoplaca K.F. Wallroth, Flora
Cryptogamica Germaniae 1:299, 1831.
=Pyrenula leucoplaca (F.G. Wallroth) G.W.
Koerber, Systema Lichenum Germaniae, p.
361. 1855. =Verrucaria alba H.A. Schrader
var. leucoplaca (F.G. Wallroth) L.E. Schaerer
in Enumeratio Critica Lichenum
Europaeorum, Quos Ex Nova Methodo
Digerit, p. 219. 1850. =Eopyrenula leuco-
placa (K.F. Wallroth) R.C. Harris, The
Michigan Botanist, Ann Arbor 12:19. 1973.

Illinois Natural History Survey Bulletin

On bark of Acer sp., Alnus sp., Carya sp.,
Corylus sp., Fraxinus sp., Juglans sp.,
Populus sp., Quercus sp., Salix sp., Sorbus
sp., Tilia sp., Ulmus sp., Aceraceae,
Juglandaceae, Betulaceae, Oliaceae,
Salicaceae, Fagaceae, Rosaceae, Tiliaceae,
Ulmaceae. Finland. (n. v.). No reply from
STR.

L. lucilla P.A. Saccardo, Nuovo Giornale Botanico

Italiano e Bolletino della Societa Botanica
Italiana, Firenze 7:310-311. 1875. Indiscrimi-
nately on wilted leaves of Pyrus communis L.,
Rosaceae. Italy.

=Phaeosphaeria lucilla (P.A. Saccardo) S.M.
Huhndorf, comb. nov.

L. mandshurica M. Miura, Flora of Manchuria and

East Mongolia, Part III, Cryptogams, Fungi
(Contributions of the Southern Manchuria
Railway Company 27):175. 1928. On leaves
of Pyrus malus L. [as Malus domestica
Borkh.], Rosaceae. China. (n. v.). Type
material was not found at TI, TKB. No reply
from TFM, TOFO, TNS.

L. minima (J.E. Duby) ex P.A. Saccardo, Michelia

Commentarium Mycologicum Fungos in
Primis Italicos Illustrans 2:320. 1881.
=Sphaeria minima J.E. Duby in C.
Roumeguere, Fungi Selecti Gallici Exsiccati,
Century 7, No. 694. Anno 1880. Nom. nud.
=Mycotodea minima (J.E. Duby ex P.A.
Saccardo) W. Kirschstein in O.C. Schmidt,
Kryptogamenflora der Mark Brandenburg und
Angrenzender Gebiete herausgegeben von den
Botanishen Verein der Provinz Brandenburg,
Leipzig 7(3):432. 1938. On leaves of
Potentilla argentea L., Rosaceae. France.
Species incertae sedis.

L. miyakeana P.A. Saccardo, Sylloge Fungorum

Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 24:999. 1928. =Phaeosphaeria
eriobotryae 1. Miyake, Botanical Magazine,
Tokyo 27:41-42. 1913. =Trematosphaerella
eriobotryae (1. Miyake) F.L. Tai, Sylloge
Fungorum Sinicorum, p. 330. 1979. On
leaves of Eriobotrya japonica (Thunb.)
Lindley, Rosaceae. China. (n. v.). Type
material was not found at TI, TKB. No reply
from TFM, TOFO, TNS. It is not known
whether type material is at PAD since this
institution has stopped lending specimens
until further notice.

L. mume K. Hara, By6chu-gai Zasschi (Journal of

Plant Protection), Tokyo 5:884. 1918. [As
mune in original publication.] On dead
branch of Prunus mume Sieb. & Zucc.,
Rosaceae. Japan. (n. v.). Type material was
not found at TI, TKB. No reply from TFM,
TOFO, TNS.

Vol. 34 Art. 5

L. nashi (K. Hara) P.A. Saccardo in A. Trotter,

Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 24:999.
1928. =Leptosphaerella nashi K. Hara,
Kajyu-byogai-ron Nihon-kankitsu-Kai,
Schiznoka [Fruit Tree Diseases], pp. 142-145.
1916. =Phaeospora nashi (K. Hara) K. Hara,
Jitsuyo-Sakumotsu-Byorigaku [Manual of
Plant Pathology], p. 270. 1925. On leaves of
Pyrus sinensis Hort., Rosaceae. Japan. (n.
v.). Type material was not found at TI, TKB.
No reply from TFM, TOFO, TNS.

L. (Clypeosphaeria) notarisii M.C. Cooke, Grevillea,

London 17:91. 1889. =Sphaeria clypeiformis
G. de Notaris, Memorie Della Accademia
delle Scienze di Torino, Series 2, No. 7, p.
113. 1853; Micromycetes Italici Novi vel
Minus Cogniti 7, p. 113. 1845. Non Sphaeria
clypeiformis L.V. de Lacroix in G.L.
Rabenhorst. =Clypeosphaeria notarisii L.
Fuckel, Symbolae Mycologicae, p. 117. 1870.
On barky runners of Rubus fruticosus L.,
Rosaceae. Italy.

=Clypeosphaeria mamillana (E.M. Fries:
E.M. Fries) J.B.E. Lambotte.

L. obesula P.A. Saccardo in V. Mouton, Bulletin.

Société R. de Botanique de Belgique,
Bruxelles 26:176. 1887. On stems of Acer
pseudoplatanus L., Angelica sp., Spiraea
ulmaria L. (=Filipendula ulmaria Maxim.),
Aceraceae, Umbelliferae, Rosaceae. Belgium.
(n. v.). Type material was not found at
DUKE. No reply from AUT, CGE, LD. It is
not known whether type material is at PAD
because this institution has stopped lending
specimens until further notice.

L. oligotheca F. Petrak & H. Sydow, Annales

Mycologici, Berlin 22:359. 1924. [Described
from the type material of Laestadia
potentillae E. Rostrup, Botanisk Tidsskrift,
Kjobenhavn 25:300. 1903.] On leaves of
Potentilla maculata Pour. (=Potentilla aurea
L.), Rosaceae. Denmark, Iceland.
=Leptosphaerulina pulchra (H.G. Winter)
M.E. Barr.

L. (Clypeosphaeria) osculanda (C.G.T. Preuss) M.C.

Cooke, Grevillea, London 17:91. 1889.
=Sphaeria osculanda G.T. Preuss, Linnaea.
Institut botanique de |’ Université de Genéve
26:713. 1853. =Clypeosphaeria osculanda
(G.T. Preuss) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:91. 1883. On
branches of Rubus sp., Rosaceae. Germany.
(n. v.). Type material was not found at B.

L. oxyspora J. Feltgen, Vorstudien zu einer Pilz-

Flora des Grossherzogthums, Luxemburg
1(2):165-166. 1901. On dry stems of Rubus
idaeus L., Rosaceae. Luxemburg. (n. v.).

May 1992

Type material is probably at LUX but was not
obtained because the collections are undergo-
ing reorganization.

L. pachytheca P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze, New Series 22:39.
1915. Nom. illeg. Art. 64.1. On dead
branches of Amygdalus communis L.
{=Prunus dulcis (Mill.) D.A. Webb],
Rosaceae. Yugoslavia. (n. v.). It is not
known whether type material is at PAD
because this institution has stopped lending
specimens until further notice.

L. (Leptosphaerella) pomona P.A. Saccardo, Nuovo
Giornale Botanico Italiano e Bolletino della
Societa Botanica Italiana, Firenze 8:176.
1876. On upper surface of leaves of Pyrus
malus L. (=Malus domestica Borkh.),
Rosaceae. Italy.
=Phaeosphaeria pomona (P.A. Saccardo)
S.M. Huhndorf, comb. nov.

L. pomona P.A. Saccardo forma transilvanica A.
Negru et I. Ditzu, Notulae Systematicae e
Sectione Cryptogamica Instituti Botanici
Nomine V.L. Komaroy Academicae
Scientiarum U.R.S.S., Petropolis 16:152-153.
1963. On fruits of Crataegus monogyna
Jacq., Rosaceae. Rumania. (n. v.). Type
material was not found at BUCM. No reply
from BUC, BUCA, BUCF, CL.

L. poterii G. Passerini, Atti dell’ Accademia
Nazionale dei Lincei. Memoire, Rome, Series
4, 6:459. (1889) 1890. On wilted stems of
Poterium sanguisorba L., Rosaceae. Italy.
(n. v.). Type material was not found at
PARMA, PISA, PAD.

L. praetermissa (P.A. Karsten) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:26.
1883. =Sphaeria praetermissa P.A. Karsten,
Fungi Fenniae Exsiccati, Century 9, No. 852.
Anno 1869. On dried canes of Rubus idaeus
L., Rubus odoratus L., Rosaceae. Finland.

L. pruni N.N. Woronichin, Vestnik Tiflisskogo
Botanicheskogo Sada, Tiflis (Moniteur du
Jardin Botanique de Tiflis) 28:17-18. 1913.
On living leaves of Prunus domestica L.,
Rosaceae. U.S.S.R. (Caucasus). (n. v.).
Type material was not found at LE, LEP, TBI.
No reply from TGM.

L. pulchra (H.G. Winter) P.A. Saccardo, Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:53—54. 1883.
=Sphaerella pulchra H.G. Winter, Hedwigia,
Dresden 11:145-146. 1872. =Mycodotea
pulchra (H.G. Winter) W. Kirschstein,
Annales Mycologici, Berlin 34:201. 1936;
Kryptogamenflora de Mark Brandenburg und
Angrenzender Gebiete herausgegeben von den

Leptosphaeria Species on the Rosaceae

529

Botanischen Verein der Provinz Brandenburg,
Leipzig 7(3):433. 1938. On dry stems and
leaves of Potentilla caulescens L., Rosaceae.
Austria. Type material was not found at B,
CORD, IBF, JE, LZ, WRSL. No reply from
FI, IB, TO, WAG.

=Leptosphaerulina pulchra (H.G. Winter)
M.E. Barr, Contributions de |’Institut
Botanique de L’ Université de Montréal 73:7.
1959.

L. puttemansti A. Maublanc, Bulletin de la Société
Mycologique de France, Paris 21:88-89.
1905. On leaves of Eriobotrya japonica
(Thunb.) Lindley, Rosaceae. Brazil. (n. v.).
Type material was not found at IBI, PC, RSA-
POM, SP. No reply from RBR.

L. rimicola (G.H. Otth) P.A. Saccardo, Hedwigia,
Dresden 35:XXIX. 1896; Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 11:XXIX. 1896. =Cladosphaeria
rimicola G.H. Otth, Mitteilungen der
Naturforschenden Gesellschaft in Bern
1870:106. 1871. Nom inval. Art. 43.1. [The
genus Cladosphaeria was validated in 1894.]
On fallen branches of Prunus avium L.,
Rosaceae. Switzerland.
=Melanomma pulvis-pyrius (C.H.
Persoon:E.M. Fries) L. Fuckel.

L. rostrupti P.A. Saccardo & D. Saccardo in P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
17:721. 1905. =Leptosphaeria dryadis E.
Rostrup. Non Rostrup 1903. On upper surface
of dried leaves of Dryas octopetala L.,
Rosaceae. Norway. (n. v.). It is not known
whether type material is at PAD since this
institution has stopped lending specimens
until further notice.

L. rustica P.A. Karsten, Fungi Fenniae Exsiccati,
Century 10, No. 964. Anno 1870.
=Metasphaeria rustica (P.A. Karsten) P.A.
Saccardo, Sylloge Fungorum Omnium
Hucusque Cognitorum Digessit P.A. Saccardo
2:157. 1883. On stems of Spiraea ulmaria L.
(=Filipendula ulmaria Maxim.), Rosaceae.
Finland.
=Leptosphaeria doliolum (C.H. Persoon:E.M.
Fries) V. Cesati & G. de Notaris.

L. saccardiana J.H. Fabre, Annales des Sciences
Naturelles, Paris, Botanique, Series 6, 9:89.
1878. =Requienella saccardiana (J.H. Fabre)
J.H. Fabre, Annales des Sciences Naturelles,
Paris, Botanique, Series 6, 15:56. 1883. On
dried stems of Crataegus oxyacantha L.,
Rosaceae. France. (n. v.). Type material was
not found at PC. No reply from L’HARMAS.

L. sanguisorbae P.A. Karsten in A.N. Berlese,
Icones Fungorum Omnium Hucusque
Cognitorum ad usum Sylloges Saccardianae

Illinois Natural History Survey Bulletin Vol. 34 Art. 5

Adcommodatae 1:78. 1892. On branches of
Sanguisorba officinalis L., Rosaceae.
Germany. (n. v.). Type material was not
found at H, UPS.

L. sanguisorbae P.A. Karsten, Enumeratio

Fungorum et Myxomycetum in Lapponia
Orientali Aestate 1861 Lectorum, p. 214.
1886; Notiser Sallskapets Pro Fauna et Flora
Fordhandlingar 8:214. 1866. [As Sphaeria
(Leptosphaeria) sangvisorbae.] On dead
stems of Sanguisorba polygama F. Nyl.,
Rosaceae. U.S.S.R. (n. v.).

L. sepincola (E.M. Fries:E.M. Fries) H.G. Winter,

Dr. L. Rabenhorst’s Kryptogaman-Flora von
Deutschland, Oesterreich und der Schweiz,
Second edition, 1(2):473. 1885. [Sphaeria
sepincola E.M. Fries, Observationes
Mycologicae. Seu Descriptiones tam
Novarum, quam Notabilium Fungorum
Exhibitae a C.H. Persoon, Lipsiae 1:181.
1815; Systema Mycologicum Sistens
Fungorum 2:498. 1823. [As saepincola.]
=Pringsheimia sepincola (E.M. Fries:E.M.
Fries) F. v. Hoehnel, Annales Mycologici,
Berlin 18:97. 1920. =Metasphaeria sepincola
(E.M. Fries:E.M. Fries) P.A. Saccardo,
Sylloge Fungorum Omnium Hucusque
Cognitorum Digessit P.A. Saccardo 2:164.
1883. [As Fr? Fuckel.] =Sphaerulina
sepincola (E.M. Fries:E.M. Fries) K.Starback,
Botaniska Notiser, Lund 1890:117. 1890;
Botanisches Zentralblatt, Jena & Dresden
46:261. 1891. =Pleosphaerulina sepincola
(E.M. Fries:E.M. Fries) H. Rehm in F. v.
Hoehnel, Annales Mycologici, Berlin 18:96.
1920. =Sclerodothis sepincola (E.M.
Fries:E.M. Fries) F. Petrak, Annales
Mycologici, Berlin 19:41. 1921. On dry
stems of Rosa sp., Rubus sp., Rosaceae.
Sweden.

=Saccothecium sepincola (E.M. Fries:E.M.
Fries) E.M. Fries, Summa Vegetabilium
Scandinaviae, Seu Enumeratio, Systematica et
Critica, Plantarum tum Cotyledonearum,
Quam Nemearum Inter Mare Occidentale Et
Album, Inter Eidoram et Nordkop, Hactenus
Lectorum, una Cum Singulae Distributione
Geographica, p. 398. 1849.

L. sieversiae (F.E. Clements) F. Petrak, Sydowia.

Annales Mycologici, Horn, N.O. 6(1-4):6.
1952. =Massaria sieversiae F.E. Clements,
Cryptogamae Formationum Coloradensium,
Century III, No. 234. Anno 1906. Nom. nud.
Art. 36.1. [Combination is not valid because
the basionym is an invalidly published
species.] On standing, dead stems of

Sieversia turbinata (Rydb.) Greene, Rosaceae.

U.S.A.
=Diadema sieversiae (C.H. Peck) S.M.
Huhndorf, comb. nov.

L. sorbi A. Jaczewski, Annales Mycologici, Berlin
1:30. 1903. On leaves of Sorbus aucuparia
L., Rosaceae. U.S.S.R.

Species incertae sedis.

L. spiraeae P.A. Karsten, Hedwigia, Dresden
27:260. 1888. On dry fallen branches of
Spiraea sorbifolia L. (=Sorbaria sorbifolia A.
Br.), Rosaceae. Finland. (n. v.). Type
material was not found at H, UPS.

L. subcutanea (M.C. Cooke & J.B. Ellis) J.B. Ellis in
N.L. Britton, Catalogue of Plants Found in
New Jersey. Geological Survey of New
Jersey, Final Report of the State Geologist,
Trenton 2(1):525. 1889. =Sphaeria
(subtectae) subcutanea M.C. Cooke & J.B.
Ellis, Grevillea, London 7:41. 1878.
=Metasphaeria subcutanea (M.C. Cooke &
J.B. Ellis) P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 2:167. 1883. On decorticated limbs
of Pyrus communis L., Rosaceae. United
States. (New Jersey).
=Lophiostoma subcutanea (M.C. Cooke &
J.B. Ellis) S.M. Huhndorf, comb. nov.

L. superficialis F.A. Hazslinszky, Matematikai és
Természettudomanyi Kozlemenyek
Vonatkozolag a Hazai Viszonyokra, Budapest
25(2):146. 1892. On unknown substrate of
Potentilla argentea L., Rosaceae. Hungary.
(n. v.). Type material was not found at B, BP,
F, FI, L, NBG, UPS, W, WAG. No reply
from IBF, TNP.

L. ternata F. Hazslinszky, Matematikai és
Természettudomanyi K6zlemenyek
Vonatkozolag a Hazai Viszonyokra, Budapest
25(2):150. 1892. On branches of Pyrus malus
L. (=Malus domestica Borkh.), Rosaceae.
Hungary. (n. v.). Type material was not
found at B, BP, F, FI, L, NBG, UPS, W,
WAG. No reply from IBF, TNP.

L. thomasiana P.A. Saccardo & C. Roumeguére,
Revue Mycologique, Toulouse 5:236. 1883.
On dead runners of Rubus sp., Rosaceae.
France. It is not known whether type material
is at PAD because this institution has stopped
lending specimens until further notice.
=Phaeosphaeria thomasiana (P.A. Saccardo
& C. Roumegueére) S.M. Huhndorf, comb.
nov.

L. umbrosa G. Niessl in G.L. Rabenhorst, Fungi
Europaei Exsiccati, Klotschii Herbarii vivi
Mycologici Continuatio, Edition 3 (Edita
Nova), Series 2, Century 20, No. 1934. Anno
1875; Just’s Botanisch Jahresberichte, Berlin
3:262. 1887. =Massaria umbrosa (G. Niessl)
H. Rehm in P.A. Saccardo, Sylloge Fungorum
Omnium Hucusque Cognitorum Digessit P.A.
Saccardo 9:761. 1891. On unknown substrate
of Actaea spicata L., Astrantia major L.,

May 1992 Leptosphaeria Species on the Rosaceae

Gentiana asclepiadea L., Solanum dulcamara
L., Spiraea aruncus L. (=Aruncus dioicus
Fern.), Ranunculaceae, Umbelliferae,
Gentianaceae, Solanaceae, Rosaceae. Austria.

L. vagabunda P.A. Saccardo, Nuovo Giornale
Botanico Italiano e Bolletino della Societa
Botanica Italiana, Firenze 7:318. 1875.
=Sphaeria fuscella P.A. Saccardo, Atti dell’
Accademia Scientifica Veneto-Trentino-
Istriana, Padova 2(1):145. 1873. [As
Sphaeria fuscella M.J. Berkeley & C.E.
Broome. Non Leptosphaeria fuscella (M.J.
Berkeley & C.E. Broome) V. Cesati & G. de
Notaris.] On branches of Abies excelsa
(Lam.) Poiret, Acer campestre L., Alnus
glutinosa (L.) Gaertner, Ampelopsis
heterophylla (Thunb.) Sieb. & Zucc.,
Clematis vitalba L., Cornus sanguinea L.,
Corylus avellana L., Dulcamara sp.,
Hypericum calycinum L., Kerria japonica (L.)
DC., Quercus pedunculata Ehrh., Salix
purpurea L., Pinaceae, Aceraceae,
Betulaceae, Vitaceae, Ranunculaceae,
Cornaceae, Solanaceae, Guttiferae, Rosaceae,
Fagaceae, Salicaceae. Italy. (n.v.). It is not
known whether type material is at PAD
because this institution has stopped lending
specimens until further notice.

L. vagabunda P.A. Saccardo var. divergens P.A.
Saccardo, Fungi Italici autographice delineati
(additis nonnullis extra-italicis asterisco
notatis), Patavii, No. 518. 1879; Sylloge
Fungorum Omnium Hucusque Cognitorum
Digessit P.A. Saccardo 2:31. 1883. On
branches of Kerria sp., Rosaceae. Italy. (n.
y.). It is not known whether type material is
at PAD because this institution has stopped
lending specimens until further notice.

L. vagabunda P.A. Saccardo var. sarmenti P.A.
Saccardo, Fungi Italici autographice delineati
(additis nonnullis extra-italicis asterisco
notatis), Patavii, No. 428. 1879; Sylloge
Fungorum Omnium Hucusque Congitorum
Digessit P.A. Saccardo 2:31. 1883. On
runners of Rubus sp., Rosaceae. Italy. (n. v.).
It is not known whether type material is at
PAD because this institution has stopped
lending specimens until further notice.

nn
Ww

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Mycologia 75:1—13.

Barr, M.E. 1987a. Prodromus to class
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Melanomma pulvis-pyrius (Pers.) Fuckel,
Melanomma fuscidulum Sacc. and Thyridaria
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Index to Taxa

abbreviata 510,526
aculeorum 526

arunci 498, 499, 526
cercocarpi 490, 491, 526
clivensis 498, 499
Clypeosphaeria 518
concentrica 492, 493, 526
coniothyrium 500, 501, 526
controversa 520, 521, 526
corticola 520,522, 526
Diadema_ 513,514,515, 516
Diaporthe 520
Discostroma 520
doliolum 483, 484
dryadis 484, 485, 527
dryadophila 484, 485, 527
eres 520,521

eriobotryae 525,527
fuscella 520, 522, 527
galiorum var. lapsanae 498
hazslinszkii 527
hendersonia 500, 501, 527
hollosiana 527

Kalmusia 498, 499, 500, 501
leiostega 520, 522, 527
lejostega 521,527
Leptosphaerulina 510, 512
leucoplaca 527
longipedicellata 498, 499
Lophiostoma 503

lucilla 494, 495, 528
mamillana 518,519
mandshurica 528
Melanomma 505, 506
minima 525, 528
miyakeana 528

mume 528

nashi 528

notaristi 518,519, 528
obesula 528

obtusa 513,514
oligotheca 511,512, 528
osculanda 528

oxyspora 528

pachytheca 529
Paraphaeosphaeria 492, 493
Phaeosphaeria 492, 494, 495
somona 492, 494, 529

pomona forma transilvanica 529
poterti 529

praetermissa 486, 487, 529
pruni 529

pulchra 510,512, 529
pulvis-pyrius 505, 506
puttemansti 529

rimicola 505, 506, 529
rostrupit 529

rustica 483, 484, 529
saccardiana 529
Saccothecium 508, 509, 510, 511
sanguisorbae 529, 530
sepincola 508, 509, 530
sepincola var. abbreviata 510, 511
sieversiae 515,516, 530

sorbi 525,530

spiraeae 530

steironematis 498

subcutanea 503, 504, 530
superficialis 530

ternata 530

thomasiana 496, 497, 530
umbrosa_ 488, 489, 530
vagabunda 531

vagabunda var. divergens 531
vagabunda var. sarmenti 531

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Catalog of Types of the Illinois
Natural History Survey
Mycological Collections (ILLS)

J.L. Crane and Pamela P.

Illinois Natural History Survey Bulletin
Volume 34, Article 6
May 1992

ILLINOIS
NATURAL
HISTORY
SURVEY

Tazik

MAY 10 1994

LIBRARY.

DEPOSITORY
aon 2 i IQG/

UNIVERSITY OF ILL INOIS
AT URBANA-CHAMPAIGC

ILLINOIS
NATURAL
HISTORY
SURVEY

Catalog of Types of the Illinois
Natural History Survey
Mycological Collections (ILLS)

ie. Crane

Center for Biodiversity
Illinois Natural History Survey

Pamela P. Tazik

Center for Aquatic Ecology
Illinois Natural History Survey

Illinois Natural History Survey Bulletin
Volume 34, Article 6
May 1992

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

A catalog of the publications of the Illinois Natural History Survey is available
without charge from the address below. A price list and an order blank are
included with the catalog.

Illinois Natural History Survey
Distribution Center

Natural Resources Building
607 East Peabody Drive
Champaign, Illinois 61820

Citation:

Crane, J.L., and P.P. Tazik. 1992. Catalog of types of the Illinois Natural
History Survey mycological collections (ILLS). Bulletin 34(6):535—550.
Editor: Eva L. Steger

P.P. Tazik’s current address: Hazardous Waste Research and Information
Center, 1 East Hazelwood Drive, Champaign, IL 61820.

US ISSN 0073-4918

Printed by Authority of the State of Illinois
(X0415-M-5-92)

Contents

Introduction 535
Catalog of Types 536
Literature Cited 550
Appendix 550

Introduction

The mycological collections of the Illinois Nat-
ural History Survey and the University of Illinois
at Urbana-Champaign originated with the rust
collection of A.B. Seymour (1881-1886) and the
powdery mildew collection of T.J. Burrill (1882—
1885). These collections were integrated and
housed in the Natural History Building on the
campus of the University of Illinois at Urbana-
Champaign until 1921, when the plant disease
specimens were segregated and became the basis
for the plant disease collection of the Illinois
Natural History Survey. The program for col-
lecting, identifying, and preserving plant disease
specimens originated in 1881 and was expanded
between 1921 and 1924 when special emphasis
was placed on obtaining information on plant dis-
ease in the state. By 1924, 18,000 plant disease
specimens had been added to the collection.

Gilbert L. Stout (1926-1930) was the first
plant pathologist to concentrate on surveying
plant disease in Illinois; he was succeeded by
Gideon H. Boewe (1930-1966). Boewe’s inter-
est was the incidence, distribution, and severity
of plant diseases. His specimens, together with
those of Leo R. Tehon (1921-1954), who special-
ized in Ascomycetes and Fungi Imperfecti that
cause plant disease, form a substantial part of
the Survey’s mycological collection.

James C. Carter (1934-1974) was an
authority on the diseases of shade trees and
ornamentals. His special interest was patho-
genic fungi of woody ornamentals and he
contributed numerous specimens to the
mycological collection. In 1967, J. Leland
Crane succeeded Boewe as mycologist, and to
date he has contributed over 4,000 specimens
of Ascomycetes and Fungi Imperfecti from
decaying substrates in aquatic systems.

The Survey’s mycological collection now
contains approximately 300 type specimens,
800 Myxomycetes, 1,200 lower fungi, 10,000
Ascomycetes, 13,200 Fungi Imperfecti, 17,000
Basidiomycetes, and 5,000 lichens, for a total
of 47,500 specimens.

This catalog provides information for
each type specimen on the binomial, the full

name and initials of the authorities, the place of
publication, the category of type, substrate or
host, locality data, date of collection, collector,
and accession number. The categories of type
specimens follow the Botanical Code (Grueter
et al. 1988, Article 7). In this code, a nomen-
clatural type is that element to which the name
of a taxon is permanently attached. A holotype
is the one specimen or illustration used or
designated by the author as the nomenclatural
type. An isotype is any duplicate of the
holotype. A syntype is any one of two or more
specimens cited by the author when no holo-
type was designated, or when any one of two or
more specimens was simultaneously designated
as types. A paratype is a specimen or illustra-
tion cited in the protologue that is neither
holotype or isotype, nor one of the syntypes if
two or more specimens were simultaneously
designated as types. A lectotype is a specimen
or illustration selected from the original
material to serve as a nomenclatural type when
no holotype was indicated at the time of
publication, or as long as it is missing. A
neotype is a specimen or illustration selected to
serve as nomenclatural type as long as all of the
material on which the name of a taxon was
based is missing.

Standardized, complete names of periodi-
cals are cited following the system of Brown
and Stratton (1963) and Porter and Koster
(1970). The titles of books and pamphlets are
cited according to Stafleu and Cowan (1976—
1988). The scientific names of host plants are
given as reported in the protologue of each type
species. Plant genera were verified in Willis
(1973) and Farr et al. (1979). Specific epithets
of vascular plants were confirmed in Fernald
(1950), Bailey and Bailey (1976), and Little
(1979). The geographic location of species is
restricted to information on the type specimen.
Type specimens missing from the mycological
collections are listed in the Appendix.

We express our sincere appreciation to
Betty A. Nelson for technical assistance in the
preparation of this manuscript.

Catalog of Types

Acrodictys martinii J.L. Crane & K.P. Dumont,
Canadian Journal of Botany 53:846. 1975.
Isotype: On rotted bark and wood along Rio
Nueve Pasos, Dr. Luis Roure’s property near
Rosario, Puerto Rico, elevation 140 m,

17.V1.1970, leg. R.P. Korf et al., ILLS 35534.

Holotype: NY.

Acrodontium myxomyceticola J.L. Crane & J.D.
Schoknecht, Transactions of the British
Mycological Society 79:346. 1982. Isotype:
On Stemonites fusca A.W. Roth var. fusca,
Territorio de Roraima, ca. 219 km north of
Boa Vista, on the Boa Vista-Sta. Elena
Venezuela Rd. (BR 174), Brazil, 1.X1I.1977,
leg. K.P. Dumont et al. BR-965, ILLS 42576.
Holotype: NY.

Actinospora jamaicensis J.L. Crane & K.P. Dumont,

Canadian Journal of Botany 53:843. 1975.
Isotype: Ex CUP-MJ 128, on wet wood,
along trail to Silver Hill Gap, near
Woodcutter’s Gap, vicinity of Newcastle,
Portland Parish, Jamaica, 9.1.1971, leg. R.P.
Korf et al., ILLS 35494. Holotype: NY.

Aniptodera chesapeakensis C.A. Shearer & M.A.
Miller, Mycologia 69:894. 1977. Holotype:
A dried culture, isolated from balsa wood
submerged in the intake canal of the Potomac
Electric Power Company, Electricity
Generating Plant, Chalk Point, Maryland,
23.V.1973, leg. C.A. Shearer CS97-26, ILLS
36523. Culture from type: ATCC 32818.

Aristastoma concentrica L.R. Tehon, Mycologia
25:249. 1933. Holotype: On leaves of Vigna
sinensis (L.) G. Engelmann, Metropolis,
Massac County, Illinois, 12.X.1927, leg. G.L.
Stout, ILLS 5453.

Ascochyta biguttulata E.Y. Daniels in L.R. Tehon
and E.Y. Daniels, Mycologia 19:125. 1927.
Holotype: On Polygonum convolvulus L.,
Urbana, Champaign County, Illinois,
11.X.1925, leg. E. Daniels, ILLS 2438.

Ascochyta elymi L.R. Tehon & E.Y. Daniels,
Mycologia 19:124-125. 1927. Holotype: On
Elymus virginicus L., Jacksonville, Morgan
County, Illinois, 25.VII.1925, leg. L.R.
Tehon, ILLS 3334.

Ascochyta maydis G.L. Stout, Mycologia 22:27 1—
272. 1930. Holotype: On leaves of Zea mays
L., Macomb, McDonough County, Illinois,

11.X.1926, leg. G.L. Stout, ILLS 19688.
Paratype: Percy, Randolph County, Illinois,
9.X1.1927, leg. G.L. Stout, ILLS 21204.

Ascochyta negundinis L.R. Tehon, Mycologia
29:442-443. 1937. Holotype: On Acer
negundo L., Mt. Carroll, Carroll County,
Illinois, 22.VI.1935, leg. G-H. Boewe, ILLS
25193.

Ascochyta plantaginella L.R. Tehon, Mycologia
25:247. 1933. Holotype: On living leaves of
Plantago rugelii J. Decaisne, Homer,
Champaign County, Illinois, 6.VI.1929, leg.
G.L. Stout, ILLS 22014.

Ascochyta rhodotypi H.W. Anderson, Transactions
of the Illinois State Academy of Science
15:129. 1922. Holotype: On leaves of
Rhodotypos scandens (C.P. Thunberg) T.
Makino [as R. kerrioides P.F. von Siebold &
J.G. Zuccarini in protologue], University of
Illinois campus, Urbana, Champaign County,
Illinois, 10.1X.1919 [as 15.IX.1919 in
protologue], leg. H.W. Anderson, ILLS 4726.

Ascochyta zeae G.L. Stout, Mycologia 22:272.
1930. Holotype: On leaves of Zea mays L..
Mt. Carmel, Wabash County, Illinois,
9.X1.1926, leg. G.L. Stout, ILLS 19581.

Asperisporium acori L.R. Tehon, Mycologia
40:32 1-322. 1948. Holotype: On Acorus
calamus L., Urbana, Champaign County,
Illinois, 17.VI.1926, leg. W.G. Solheim, ILLS
30124.

Brachysporiella dennisii J.L. Crane & K.P. Dumont,
Canadian Journal of Botany 56:2613. 1978.
Isotype: On unidentified stem, path between
hotel and hotel’s water source, Rancho
Grande, Parq. Nac. Henry Pittier, Edo.
Aragua, Venezuela, 3. VII.1971, leg. K.P.
Dumont, J.H. Haines, & G.J. Samuels VE-
1174, ILLS 36938, VEN. Holotype: NY.

Camposporium marylandicum C.A. Shearer,
Mycologia 66:16. 1974. Holotype: On balsa
wood [Ochroma pyramidale (A.J. Cavanilles)
I. Urban] submerged in the Patuxent River at
the Wildlife Refuge, Laurel, Maryland,
22.VII.1969 [as 22.V.1969 in protologue],
leg. C.A. Shearer, ILLS 35538. Paratype:
ILLS 35591.

May 1992

Carpenterella molinea L.R. Tehon & H.A. Harris,
Mycologia 33:128. 1941. Holotype: On
Ulmus americana L., Madison, Wisconsin,
4.V.1932, leg. E.L. Champers, Wisconsin
Department of Agriculture & Markets, ILLS
22722.

Catosphaeropsis caulivora L.R. Tehon, Mycologia
31:542. 1939. Holotype: On Lespedeza
stipulacea C.J. Maximowicz, Crossville,
White County, Illinois, 22. VII.1937, leg. G.H.
Boewe, ILLS 26978. Paratype: Metropolis,
Massac County, Illinois, 9.IX.1937, leg. G.H.
Boewe, ILLS 26979.

Cercophora septentrionalis N. Lundqvist, Symbolae
Botanicae Upsalienses 20(1):100. 1972.
Paratype: Sweden, Gotland: Gerum par., 2
km E. of Hejdes in pine forest, on horse dung
in moist chamber, Uppsala, 9.VI.1959, leg. N.
Lundqvist 2126-e, ILLS 35156.

Cercospora abutilonis L.R. Tehon & E.Y. Daniels,
Mycologia 17:246. 1925. Holotype: On
Abutilon theophrasti F.C. Medicus, Spring
Valley, Bureau County, Illinois, 17.VIII.1922,
leg. C.L. Porter, ILLS 963.

Cercospora arborescentis L.R. Tehon & E.Y.
Daniels, Mycologia 17:246. 1925. Holotype:
On leaves of Hydrangea arborescens L.,
Thebes, Alexander County, Illinois,

17. VIII.1922, leg. P.A. Young, ILLS 599.

Cercospora cercidicola J.B. Ellis var. coremioideas
L.R. Tehon, Mycologia 16:140. 1924.
Holotype: On Cercis canadensis L., Boaz,
Massac County, Illinois, 8. VIII.1922, leg.
P.A. Young, ILLS 1129.

Cercospora difformis L.R. Tehon, Mycologia
40:322-323. 1948. Holotype: On Viola sp.,
wild, Vandalia, Fayette County, Illinois,
11.X.1944, leg. G.H. Boewe, ILLS 30224.

Cercospora hemerocallidis L.R. Tehon, Mycologia
16:139. 1924, [as hemerocallis]. Holotype:
On Hemerocallis fulva L., Bloomfield,
Johnson County, Illinois, 25. VII.1922, leg.
P.A. Young, ILLS 2897.

Cercospora hyperici L.R. Tehon & E.Y. Daniels,
Mycologia 19:127—128. 1927. Holotype: On
Hypericum adpressum B.S. Barton, Bement,
Piatt County, Illinois, 6. VII.1925, leg. L.R.
Tehon, ILLS 13035.

Cercospora menthicola L.R. Tehon & E.Y. Daniels,
Mycologia 17:247. 1925. Holotype: On
leaves of Mentha canadensis L., Vandalia,
Fayette County, Illinois, 14.VII.1924, leg.
P.A. Young, ILLS 13699,

Cercospora nepetae L.R. Tehon, Mycologia 16:140.
1924. Holotype: On Nepeta cataria L., Ullin,
Pulaski County, [llinois, 11.VIII.1922, leg.
P.A. Young, ILLS 1419.

Cercospora paeoniae L.R. Tehon & E.Y. Daniels,
Mycologia 17:247. 1925. Holotype: On
leaves of Paeonia officinalis L., Prairie du

Catalog of Types 537

Rocher, Randolph County, Illinois,
24. VIII.1922, leg. P.A. Young, ILLS 5645.

Cercospora plantaginella L.R. Tehon, Mycologia
16:139. 1924. Holotype: On Plantago rugelii
J. Decaisne, Boaz, Massac County, Illinois,
8.VIIL.1922, leg. P.A. Young, ILLS 1140.

Cercospora podophylli L.R. Tehon & E.Y. Daniels,
Mycologia 19:128. 1927. Holotype: On
Podophyllum peltatum L., Jersey County,
Illinois, 2. VIII.1922, leg. C.O. Peake, ILLS
12950.

Cercospora psedericola L.R. Tehon, Mycologia
16:139. 1924. Holotype: On Parthenocissus
quinquefolia (L.) J.E. Planchon, [as Psedera],
Buckner, Franklin County, Illinois,
20.VII.1922, leg. PA. Young, ILLS 2987.

Cercospora rhapontici L.R. Tehon & E.Y. Daniels,
Mycologia 17:248. 1925. Holotype: On
leaves of Rheum rhaponticum L., Coxeyville,
Monroe County, Illinois, 24. VIII. 1922, leg.
P.A. Young, ILLS 5111.

Cercospora saccharini A.E. Liberta & G.H. Boewe,
Mycologia 52:345—347. 1960. Holotype: On
Acer saccharinum L., Metropolis, Massac
County, Illinois, 15.X.1959, leg. G.H. Boewe,
ILLS 33219.

Cercospora setariicola L.R. Tehon & E.Y. Daniels,
Mycologia 19:128-129. 1927. Holotype: On
Setaria glauca (L.) A. Beauvois, Macomb,
McDonough County, Illinois, 16. VIII.1924,
leg. P.A. Young, ILLS 11542. Paratype:
Oquawka, Henderson County, Illinois,
15.1X.1924, leg. P.A. Young, ILLS 7905.

Cercospora silphii J.B. Ellis & B.M. Everhart var.
laciniati L.R. Tehon & E.Y. Daniels,
Mycologia 19:128. 1927. Holotype: On
Silphium laciniatum L., Bement, Piatt County,
lilinois, 6.VII.1925, leg. L.R. Tehon, ILLS
15262.

Cercospora sororiae L.R. Tehon, Mycologia
40:323-324. 1948. Holotype: On leaves of
Viola sororia K.L. Willdenow, Kinderhook,
Pike County, Illinois, 28.VI.1933, leg. G.H.
Boewe, ILLS 30126.

Cercospora vignicaulis L.R. Tehon, Mycologia
29:436-437. 1937. Holotype: On Vigna
sinensis (L.) G. Engelmann, Equality, Gallatin
County, Illinois, leg. G.H. Boewe, 8.1X.1932,
ILLS 23703 (with Glomerella vignicaulis).

Cercospora viminei L.R. Tehon, Mycologia 16:141.
1924. Holotype: On Aster vimineus J.B.
Lamarck, Waltonville, Jefferson County,
Illinois, 23.VI.1922, leg. P.A. Young, ILLS
2600.

Cercospora zeae-maydis L.R. Tehon & E.Y.
Daniels, Mycologia 17:248. 1925. Holotype:
On leaves of Zea mays L., McClure,
Alexander County, Illinois, 29. VIII.1924, leg.
P.A. Young, ILLS 4276.

538 Illinois Natural History Survey Bulletin

Chaetomella tritici L.R. Tehon & E.Y. Daniels,
Mycologia 17:242. 1925. Holotype: On inner
surface of glumes of Triticum aestivum L.,
Waterloo, Monroe County, Illinois,
24.VIII.1922, leg. P.A. Young, ILLS 971.

Chaetopsina ludovicina J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
60:372. 1982. Holotype: On submerged,
decayed? magnolia leaf, freshwater roadside
swamp along Louisiana Route 51 about 20
miles north of New Orleans, St. John Baptist
Parish, Louisiana, 1.VIII.1974, leg. J.-L. Crane
& J.D. Schoknecht, ILLS 40844.

Chaetoseptoria vignae L.R. Tehon, Mycologia
29:444. 1937. Holotype: On Vigna sinensis
(L.) G. Engelmann, Eldorado, Saline County,
Illinois, 10.1X.1934, leg. G-H. Boewe, ILLS
25080.

Chaetosphaeria aspergilloides M.E. Barr & J.L.
Crane, Canadian Journal of Botany 57:835—
836. 1979. Isotype: On decayed woody
material, tropical greenhouse, Univ. Nac.
Auton. de México, Distrito Federal, México,
7.1X.1972, leg. J.M. Trappe 3462, ILLS
37868. Holotype: NY.

Cladosporium calotropidis F.L. Stevens, Transac-
tions of the Illinois State Academy of Science
10:207. 1917. Isotype: On Calotropis
procera (W. Aiton) W.T. Aiton f., Guayanilla,
Puerto Rico, VII.1915, leg. F.L. Stevens 9130,
ILLS 41985. Holotype: BPI 70873.

Clavatospora stellatacula P. Kirk, Mycologia
61:178. 1969. Isotype: Isolated from
submerged wood (Pinus ponderosa D.
Douglas ex C. Lawson) in Chesapeake Bay,
Fort Wood, Hampton, Virginia, leg. P. Kirk,
VPI F-83, ILLS 34567. Holotype: P. Kirk
68-137, NY.

Clithris leucothoicola L.R. Tehon, Mycologia
31:676-677. 1939. Holotype: On Leucothoe
catesbei (T. Walter) A. Gray [=L. axillaris
(A.B. Lambert) D. Don], Stroudsburg,
Pennsylvania, 24.V.1931, leg. B.H. Davis,
ILLS 25091.

Codinaea gonytrichodes C.A. Shearer & J.L. Crane,
Mycologia 63:245-247. 1971. Holotype: On
fruit of Carya sp., cypress swamp, northwest
of Karnak, Pulaski County, Illinois,
22.V.1969, leg. J.L. Crane 84-69, ILLS
34867. Paratypes: On balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban]
submerged in Patuxent River at Naval
Ordinance Laboratory Acoustics Facility,
Brighton, Maryland, 22.1X.1967, C.A.
Shearer & J.L. Crane, ILLS 34467; on
samaras of Acer sp. and other decayed plant
material, cypress swamp northwest of Karnak,
Pulaski County, Illinois, 22.V.1969, leg. C.A.

Vol. 34 Art. 6

Shearer and J.L. Crane 94-69, ILLS 34868;
saprophytic on seeds of Liriodendron
tulipifera L., Forman Cypress Swamp,
northwest of Belknap, Johnson County,
Illinois, 18.X1.1969, C.A. Shearer & J.L.
Crane 201-69, ILLS 34869.

Codinaea illinoensis A.D. Hewings & J.L. Crane,
Mycotaxon. An International Journal
Designed to Expedite Publication of Research
on Taxonomy & Nomenclature of Fungi &
Lichens 13:419-421. 1981. Holotype: On
decayed wood, cypress swamp, northwest of
Vienna, Johnson County, Illinois, 9. VII.1969.
leg. J.L. Crane 98-69, ILLS 42169.

Codinaea matsushimae A.D. Hewings & J.L. Crane,
Mycotaxon. An International Journal
Designed to Expedite Publication of Research
on Taxonomy & Nomenclature of Fungi &
Lichens 13:423-425. 1981. Holotype: On
exocarp of Juglandaceae (Carya sp.),
Benson's Bluff, southeast of Goreville,
Johnson County, Illinois, 4.1V.1969, leg. J.L.
Crane 12-1-69, ILLS 42170. Paratype: On
decayed wood, Elvira Cypress Swamp (Deer
Pond), Johnson County, Illinois, 22 May
1969, leg. J.L. Crane, ILLS 42171.

Colletotrichum aeciicolum L.R. Tehon, Mycologia
25:254. 1933. Holotype: On unruptured
peridia of aecidia of Puccinia asterum (L.D.
von Schweinitz) F.D. Ker, in leaves of
Solidago canadensis L., Beechville, Calhoun
County, Illinois, 16.[X.1926, leg. G.L. Stout,
ILLS 22416.

Colletotrichum dioscoreae L.R. Tehon, Mycologia
25:255. 1933. Holotype: On Dioscorea
villosa L., Marlow, Jefferson County, Illinois,
7.1X.1926, leg. G.L. Stout, ILLS 22420.

Colletotrichum smilacinae L.R. Tehon & E.Y.
Daniels, Mycologia 17:245-246. 1925.
Holotype: On leaves of Smilacina racemosa
(L.) R.L. Desfontaines, Goreville, Johnson
County, Illinois, 22.V1.1924, leg. P.A. Young,
ILLS 7259.

Colletotrichum smilacis L.R. Tehon, Mycologia
25:254-255. 1933. Holotype: On living
leaves of Smilax hispida G. Muhlenberg,
Olney, Richland County, Illinois, 21.0X.1932,
leg. G.L. Stout, ILLS 22849.

Colletotrichum trillii L.R. Tehon, Mycologia 16:141.
1924. Holotype: On Trillium recurvatum
Beck, Ryder, Jefferson County, Illinois,
23.VI.1922, leg. P.A. Young, ILLS 2350.

Confertopeltis asparagi L.R. Tehon, Mycologia
25:251. 1933. Holotype: On dry stems of
Asparagus officinalis L., Villa Ridge, Pulaski
County, Illinois, 10.X1.1927, leg. G.L. Stout,
ILLS 7295.

May 1992

Conioscypha varia C.A. Shearer, Mycologia 65:133.
1973. Holotype: A dried colony on cantino
peptone yeast glucose agar, isolated from
balsa wood [Ochroma pyramidale (A.J.
Cavanilles) I. Urban] submerged in the
Patuxent River at Triadelphia Reservoir,
Brighton, Maryland, 18.XII.1968, leg. C.A.
Shearer CS-125-6, ILLS 35119, culture from
the type: ATCC 22765.

Coniothyrium fagi L.R. Tehon, Mycologia 25:247.
1933. Holotype: On leaves of Fagus
grandifolia F. Ehrhart, Alto Pass, Union
County, Illinois, 13.V1.1927, leg. G.L. Stout,
JEES'22555:

Coniothyrium negundinis L.R. Tehon & E.Y.
Daniels, Mycologia 17:243. 1925. Holotype:
On Acer negundo L., Urbana, Champaign
County, Illinois, 3.VI.1922, leg. P.A. Young,
ILLS 13413.

Coniothyrium radicicola L.R. Tehon, Mycologia
29:442. 1937. Holotype: On U/mus
americana L., Dayton, Ohio, X1.1934, leg.
H.L. Jacobs, ILLS 24540.

Coniothyrium ulmi B.C. Tharp, Mycologia 9:116.
1917. Isotypes: On Ulmus campestris L.,
Bonham, Texas, 21.VIII.1916, ILLS 22805,
TEX. Holotype: BPI.

Coniothyrium zeae G.L. Stout, Mycologia 22:273.
1930. Holotype: On Zea mays L., Putnam,
Putnam County, Illinois, 6.X.1926, leg. G.L.
Stout, ILLS 19686. Paratype: On leaves of
Zea mays L., Casey, Clark County, Illinois,
24.X.1927, leg. G.L. Stout, ILLS 21159.

Cribopeltis citrullina L.R. Tehon, Mycologia
25:252. 1933. Holotype: On the fruit of
Citrullus vulgaris H.A. Schrader, Spring Bay,
Woodford County, Illinois, 30.1X.1927, leg.
G.L. Stout, ILLS 22882.

Cryptostictis paeoniae L.R. Tehon & E.Y. Daniels,
Mycologia 17:243—244. 1925. Holotype: On
leaves of Paeonia officinalis L., Bloomfield,
Johnson County, Illinois, 25. VII.1922, leg.
P.A. Young, ILLS 6024.

Cylindrosporium quercinum J.C. Carter, Illinois
Natural History Survey Bulletin 21(6):221.
1941. Holotype: On Quercus imbricaria A.
Michaux, Carter, Marion County, Illinois,
17.1X.1936, leg. GH. Boewe, ILLS 27081.

Cyphellopycnis pastinacea L.R. Tehon & G.L. Stout,
Mycologia 21:189. 1929. Holotype: On
Pastinaca sativa L., Arnold, Morgan County,
Illinois, 20.VII.1926, leg. G.L. Stout, ILLS
13257.

Cytospora sambucina L.R. Tehon & G.L. Stout,
Mycologia 19:122. 1927. Holotype: On
Sambucus canadensis L., Oregon, Ogle
County, Illinois, 26. VIII.1926, leg. C.L.
Porter, ILLS 7426.

Catalog of Types

539

Cytospora sassafrasicola L.R. Tehon & E.Y.
Daniels, Mycologia 19:122. 1927. Holotype:
On Sassafras variifolium (R.A. Salisbury) O.
Kuntze [=S. al/bidum (T. Nuttall) C.G. Nees von
Esenbeck], Olney, Richland County, Illinois,
14.VI.1924, leg. P.A. Young, ILLS 12161.

Dactylaria fusiformis C.A. Shearer & J.L. Crane,
Mycologia 63:243. 1971. Holotype: Isolated
from balsa wood blocks |Ochroma
pyramidale (A.J. Cavanilles) I. Urban]
submerged in the Patuxent River at the Naval
Ordnance Laboratory Acoustics Facility,
Triadelphia Reservoir, Brighton, Maryland,
20.X.1967, leg. C.A. Shearer & J.L. Crane,
ILLS 34519.

Dendrophoma zeae L.R. Tehon, Mycologia 25:246.
1933. Holotype: On arid leaves of Zea mays
L., Clay City, Clay County, Illinois,
8.X1.1926, leg. G.L. Stout, ILLS 7742.

Diderma diadematum J.D. Schoknecht & J.L. Crane,
Transactions of the British Mycological
Society 70:146—147. 1978. Holotype: On
submerged, decayed leaves of angiosperms
(Acer sp.) and Taxodium distichum (L.) L.C.
Richard, Elvira Cypress Swamp (Deer Pond),
Johnson County, Illinois, 9.111.1977, leg. J.D.
Schoknecht & J.L.Crane, ILLS 36664.
Isotype: NY, TER, K. Paratypes: ILLS
36883, ILLS 36884, ILLS 36885.

Diplodia acericola L.R. Tehon & G.L. Stout,
Mycologia 21:190. 1929. Holotype: On Acer
saccharum H. Marshall, Mt. Pleasant, Union
County, Illinois, 7.VII.1926, leg. G.L. Stout,
ILLS 14104.

Diplodia sambuci L.R. Tehon & E.Y. Daniels,
Mycologia 19:125. 1927. Holotype: On
Sambucus canadensis L., Oregon, Ogle
County, Illinois, 26. VIII.1922, leg. C.L.
Porter, ILLS 7073.

Diplodia ulmi J.B. Dearness, Mycologia 8:102.
1916. Isotype: On dead branchlets of U/mus
americana L., London, Ontario, X.1903, J.B.
Dearness 3052, ILLS 22806.

Diplopeltis sassafrasicola L.R. Tehon & G.L. Stout,
Mycologia 21:193. 1929. Holotype: On
Sassafras albidum (T. Nuttall) C.G. Nees von
Esenbeck, Thebes, Alexander County, Illinois,
17.VII.1922, leg. P.A. Young, ILLS 581.

Discosia potentillae L.R. Tehon, Mycologia 25:253.
1933. Holotype: On leaves of Potentilla
canadensis L., Marlow, Jefferson County,
Illinois, 7.1X.1926, leg. L.R. Tehon, ILLS
22390.

Dothistroma pini R.L. Hulbary, Illinois Natural
History Survey Bulletin 21(7):236. 1941.
Holotype: On Pinus nigra J. Arnold var.
austrica (F. Hoess) H. Badoux, Waterman,
DeKalb County, Illinois, 29.X1.1938, leg. J.C.
Carter, ILLS 27093.

Illinois Natural History Survey Bulletin

phragmia boewei J.L. Crane, Mycologia
64:658-661. 1972. Holotype: On decayed
plant material, Grantsville Cypress Swamp,
east of Grantsville, Johnson County, Illinois,
2.V.1969, leg. J.L. Crane 64-69, ILLS 34948.

Endophragmia triseptata C.A. Shearer, J.L. Crane &
M.A. Miller, Mycologia 68:184—186. 1976.
Holotype: On decaying submerged wood,
Elvira Cypress Swamp (Deer Pond), Johnson
County, Illinois, 27.V1.1970, leg. J.L. Crane
28-70, ILLS 36154. Isotype: NY.

Epicoccum nigrum J.H. Link, Observationes in
Ordines Plantarum Naturales, Dissertatio
Secunda 2:32. 1815. Isotype: Communicated
by H. Sydow, date unknown, ILLS 24668.

Epidermella hansbroughii L.R. Tehon, Mycologia
31:688-689. 1939. Holotype: On Rubus
idaeus L. var. aculeatissimum E.A. von Regel
& H.S. Tiling, Bennington, Vermont,
8.VIL.1937, leg. J.R. Hansbrough 3069, ILLS
25862.

Erikssonia protii E.K. Cash, Mycologia 35:634—635.
1943. Holotype: On Protium asperum P.C.
Standley, Almirante, Panama, 28.VIII.1940,
leg. R.C. Lorenz 3070, BPI 71385. Isotype:
ILLS 41994.

Exilispora plurisepta L.R. Tehon & E.Y. Daniels,
Mycologia 19:113. 1927. Holotype: On
Erigeron sp., McNabb, Putnam County,
Illinois, 28.VI.1922, leg. C.L. Porter, ILLS
8404.

Exophoma astericola L.R. Tehon, Mycologia
21:188—189. 1929. Holotype: On Aster
tardiflorus L., Paris, Edgar County, Illinois,
4.X1.1926, leg. G.L. Stout, ILLS 19386.

Fusicoccum elaeagni J.C. Carter & C.M. Sacamano,
Mycologia 59:537. 1967. Holotype: On
Elaeagnus angustifolia L., J.F. Neil property,
9007 McKnight Woods, Clayton, Missouri,
18.VI.1963, leg. C.M. Sacamano, ILLS
34453.

Gloeosporium impatientis H.W. Anderson, Transac-
tions of the Illinois State Academy of Science
15:128. 1922. Holotype: On /mpatiens

biflora T. Walter, Brownfield Woods, Urbana,

Champaign County, Illinois, 19.VII.1921,
leg. H.W. Anderson, ILLS 4722.

Glomerella vignicaulis L.R. Tehon, Mycologia
29:435-436. 1937. Holotype: On Vigna
sinensis (L.) S.L. Endlicher, Equality, Gallatin
County, Illinois, 8.1X.1932, leg. G.H. Boewe,
ILLS 23703. Paratypes: Carmi, White
County, Illinois, 10.1X.1934, leg. G.H.
Boewe, ILLS 24809; Olmstead, Pulaski
County, Illinois, 17.1X.1933, leg. G.H.
Boewe, ILLS 25450.

Grovesinia pyramidalis M.N. Cline, J.L. Crane &
S.D. Cline, Mycologia 75:991. 1983.
Holotype: Dried sclerotia and apothecia from

Vol. 34 Art. 6

sterilized vermiculite in deep dishes and
derived dried cultures of micro- and
macroconidia, Champaign County, Illinois,
VIII.1982, leg. M.N. Cline, ILLS 43169.
Isotypes: BPI, CUP, IMI.

Gymnosporangium vauqueliniae W.H. Long & L.N.
Goodding, Mycologia 31:671—673. 1939.
Paratype: On Juniperus monosperma (G.
Engelmann) C.S. Sargent, Superstition
Mountain, Arizona, 5.V.1939, leg. L.N.
Goodding 8371, ILLS 41921.

Halosarpheia cincinnatula C.A. Shearer & J.L.
Crane, Botanica Marina 23:613. 1980.
Holotype: On balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban]
submerged in intake canal (Patuxent River) of
the Potomac Electricity Generating Plant at
Chalk Point, Ann Arundel County, Maryland,
16.VI.1969, leg. C.A. Shearer C.S. 93-2, ILLS
39006.

Halosarpheia retorquens C.A. Shearer & J.L. Crane,
Botanica Marina 23:608-610. 1980. Holotype:
On dried culture of alfalfa (Medicago sativa L),
originally isolated from balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban}
submerged in the Sangamon River below the
sewage treatment plant at Decatur, Macon
County, Illinois, 21.X.1975, leg. J.L. Crane &
C.A. Shearer C.S. 549-2, ILLS 38994.

Helminthosporium zeicola G.L. Stout, Mycologia
22:273-274. 1930. Holotype: On stalks of
Zea mays L., Dixon, Lee County, Illinois,
27.1X.1926, leg. G.L. Stout, ILLS 19884.
Paratypes: Eichorn, Hardin County, Illinois,
21.X.1926, leg. G.L. Stout, ILLS 20180;
Shelbyville, Shelby County, Illinois,
16.X1.1926, leg. G.L. Stout, ILLS 20181;
Mount Carroll, Carroll County, Illinois,
27.1X.1926, leg. G.L. Stout, ILLS 20182.

Hyalocylindrophora venezuelensis J.L. Crane & K.P.
Dumont, Canadian Journal of Botany
56:2616. 1978. Isotype: On unidentified
wood, 30 km north of San Cristobal, on San
Cristobal-LaGrita Road, Edo. Tachira,
Venezuela, 28. VII.1971, leg. K.P. Dumont,
G.J, Samuels, and L. Borjas VE-3249, ILLS
36940. Holotype: NY.

Hydnellum pineticola K.A. Harrison, Canadian
Journal of Botany 42:1226. 1964. Isotype:
North of Sheephead Lake, Chippewa County,
Michigan, 23.VIII.1961, leg. A-H. Smith &
K.A. Harrison, A.H.S. 63973, ILLS 47446.

Hydnum calvatum K.A. Harrison var. calvatum,
Canadian Journal of Botany 42:1216—1217.
1964. Paratype: Under spruce, Middle Fork
of Lake Fork Creek, Idaho National Forest,
Valley County, Idaho, 5. VIIL.1941, leg. A.H.
Smith 16023, ILLS 44310. Holotype: MICH.

May 1992

Hypoderma apocyni L.R. Tehon, Mycologia 31:679—

680. 1939. Holotype: On dead stems of
Apocynum medium E.L. Greene, Pine Plains,
New York, 27.VIII.1935, leg. J.R.
Hansbrough 1766, ILLS 25094.

Hypoderma caryae L.R. Tehon, Mycologia 31:680—
681. 1939. Holotype: On fallen petioles of
Carya glabra (P. Miller) R. Sweet, Hamilton,
Massachusetts, 7.X1.1935, leg. H.G. Eno,
communicated by J.R. Hansbrough 1772,
ILLS 2509S.

Hysterium pulcherrimum L.R. Tehon & P.A. Young,

Mycologia 16:31—32. 1924. Holotype: On
bark of Platanus occidentalis L., White
Heath, Piatt County, Illinois, 12.V.1923, leg.
P.A. Young, ILLS 4949.

Intercalarispora nigra J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
61:2243-2244. 1983. Holotype: On decayed
wood submerged in Forman Cypress Swamp,
NW of Belknap, Johnson County, Illinois,
29.X.1970, leg. J.L. Crane 203-69,

ILLS 43047. Isotype: NY. Paratypes: ILLS
43045, ILLS 43046, ILLS 43048.

Kaskaskia gleditsiae G.L. Born & J.L. Crane,

Phytopathology 62:927—929. 1972. Holo-

type: Parasitic on Gleditsia triacanthos L. var.

inermis C.K. Schneider, Freer Gymnasium,
University of Illinois, Urbana, Champaign
County, Illinois, leg. J.C. Carter, 17.V1.1968,
ILLS 34832. Isotype: IMI 151725, NY,
culture from the type: ATCC 22647.

Labrella aspidistrae L.R. Tehon & E.Y. Daniels,
Mycologia 19:126. 1927. Holotype: On
Aspidistra sp., Libertyville, Lake County,
Illinois, 9.VI.1922, leg. C.L. Porter, ILLS
7648.

Lactarius areolatus L.R. Hesler & A.H. Smith,
North American Species of Lactarius,
University of Michigan Press, Ann Arbor, pp.
515-516. Paratypes: Highland Recreation
Area, Oakland County, Michigan,

10. VIII.1972, leg. A.LH. Smith & N.S. Weber,

A.HLS. 81439, ILLS 47447; 27.VIII.1972, leg.

A.H. Smith & N.S. Weber, A.H.S. 81784,
ILLS 47452; Mill Lake, Waterloo Recreation
Area, Washtenaw County, Michigan,
11.VIII.1972, leg. A-H. Smith & N.S. Weber,
A.H.S. 81456, ILLS 47451; Gorman Lake,
Washtenaw County, Michigan, 20. VIII.1972,

leg. A.H. Smith & N.S. Weber, A.H.S. 81635,

ILLS 47448; Cedar Lake, Waterloo Recre-
ation Area, Washtenaw County, Michigan,
11.VII.1973, leg. A.H. Smith & N.S. Weber,
A.H.S. 84178, ILLS 47450; Haven Hill,
Highland Recreation Area, Oakland County,
Michigan, 12.VIII.1973, leg. A.H. Smith &
N.S. Weber, A.H.S. 84424, ILLS 47449.

Catalog of Types 541

Lepidopterella palustris C.A. Shearer & J.L. Crane,
Transactions of the British Mycological
Society 75:194—195. 1980. Holotype: On
unidentified submerged twigs and cultured on
alfalfa (Medicago sp.) stems, Elvira Cypress
Swamp, Johnson County, Illinois,
3.VIIL.1977, leg. J.L. Crane & C.A. Shearer
CS-470, ILLS 38990. Culture from type:
ATCC 38017. Isotype: NY. Paratype: On
twigs submerged in Elvira Cypress Swamp,
Johnson County, Illinois, 6 VIII 1974, leg.
C.A. Shearer & J.L. Crane CS-470-1, ILLS
38991.

Leptosphaeria maydis G.L. Stout, Mycologia
22:275. 1930. Holotype: On leaves of Zea
mays L., Vandalia, Fayette County, Illinois,
6.X1.1926, leg. G.L. Stout, ILLS 19423.
Paratypes: Shelbyville, Shelby County,
Illinois, 6.X1.1926, leg. G.L. Stout, ILLS
19669; Moline, Rock Island County, Illinois,
8.X.1926, leg. G.L. Stout, ILLS 19716;
Streator, La Salle County, Illinois,
23.1X.1926, leg. G.L. Stout, ILLS 19671;
Elgin, Kane County, Illinois, 24.1X.1926, leg.
G.L. Stout, ILLS 19725; Mt. Carmel, Wabash
County, Illinois, 6.X.1927, leg. G.L. Stout,
TELS 21223:

Leptosphaeria variiseptata G.L. Stout, Mycologia
22:276-277. 1930. Holotype: On leaves of
Zea mays L., Roscoe, Winnebago County,
Illinois, 25.1X.1926, leg. G.L. Stout, ILLS
19726. Paratype: Carmi, White County,
Illinois, 10.X1.1926, leg. G.L. Stout, ILLS
19727.

Leptosphaeria zeae G.L. Stout, Mycologia 22:277.
1930. Holotype: On leaves of Zea mays L.,
Sandoval, Marion County, Illinois, 6.X1.1926,
leg. G.L. Stout, ILLS 19421. Paratype: Clay
City, Clay County, Illinois, 8.X1.1926, leg.
G.L. Stout, ILLS 19422.

Leptosphaerulina vignae L.R. Tehon & G.L. Stout,
Phytopathology 18:703. 1928. Holotype: On
Vigna sinensis Hassk., Huey, Clinton County,
Illinois, 5.1X.1927, leg. G.L. Stout, ILLS
20937.

Leptostroma querci L.R. Tehon, Mycologia 29:446.
1937. Holotype: On leaves of Quercus
imbricaria A. Michaux, Ramsey, Fayette
County, Illinois, 7.V1I.1927, leg. L.R. Tehon,
ILLS 9455.

Leptostromella solani L.R. Tehon, Mycologia
40:317-318. 1948. Holotype: On stems of
Solanum carolinense L., LeRoy, McLean
County, Illinois, 11.X.1943, leg. G.H. Boewe,
ILLS 30051.

Leptothyriella liquidambaris L.R. Tehon & G.L.
Stout, Mycologia 21:192—193. 1929.
Holotype: On Liguidambar styraciflua L.,
Olmstead, Pulaski County, Illinois,

9. VIII.1922, leg. P.A. Young, ILLS 1445.

Illinois Natural History Survey Bulletin

sothyrium anthelmintict L.R. Tehon, Mycologia
40:314-315. 1948. Holotype: On dead stems
of Chenopodium ambrosioides L. var.
anthelminticum (L.) A. Gray, Harrisburg,
Saline County, Illinois, 16.VIII.1943, leg.
G.H. Boewe, ILLS 30049.

Leptothyrium avenae L.R. Tehon, Mycologia
40:315. 1948. Holotype: On Avena sativa L.,
Woodford, Woodford County, Illinois,

1. VII.1938, leg. G.H. Boewe, ILLS 29926.

Leptothyrium brunnichiae L.R. Tehon, Mycologia
16:136. 1924. Holotype: On Brunnichia
cirrhosa J. Gaertner, Karnak, Pulaski County,
Illinois, 8. VIII.1922, leg. PA. Young, ILLS
1120.

Leptothyrium fimbriatum L.R. Tehon, Mycologia
40:315-317. 1948. Holotype: On leaves of
Smilax hispida (G. Muhlenberg) M.L.
Fernald, Topeka, Mason County, Illinois,
20.X.1937, leg. G.H. Boewe, ILLS 30050.

Leptothyrium maximum L.R. Tehon & E.Y. Daniels,
Mycologia 17:245. 1925. Holotype: On
diseased twigs of Acer negundo L., Urbana,
Champaign County, Illinois, 3.V1I.1922, leg.
P.A. Young, ILLS 1795.

Leptothyrium zeae G.L. Stout, Mycologia 22:278.
1930. Holotype: On leaves of Zea mays L.,
Vandalia, Fayette County, Illinois, 6.X1.1926,
leg. G.L. Stout, ILLS 19445. Paratypes:
Sullivan, Moultrie County, Illinois,
16.X1.1926, leg. G.L. Stout, ILLS 19670;
Belleview, Calhoun County, Illinois,
7.X1.1927, leg. G.L. Stout, ILLS 21154.

Lophodermina melaleucaum (E.M. Fries:E.M. Fries)
F. v. Hohnel var. epiphyllum S.M. Zeller,
Mycologia 26:293. 1934. Holotype: On
Vaccinium ovatum F.T. Pursh, Waldport,
Oregon, X.1929, leg. S.M. Zeller, ILLS
24405.

Lophodermina septata L.R. Tehon, Illinois Biological
Monographs, University of Illinois at Urbana-
Champaign 13(4):109. 1935. Holotype: On
Picea sitchensis (A.H.G. Bongard) E.A.
Carriere, Knappa, Clatsop County, Oregon,
3.V.1919, leg. E.J. Perkins, communicated by
W.W. Wagener, ILLS 23480.

Lophodermium antarcticum C.L. Spegazzini, Fungi
Fuegani No. 304. 1887; Boletin de la
Academia Nacional de Ciencias en Cordoba
11:249-250. 1887. Isotype: On Rostkovia
grandiflora W.J. Hooker, Staten Island, Tierra
del Fuego, Argentina, Mart. 1882, leg. C.L.
Spegazzini, ILLS 23567 as a microscopic
preparation. Holotype: LPS.

Lophodermium camelliae S.C. Teng, Sinensia.

Special Bulletin of the Metropolitan Museum

of Natural History 4:138. 1933. Isotype: On

fallen leaves of Camellia sp., Teng’s garden,

Foochow, Fukien Province, China,

18. VIIT.1933, leg. S.C. Teng 1904, ILLS

25087.

Vol. 34 Art. 6

Lophodermium clavuligerum C.L. Spegazzini, Fungi
Fuegani No. 305. 1887; Boletin de la Academia
Nacional de Ciencias en Cordoba 11:250.
1887. Presumed isotype: On Pernettia
mucronata (L. f) C. Gaudichaud-Beaupré ex
K. Sprengel, Staten Island, Tierra del Fuego,
Argentina, date unknown [as Mart. 1882 in
protologue], leg. unknown, ILLS 23564.

Lophodermium danthoniae L.R. Tehon, Mycologia
31:690-691. 1939. Isotype: On dead leaves
of Danthonia spicata (L.) A. Beauvois, Brule,
Wisconsin, 11.VIII.1934, leg. J.J. Davis, ILLS
25090. Holotype: WIS. Paratype: Carbon-
dale, Jackson County, Illinois, 28.1V.1938,
leg. G.H. Boewe, ILLS 27027.

Lophodermium fuegianum C.L. Spegazzini, Fungi
Fuegiani No. 306. 1887; Boletin de la
Academia Nacional de Ciencias en Cordoba
11:250. 1887. Presumed isotype: On
Rostkovia grandiflora W.J. Hooker, Staten
Island, Tierra del Fuego, Argentina, date
unknown [as Agaia, Jun. 1882 in protologue],
leg. unknown, ILLS 23565 as a microscopic
preparation.

Lophodermium leptothecium C.L. Spegazzini, Fungi
Guaranitici I, No. 307. 1885; Anales de la
Sociedad Cientifica Argentina 19:261. 1885.
Isotype: On Laurinea sp., Guarapi, Paraguay,
VI.1883, leg. B. Balansa, ILLS 23569.
Holotype: LPS.

Lophodermium rosae S.C. Teng, Sinensia. Special
Bulletin of the Metropolitan Museum of
Natural History, 4:138—139. 1933. Isotype:
On Rosa sp. (on dead twigs), Heng-shan,
Hunan Province, China, [IX.1933, leg. C.I.
Shen 390, communicated by S.C. Teng, ILLS
25086. Holotype: Metropolitan Museum
Academia Sinica, Nanjing, China.

Lophodermium sesleriae A. Hilitzer, Védecké spisy
Vydavané Céskoslovenskou Akademii
Zemedelskou 3:91. 1929. Isotype: On
Sesleria caerulea (L.) P. Arduino, Prague,
Czechoslovakia, 13.X.1927, leg. A. Hilitzer,
ILLS 24394.

Lophodermium subtropicale C.L. Spegazzini, Anales
del Museo Nacional de Historia Natural de
Buenos Aires 23:88-89. 1912; Mycetes
Argentinenses, Series 6, No. 1443. 1912.
Isotype: On Psidium sp., Puerto Leon,
Missiones, Argentina, IX.1909, leg. unknown,
ILLS 23566 as a microscopic preparation.
Holotype: LPS.

Lophodiscella asparagi L.R. Tehon, Mycologia
25:253. 1933. Holotype: On arid stems of
Asparagus officinalis L., Villa Ridge, Pulaski
County, Illinois, 10.X1.1927, leg. G.L. Stout,
ILLS 7576. Paratypes: Anna, Union County,
Illinois, 10.X1.1927, leg. G.L. Stout, ILLS
7597; Alto Pass, Union County, Illinois,
10.X1.1927, leg. G.L. Stout, ILLS 7603;
26.X1.1927, leg. G.L. Stout, ILLS 7740.

May 1992

Luttrellia estuarina C.A. Shearer, Mycologia 70:692—
693. 1978. Holotype: On balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban] from
intake canal of Potomac Electric Power
Company, Electricity Generating Plant, Chalk
Point, Patuxent River, Maryland, 26. VII.1973,
leg. C.A. Shearer CS-80-4, ILLS 36979.

Macrophoma cercis L.R. Tehon & G.L. Stout,
Mycologia 21:188. 1929. Holotype: On
Cercis canadensis L., Venedy, Washington
County, Illinois, 8.1X.1926, leg. G.L. Stout,
TELS 19972.

Macrophoma oblongata L.R. Tehon, Mycologia
29:438. 1937. Holotype: On Poa pratensis
L., Rushville, Schuyler County, Illinois,

2. VIII.1935, leg. G.H. Boewe, ILLS 25381.

Macrophoma phlei L.R. Tehon & G.L. Stout,
Mycologia 21:188. 1929. Holotype: On
Phleum pratense L., Wayne City, Wayne
County, Illinois, 8.X1.1926, leg. G.L. Stout,
ILLS 19413.

Macrophoma rubi L.R. Tehon, Mycologia 29:441.
1937. Holotype: On Rubus idaeus L. var.
strigosus (A. Michaux) L.J. Maximowicz
(Latham raspberry, cultivated), Barry, Pike
County, Illinois, 30. VIHI.1934, leg. G.H.
Boewe, ILLS 24802.

Macrophoma secalina L.R. Tehon, Mycologia
29:438-440. 1937. Holotype: On dead leaves
and sheaths of Secale cereale L., Liberty,
Adams County, Illinois, 23.V1.1931, leg. G.H.
Boewe, ILLS 23913. Paratype: Westville,
Vermilion County, Illinois, 3.VII.1935, leg.
G.H. Boewe, ILLS 25073.

Marssonina salicina L.R. Tehon, Mycologia 25:255—
256. 1933. Holotype: On leaves of Salix nigra
H. Marshall, Lincoln, Logan County, Illinois,
18.V1.1930, leg. L.R. Tehon, ILLS 22258.

Masoniomyces claviformis J.L. Crane & K.P.
Dumont, Canadian Journal of Botany 53:847.
1975. Isotype: On wood, Dolphin Head,
Hanover Parish, Jamaica, 22.1.1971, leg.

R.P. Korf et al., 715, ILLS 35742. Holotype:
NY.

Melanospora interna L.R. Tehon & G.L. Stout,
Mycologia 21:181—182. 1929. Holotype: On
Lycopersicon esculentum P. Miller, Mound
City, Pulaski County, Illinois, 13.X1.1927,
leg. G.L. Stout, ILLS 20939.

Metasphaeria asparagi L.R. Tehon & G.L. Stout,
Mycologia 21:182—183. 1929. Holotype: On
Asparagus officinalis L., Anna, Union
County, Illinois, 11.X1.1926, leg. G.L. Stout,
ILLS 19944.

Metasphaeria sassafrasicola L.R. Tehon & G.L.
Stout, Mycologia 21:183. 1929. Holotype:
On Sassafras albidum (T. Nuttall) C.G. Nees
von Esenbeck, Seymour, Champaign County,
Illinois, 15.X.1925, leg. L.R. Tehon, ILLS
20103 [as S. variifolium (R.A. Salisbury) O.
Kuntze in protologue}.

Catalog of Types 543

Microstroma pithecolobii E.M.R. Lamkey in F.L.
Stevens, Mycologia 12:52. 1920. Isotype: On
Samanea saman (N.J. Jacquin) E.D. Merrill
{as Pithecollobium saman (N.J. Jacquin) G.
Bentham], Mayaguez, Puerto Rico, XII.1913,
leg. F.L. Stevens 6734, ex 70896, ILLS
41770.

Mollisia lithocarpi E.K. Cash, Mycologia 50:647—
648. 1958. Isotype: On dead leaves attached
to fallen branches of Lithocarpus densiflora
(W.J. Hooker & G. Arnott) A. Rehder, Big
Basin State Park, Santa Cruz County,
California, 15. VII.1954, leg. L. Bonar, ILLS
40594.

Monodictys cruciseptata J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
61:2244-2247. 1983. Holotype: On
submerged sticks incubated on filter paper,
also on filter paper, Shark Valley, Everglades
National Park, Florida, 3.[11.1976, leg.
J.L.Crane & J.D. Schoknecht, ILLS 42713.

Mycosphaerella cornicola L.R. Tehon & E.Y.
Daniels, Mycologia 17:240—241. 1925.
Holotype: On bark of Cornus stolonifera A.
Michaux, Apple River Canyon, Jo Daviess
County, Illinois, 17.VII.1924, leg. P.A.
Young, ILLS 13596.

Mycosphaerella holci L.R. Tehon, Mycologia
29:434435. 1937. Holotype: On living
leaves of Sorghum vulgare C.H. Persoon var.
technicum (F.A. Kornicke) O. Stapf ex J.H.
Holland, Oakland, Coles County, Illinois,
5.X.1927, leg. G.L. Stout, ILLS 11591.
Paratype: Mattoon, Coles County, Illinois,
5.X.1927, leg. G.L. Stout, ILLS 11682.

Mycosphaerella juglandis K.J. Kessler, Mycologia
76:363. 1984. Isotype: On Juglans nigra L.,
SW'/2 Sec. 28, T14S, R2W, Alexander
County, Illinois, 23.V.1979, leg. K.J. Kessler,
ILLS 42084.

Mycosphaerella zeicola G.L. Stout, Mycologia
22:278-279. 1930. Holotype: On Zea mays
L., Vandalia, Fayette County, Illinois,
6.X1.1926, leg. G.L. Stout, ILLS 13803.
Paratypes: Belleview, Calhoun County,
Illinois, 7.X1.1927, leg. G.L. Stout, ILLS
21154; Bruce, Moultrie County, Illinois,
21.X.1927, leg. G.L. Stout, ILLS 21194;
Champaign County, Illinois, 23.1X.1927, leg.
G.L. Stout, ILLS 21151; Effingham,
Effingham County, Illinois, 20.1X.1927, leg.
G.L. Stout, ILLS 21166; Gibson City, Ford
County, Illinois, 4.X.1926, leg. G.L. Stout,
ILLS 19697; Harrisburg, Saline County,
Illinois, 10.X.1927, leg. G.L. Stout, ILLS
21212; Mattoon, Coles County, Illinois,
15.1X.1927, leg. G.L. Stout, ILLS 21164;
Minonk, Woodford County, Illinois,
29.1X.1926; leg. G.L. Stout, ILLS 19685;
McLeansboro, Hamilton County, Illinois,
10.X1.1926, leg. G.L. Stout, ILLS 20136;

Illinois Natural History Survey Bulletin

Mt. Carmel, Wabash County, Illinois,
6.X.1927, leg. G.L. Stout, ILLS 21222;
Riverton, Sangamon County, Illinois,
19.X.1927, leg. G.L. Stout, ILLS 21216; West
City, Franklin County, Illinois, 12.X1.1926,
leg. G.L. Stout, ILLS 19629.

Nais glitra J.L. Crane & C.A. Shearer, Transactions
of the British Mycological Society 86:509.

1986. Holotype: Red mangrove wood
(Rhizophora mangle L.), submerged in Coot
Bay, Everglades National Park, Florida,
14.11.1984, leg. J.L. Crane & C.A. Shearer
CS-720-1, ILLS 44900.

Nakataea serpens C.A. Shearer & J.L. Crane,
Transactions of the British Mycological
Society 73:370. 1979. Holotype: Isolated
from plant debris collected from Quiver
Creek, Mason County, Illinois, 11.VIII.1978,
leg. C.A. Shearer, ILLS 38410. Isotype:
ILLS 42909, culture from the type: ATCC
38018.

Neta lignicola C.A. Shearer, Mycologia 66:21—23.
1974. Holotype: On balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban]
submerged in Patuxent River at Triadelphia
Reservoir, Brighton, Maryland, 22.VII.1969,
leg. C.A. Shearer CS-223, ILLS 35536.

Neta patuxentica C.A. Shearer & J.L. Crane,
Mycologia 63:241—242. 1971. Holotype:
Isolated from balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban]
submerged in the Patuxent River at the
Patuxent Wildlife Refuge Center, U.S.
Department of Interior, Laurel, Maryland,
U.S.A., 10.X.1967, leg. C.A. Shearer & J.L.
Crane A-64, ILLS 34520. Paratypes: DAOM
127056, ILLS 34827; DAOM 127053, ILLS
34809; DAOM 127057, ILLS 34828; DAOM
127054, ILLS 34825; DAOM 127055, ILLS
34826, culture from the type: ATCC 18854.

Nitschkia polygoni L.R. Tehon & E.Y. Daniels,
Mycologia 19:111. 1927. Holotype: On
Polygonum sp., Villa Ridge, Pulaski County,
Illinois, 21.V1.1925, leg. PA. Young, ILLS
9239.

Parasympodiella minima J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
60:369. 1982. Holotype: On submerged,
decayed plant material, cypress swamp,
Florida Route 6, Hamilton County, Florida,
17.V.1977, leg. J.L. Crane & J.D. Schoknecht,
ILLS 39789.

Paratomenticola georgiana J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
60:369-371. 1982. Holotype: On a dead
frond of Serenoa repens (W. Bartram) J.K.
Small, Monkey Lake Landing, Suwannee
Canal, Okefenokee National Wildlife Refuge,
Georgia, 28. VIII.1978, leg. J.L. Crane & J.D.
Schoknecht, ILLS 39897. Isotype: NY.

Vol. 34 Art. 6

Pestalozzia heucherae L.R. Tehon & E.Y. Daniels,
Mycologia 19:126—127. 1927. Holotype: On
Heuchera parviflora F.G. Bartling, Fountain
Bluff, Jackson County, Illinois, 20.V1.1924,
leg. P.A. Young, ILLS 2873.

Phacidium negundinis L.R. Tehon & E.Y. Daniels,
Mycologia 17:240. 1925. Holotype: On
diseased twigs of Acer negundo L., Urbana,
Champaign County, Illinois, 3.V1.1922, leg.
P.A. Young, ILLS 8890.

Phaeocytosporella zeae G.L. Stout, Mycologia
22:280-281. 1930. Holotype: On culms of
Zea mays L., Mattoon, Coles County, Illinois,
19.X.1926, leg. G.L. Stout, ILLS 20039.

Phaeoseptoria caricis L.R. Tehon & E.Y. Daniels,
Mycologia 17:245. 1925. Holotype: On
leaves of Carex sp., Ursa, Adams County,
Illinois, 28. VI.1922, leg. O.A. Plunkett, ILLS
15455.

Phialocephala fluminis C.A. Shearer, J.L. Crane, &
M.A. Miller, Mycologia 68:186—-188. 1976.
Holotype: On balsa wood [Ochroma
pyramidale (A.J. Cavanilles) I. Urban}
submerged in the Sangamon River, Station 15,
at Decatur, Macon County, Illinois,
15.1V.1975, leg. C.A. Shearer & J.L. Crane
466-1, ILLS 36160. Isotype: NY, culture
from the type: ATCC 32105.

Phialocephala illini J.L. Crane, Transactions of the
British Mycological Society 56:162—163.
1971. Holotype: On decayed wood of
Taxodium disticum (L.) L.C. Richard, Elvira
Cypress Swamp, northwest of Vienna,
Johnson County, Illinois, 18.X1.1969, leg. J.L.
Crane 202-69, ILLS 34911.

Phoma asparagina L.R. Tehon & G.L. Stout,
Mycologia 21:187. 1929. Holotype: On
Asparagus officinalis L., Anna, Union
County, Illinois, 11.X1.1926, leg. G.L. Stout,
ILLS 19943.

Phomopsis callistephi L.R. Tehon & E.Y. Daniels,
Mycologia 17:242. 1925. Holotype: On
stems of Callistephus hortensis (L.) C.G.
Nees von Esenbeck, Shelbyvilie, Shelby
County, Illinois, 20.1X.1924, leg. P.A. Young,
ILLS 2014.

Phomopsis ganjae J.M. McPartland, Mycotaxon. An
International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens 18:527—
528. 1983. Holotype: On living leaves of
Cannabis sativa L., Hanna City, Peoria
County, Illinois, 8.VIIT.1982, leg. P.L. Pruitt
& J.M. McPartland, ILLS 43621.

Phyllachora cinnae L.R. Tehon & E.Y. Daniels,
Mycologia 19:110—111. 1927. Holotype: On
Cinna arundinacea L., Granite City, Madison

County, Illinois, VII.1924 [as 1920 in
original publication], leg. PA. Young, ILLS
9316.

May 1992

Phyllosticta allegheniensis L.R. Tehon & G.L. Stout,

Mycologia 21:185. 1929. Holotype: On
Rubus allegheniensis T.C. Porter, Nashville,
Washington County, Illinois, 29. VII.1926,
leg. G.L. Stout, ILLS 20940.

Phyllosticta allii L.R. Tehon & E.Y. Daniels,
Mycologia 17:241—242. 1925. Holotype: On
Allium cepa L., Columbia, Monroe County,
Illinois, 24. VIII.1922, leg. P.A. Young, ILLS
11132.

Phyllosticta anserinae L.R. Tehon, Mycologia
25:243. 1933. Holotype: On living leaves of
Potentilla anserina L., Witt, Montgomery
County, Illinois, 23. VI.1927, leg. G.L. Stout,
ILLS 22762.

Phyllosticta aquilegiae L.R. Tehon & E.Y. Daniels,
Mycologia 17:241. 1925. Holotype: On
leaves of Aquilegia canadensis L., Marion,

Williamson County, Illinois, 19.VII.1922, leg.

P.A. Young, ILLS 2098.

Phyllosticta atomata L.R. Tehon, Mycologia 29:437.

1937. Holotype: On living leaves of
Plantago rugelii J. Decaisne, Elizabeth, Jo
Daviess County, Illinois, 24. VII.1927, leg.
L.R. Tehon, ILLS 10349.

Phyllosticta avenophila L.R. Tehon & E.Y. Daniels,
Mycologia 19:118-119. 1927. Holotype: On
Avena sativa L., Piper City, Ford County,
Illinois, 19.V1.1925, leg. L-.R. Tehon, ILLS
2415.

Phyllosticta chenopodiicola L.R. Tehon & E.Y.
Daniels, Mycologia 19:121. 1927. Holotype:
On Chenopodium album L., Lincoln, Logan
County, Illinois, 22.VII.1925, leg. L.R.
Tehon, ILLS 6309. Paratypes: Mason City,

Mason County, Illinois, 22.VII.1925, leg. L.R.

Tehon, ILLS 10190; Nashville, Washington
County, Illinois, 13. VIII.1925, leg. L.R.
Tehon, ILLS 16667.

Phyllosticta circuligerens L.R. Tehon & E.Y.
Daniels, Mycologia 19:120—121. 1927.
Holotype: On Rumex altissimus A. Wood,
Dongola, Union County, Illinois,
12.VIII.1922, leg. PA. Young, ILLS 1007.

Phyllosticta dispergens L.R. Tehon, Mycologia
25:242-243. 1933. Holotype: On leaves of
Rubus flagellaris K.L. Willdenow, Belleville,
St. Clair County, Illinois, 9. VIII.1927, leg.
G.L. Stout, ILLS 829.

Phyllosticta glycineum L.R. Tehon & E.Y. Daniels,
Mycologia 19:117. 1927. Holotype: On
Glycine hispida C.J. Maximowicz [=G. max
(L.) E.D. Merrill], Arthur, Douglas County,
Illinois, 6.VII.1925, leg. L.R. Tehon, ILLS
4392.

Phyllosticta gymnocladi L.R. Tehon & E.Y. Daniels,

Mycologia 19:114. 1927. Holotype: On
Gymnocladus dioica (L.) K. Koch,
Lawrenceville, Lawrence County, Illinois,
27.VIII.1925, leg. L.R. Tehon, ILLS 10319.

Catalog of Types 545

Phyllosticta illinoensis L.R. Tehon & E.Y. Daniels,
Mycologia 19:120. 1927. Holotype: On
Sassafras variifolium (R.A. Salisbury) O.
Kuntze [=S. albidum (T. Nuttall) C.G. Nees
von Esenbeck], Grand Tower, Jackson
County, Illinois, 17.VIII.1922, leg. P.A.
Young, ILLS 574. Paratype: Coxeyville,
Monroe County, Illinois, 23. VIII. 1922, leg.
P.A. Young, ILLS 5096.

Phyllosticta menispermicola L.R. Tehon & E.Y.
Daniels, Mycologia 19:119. 1927. Holotype:
On Menispermum canadense L., Porterfield,
Marshall County, Illinois, 24.VIII.1922, leg.
C.L. Porter, ILLS 8038 [as 8039 in original
publication].

Phyllosticta neuroterigallicola L.R. Tehon,
Mycologia 25:240-241. 1933. Holotype: On
leaves of Quercus imbricaria A. Michaux,
Dongola, Union County, Illinois,

11. VIII.1927, leg. G.L. Stout, ILLS 22830.

Phyllosticta nymphaeicola L.R. Tehon & E.Y.
Daniels, Mycologia 19:117. 1927. Holotype:
On Nuphar luteum (L.) J. Sibthorp ssp.
macrophyllum (J.K. Small) E.O. Beal [as
Nymphaea advena (D.C. Solander) R. Brown
in protologue], Olney, Richland County,
Illinois, 28. VIII.1925, leg. L.R. Tehon, ILLS
11093.

Phyllosticta plantaginicola L.R. Tehon & E.Y.
Daniels, Mycologia 19:118. 1927. Holotype:
On Plantago virginica L., Marshall, Clark
County, Illinois, 1.VII.1925, leg. L.R. Tehon,
ILLS 3188.

Phyllosticta podophyllina L.R. Tehon & G.L. Stout,
Mycologia 21:184—185. 1929. Holotype: On
Podophyllum peltatum L., Columbia, Monroe
County, Illinois, 24.VI.1926, leg. L.R. Tehon,
ILLS 19480.

Phyllosticta porteri L.R. Tehon & E.Y. Daniels,
Mycologia 19:113—114. 1927. Holotype: On
Syringa vulgaris L., Oregon, Ogle County,
Illinois, 26. VIII.1922, leg. C.L. Porter, ILLS
3097.

Phyllosticta pteleicola L.R. Tehon & E.Y. Daniels,
Mycologia 17:241. 1925. Holotype: On
leaves of Prelea trifoliata L., Starved Rock,
LaSalle County, Illinois, 27.V1.1924, leg. P.A.
Young, ILLS 6807.

Phyllosticta rafinesquii H.W. Anderson, Transac-
tions of the Illinois State Academy of Science
15:128-129. 1922. Holotype: On Viola
rafinesquii E.L. Greene, Anna, Union County,
Illinois, 23.1V.1921, leg. H.W. Anderson,
ILLS 4724.

Phyllosticta rugelii L.R. Tehon & G.L. Stout,
Mycologia 21:184. 1929. Holotype: On
Plantago rugelii J. Decaisne, Lawrenceville,
Lawrence County, Illinois, 27.V1I.1926, leg.
L.R. Tehon, ILLS 19477.

Illinois Natural History Survey Bulletin

losticta scariolicola L.R. Tehon, Mycologia

25:245. 1933. Holotype: On living leaves of
Lactuca scariola L., West Union, Clark
County, Illinois, 8.V1.1927, leg. G.L. Stout,
ILLS 22503.

Phyllosticta solidaginicola L.R. Tehon & E.Y.
Daniels, Mycologia 19:116. 1927. Holotype:
On Solidago sp., Sumner, Richland County,
Illinois, 27. VIII.1925, leg. L.R. Tehon, ILLS
10114.

Phyllosticta zeae G.L. Stout, Mycologia 22:28 1—

282. 1930. Holotype: On leaf of Zea mays L..,

Robinson, Crawford County, Illinois,
5.X1.1926, leg. G.L. Stout, ILLS 19359.
Paratypes: DuQuoin, Perry County, Illinois,
8.1X.1927, leg. G.L. Stout, ILLS 21196;
Mattoon, Coles County, Illinois, 15.[X.1927,
leg. G.L. Stout, ILLS 21165.

Physalospora zeae G.L. Stout, Mycologia 22:282.
1930. Holotype: On leaves of Zea mays L.,
Vandalia, Fayette County, Illinois, 6.X1.1926,
leg. G.L. Stout, ILLS 19883.

Pirostoma nyssae L.R. Tehon, Mycologia 16:137.
1924. Holotype: On Nyssa sylvatica Marsh.,
Tunnel Hill, Johnson County, Illinois,
25.VII.1922, leg. P.A. Young, ILLS 2940.

Placosphaeria medicaginis L.R. Tehon, Mycologia
31:538-539. 1939. Holotype: On Medicago
sativa L., Freeburg, St. Clair County, Illinois,
22.X.1935, leg. GLH. Boewe, ILLS 25276.
Paratypes: Malta, De Kalb County, Illinois,
17.VII.1936, leg. G.H. Boewe, ILLS 26977;
Centralia, Marion County, Illinois,
29.1V.1938, leg. G.H. Boewe, ILLS 26704.

Pleosphaerulina zeicola G.L. Stout, Mycologia
22:284. 1930. Holotype: On leaf of Zea mays
L., Highland, Madison County, Illinois,
26.X.1927, leg. G.L. Stout, ILLS 21182.

Pleospora aquatica D., Griffiths, Bulletin of the
Torrey Botanical Club (and Torreya) 26:443.
1899. Isotype: On dead Eleocharis palustris
(L.) J.J. Roemer & J.A. Schultes, Aberdeen,
Brown County, South Dakota, V.1896, leg. D.
Griffiths, ILLS 42699.

Pleospora oleraceae L.R. Tehon & G.L. Stout,
Mycologia 21:183—184. 1929. Holotype: On
Brassica oleracea L. var. capitata L., West
Vienna, Johnson County, Illinois, 7.VII.1926,
leg. G.L. Stout, ILLS 19358.

Podospora pectinata N. Lundqvist, Svensk Botanisk
Tidskrift 64:417. 1970. Paratype: U.S.A.
Colorado, Boulder County, west base of Mt.
Steamboat at foothills of Front Range, 3 km
northwest of Lyons, near mouth of St. Vrain
{as Vain on label] Creek, altitude 1900 m, on
old cow dung incubated in moist chamber at
Uppsala, 30.1V.1966, leg. R. Santesson
18499-e, ILLS 35158, FH, IMI, TRTC, UC,
UPS.

Vol. 34 Ant. 6

Pseudodictya sassafrasicola L.R. Tehon & G.L.
Stout, Mycologia 21:192. 1929. Holotype:
On Sassafras varrifolium (R.A. Salisbury) O.
Kuntze [=S. albidum (T. Nuttall) C.G. Nees
von Esenbeck], Seymour, Champaign County,
Illinois, 15.X.1925, leg. L.R. Tehon, ILLS
9353.

Puccinia puritanica G.B. Cummins, Bulletin of the
Torrey Botanical Club (and Torreya) 68:45.
1941. Isotype: On Carex pensylvanica J.B.
Lamarck, Waltham, Massachusetts, 1.X.1910,
leg. A.B. Seymour, ILLS 41662, BPI.
Holotype: PUR.

Pyrenochaeta minuta J.C. Carter, Illinois Natural
History Survey Bulletin 21(6):219. 1941.
Holotype: On branch of Quercus palustris O.
von Muenchhausen, Xenia, Clay County,
Illinois, 8.X.1937, leg. J.C. Carter, ILLS
27082.

Rhizopogon oswaldii A.H. Smith in A.H. Smith and
S.M. Zeller, Memoirs of the New York
Botanical Garden, Bronx 14(2):107—108.
1966. Paratype: In an old horse corral, Mt.
Wilson Road, Bear Springs, Mt. Hood
National Forest, Wasco County, Oregon,
26.X.1946, leg. W.B. Gruber (A.H. Smith
25065), ILLS 44316.

Rhizopogon pinyonensis K.A. Harrison & A.H.
Smith var. pinyonensis, Canadian Journal of
Botany 46:890. 1968. Paratype: Under pinion
pine, Hyde Park Road, near Santa Fe, Santa
Fe County, New Mexico, 7.X.1967, leg. K.A.
Harrison 7321, ILLS 44325.

Rhizopogon pseudoaffinis A.H. Smith in A.H. Smith
and S.M. Zeller, Memoirs of the New York
Botanical Garden 14(2):138—139. 1966.
Paratype: Brundage Mountain, Valley
County, Idaho, 8.VIII.1962, leg. A.H. Smith
65780, ILLS 44320.

Rhizopogon rubescens E.L.R. Tulasne var.
ochraceus A.H. Smith, in A-H. Smith and
S.M. Zeller, Memoirs of the New York
Botanical Garden 14(2):99-100. 1966.
Paratype: Under white pine bark and alpine
fir, Heaven’s Gate Ridge, Seven Devils
Mountains, Idaho County, Idaho, 2. VIII.1958,
leg. A.H. Smith 59481, ILLS 44315.

Rhodosticta quercina J.C. Carter, Illinois Natural
History Survey Bulletin 21(6):223. 1941.
Holotype: On branch of Quercus palustris O.
von Muenchhausen, Onarga, Iroquois County,
Illinois, 25.X.1935, leg. J.C. Carter, ILLS
27083.

Rogersia annelidica C.A. Shearer & J.L. Crane,
Mycologia 68:949-950. 1976. =Filosporella
annelidica (C.A. Shearer & J.L. Crane) J.L.
Crane & C.L. Shearer, Mycotaxon 6:28. 1977.
Holotype: On sycamore leaves (Platanus
occidentalis L.) submerged in the Sangamon

May 1992

River at Brigham Station No. 1, Mahomet,
Champaign County, Illinois, 16.11.1976, leg.
C.A. Shearer S-77-8, ILLS 36352.
Paratypes: ILLS 36343, ILLS 36344, ILLS
36345, ILLS 36346, ILLS 36347, ILLS
36348, ILLS 36349, ILLS 36350, ILLS
36351, culture from the type: ATCC 32834.

Rogersiomyces okefenokeensis J.L. Crane & J.D.
Schoknecht, American Journal of Botany
65:903. 1978. Holotype: On submerged,
decayed leaves of angiosperms and Taxodium
sp., Trembling Earth Nature Trail, Steven
Foster State Park, Okefenokee National
Wildlife Refuge, Georgia, 10.V.1976, leg. J.L.
Crane & J.D. Schoknecht, ILLS 37095.
Isotype: NY, culture from the type: ATCC
36118.

Rostrosphaeria phlei L.R. Tehon & E.Y. Daniels,
Mycologia 19:112. 1927. Holotype: On
Phleum pratense L., Jacksonville, Morgan
County, Illinois, 25.VII.1925, leg. L.R.
Tehon, ILLS 893.

Scolecobasidium cylindrosporum J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
60:372. 1982. Holotype: On submerged,
decayed wood, Minnie’s Lake, Okefenokee
National Wildlife Refuge, Georgia,
17.V.1977, leg. J.L. Crane & J.D. Schoknecht,
ILLS 40889.

Scolecobasidium salmonicolor C.A. Shearer,
Mycologia 66:18—20. 1974. Holotype: On
Cantino PYF agar, isolated from balsa wood
submerged in the Patuxent River at Lower
Marlboro, Maryland, 6.V.1969, leg. C.A.
Shearer, CS-180, ILLS 35537, culture from
the type: ATCC 24299.

Sebacina farinacea D.P. Rogers, Pacific Science
1:97. 1947. Paratype: On sheath of Cocos
nucifera L., on campus, Mauoa, Oahu,
Hawaii, 20.11.1946, leg. L.A. Abbott & D.P.
Rogers (DPR 1175), ILLS 39247.

Sebacina petiolata D.P. Rogers, Pacific Science
1:99. 1947. Paratypes: On bark of Acacia
koa A. Gray, Tautalus Tr., Pauoa (c. 1500 ft),
Oahu, Hawaii, 21.VII.1946, leg. D.P. Rogers
(DPR 1331), ILLS 39249. On Cocos nucifera
L., fallen log, Ormed I., Wotje, Atoll,
Marshall Islands, 4.1X.1946, leg. D.P. Rogers
(DPR 1385), ILLS 39210.

Septocylindrium hydrophylli E.Y. Daniels,
Mycologia 19:127. 1927. Holotype: On
Hydrophyllum canadense L., Seymour,
Champaign County, Illinois, 15.X.1925, leg.
E.Y. Daniels, ILLS 9531.

Septogloeum equiseti L.R. Tehon, Mycologia
29:445-446. 1937. Holotype: On living
stems of Equisetum laevigatum A. Braun,
Sterling, Whiteside County, Illinois,
23.VII.1927, leg. L.R. Tehon, ILLS 10369.

Catalog of Types 547

Septoria atropurpurei L.R. Tehon, Mycologia
16:135—136. 1924. Holotype: On Euonymus
atropurpureus N.J. Jacquin, Marion,
Williamson County, Illinois, 19.VII.1922, leg.
P.A. Young, ILLS 2064.

Septoria collinsiae H.W. Anderson, Transactions of
the Illinois State Academy of Science 15:127.
1922. Holotype: On Collinsia verna T.
Nuttall, Brownfield Woods, Urbana,
Champaign County, Illinois, 18.V.1919, leg.
H.W. Anderson, ILLS 4723.

Septoria cunillae L.R. Tehon, Mycologia 25:249-250.
1933. Holotype: On Cunila origanoides (L.)
N.L. Britton, Alto Pass, Union County, Illinois,
13.VI.1927, leg. G.L. Stout, ILLS 22557.

Septoria eupatoriicola L.R. Tehon, Mycologia
25:250. 1933. Holotype: On leaves of
Eupatorium perfoliatum L., DuQuoin, Perry
County, Illinois, 13.V1.1930, leg. L.R. Tehon,
ILLS 22260.

Septoria festucina L.R. Tehon & E.Y. Daniels,
Mycologia 19:125-126. 1927. Holotype: On
Festuca elatior L., Tallula, Menard County,
Illinois, 16.VI.1922, leg. O.A. Plunkett, ILLS
9224.

Septoria pinicola J.B. Dearness, Mycologia 20:237.
1928. Isotype: On Pinus virginiana P. Miller,
Chain Bridge, Virginia, 10.V.1927, leg. G.G.
Hedgcock, communicated by Paul V. Siggers,
ILLS 27098. Holotype: DAOM.

Septoria septentrionalis H.W. Anderson, Transac-
tions of the Illinois State Academy of Science
15:127—128. 1922. Holotype: On lower
leaves of Ranunculus septentrionalis J.L.M.
Poiret, Brownfield Woods, Urbana,
Champaign County, Illinois, 21.1V.1921, leg.
H.W. Anderson, ILLS 4725.

Septoria tecomaxochitl L.R. Tehon & G.L. Stout,
Mycologia 21:191. 1929. Holotype: On
Campsis radicans (L.) B.C. Seemann ex
Bureau [as Tecoma radicans (L.) A.L. de
Jussieu in protologue}], Lawrenceville, Law-
rence County, Illinois, 20.X.1926 [as 26.X in
protologue], leg. G.L. Stout, ILLS 20946.

Septoria zeae G.L. Stout, Mycologia 22:284-285.
1930. Holotype: On leaf of Zea mays L.,
Joliet, Will County, Illinois, 24.1X.1926, leg.
G.L. Stout, ILLS 19673. Paratypes: Dixon,
Lee County, Illinois, 27.1X.1926, leg. G.L.
Stout, ILLS 19681; Elgin, Kane County,
Illinois, 24.1X.1926, leg. G.L. Stout, ILLS
19725; Moline, Rock Island County, Illinois,
8.X.1926, leg. G.L. Stout, ILLS 19716; Mt.
Carroll, Carroll County, Illinois, 27.1X.1926,
leg. G.L.Stout, ILLS 19682; Rockford,
Winnebago County, Illinois, 25.1X.1926, leg.
G.L. Stout, ILLS 19677; Stockton, Jo Daviess
County, Illinois, 27.1X.1926, leg. G.L.Stout,
ILLS 19683; Streator, La Salle County, Illinois,
23.1X.1926, leg. G.L.Stout, ILLS 20100.

Illinois Natural History Survey Bulletin

septoria zeicola G.L. Stout, Mycologia 22:286.
1930. Holotype: On leaf of Zea mays L.,
Vandalia, Fayette County, Illinois, 6.X.1926,
leg. G.L. Stout, ILLS 20102. Paratypes:
Casey, Clark County, Illinois, 24.X.1927, leg.
G.L. Stout, ILLS 21160; Harrisburg, Saline
County, Illinois, 10.X.1927, leg. G.L. Stout,
ILLS 21211; Mattoon, Coles County, Illinois,
15.1X.1927, leg. G.L. Stout, ILLS 21162;
Toulon, Stark County, Illinois, 7.X.1926, leg.
G.L. Stout, ILLS 20138; West City, Franklin
County, Illinois, 12.X1.1926, leg. G.L. Stout,
ILLS 19629.

Septoria zeina G.L. Stout, Mycologia 22:287. 1930.
Holotype: On leaf of Zea mays L.,
Taylorville, Christian County, Illinois,
20.X.1927, leg. G.L. Stout, ILLS 21231.

Sirococcus phlei L.R. Tehon & E.Y. Daniels,
Mycologia 19:122. 1927. Holotype: On
Phleum pratense L., New Berlin, Sangamon
County, Illinois, 25. VII.1925, leg. L.R.
Tehon, ILLS 14799.

Sphaeropsis ampelopsidis E.Y. Daniels in L.R.
Tehon & E.Y. Daniels, Mycologia 19:123—
124. 1927. Holotype: On Parthenocissus
quinquefolia (L.) J.L. Planchon [as
Ampelopsis quinquefolia (L.) A. Michaux in
original publication], Fisher, Champaign
County, Illinois, 20.X.1925, leg. L.R. Tehon,
ILLS 2541.

Sphaeropsis negundinis L.R. Tehon & E.Y. Daniels,
Mycologia 17:242—243. 1925. Holotype: On
diseased twigs of Acer negundo L., Urbana,
Champaign County, Illinois, 3.VI.1922, leg.
P.A. Young, ILLS 15198.

Sphaeropsis profundae L.R. Tehon & E.Y. Daniels,
Mycologia 19:124. 1927. Holotype: On
Fraxinus profunda (B.F. Bush) B.F. Bush
[=F. tomentosa F.A. Michaux], Fountain
Bluff, Jackson County, Illinois, 20.V1I.1924,
leg. P.A. Young, ILLS 17554.

Sporidesmium taxodii J.L. Crane, Transactions of the
British Mycological Society 58:425-426.
1972. Holotype: On submerged decaying
leaves of Taxodium distichum (L.) L.C.
Richard, Elvira Cypress Swamp, northwest of
Vienna, Johnson County, Illinois, 29.X.1969,
leg. J.L. Crane 222-69, ILLS 34936. Isotype:
NY.

Stagonospora heteroderae L.M. Carris, D.A. Glawe
& G. Morgan-Jones, Mycotaxon. An
International Journal Designed to Expedite
Publication of Research on Taxonomy &
Nomenclature of Fungi & Lichens 29:45 1—
452. 1987. Holotype: Isolated from cyst of
Heterodera glycines, Sidney, Champaign
County, Illinois, 22. VIII.1985, leg. L.M.
Carris, ILLS 46332, culture from the type:

ATCC 62861.

Vol. 34 Art. 6

Stagonospora scirpi L.R. Tehon, Mycologia 25:247—
248. 1933. Holotype: On leaves and sheaths
of Scirpus atrovirens K.L. Willdenow,
DuQuoin, Perry County, Illinois, 13.VI.1930,
leg. L.R. Tehon, ILLS 22259.

Stictopatella iridis L.R. Tehon, Mycologia 40:320—
321. 1948. Holotype: On leaves of /ris
virginica L. var. shrevei (J.K. Small) E. Ander-
son, Urbana, Champaign County, Illinois,
11.VI.1947, leg. R.A. Evers, ILLS 30048.

Stigmatea plantaginis L.R. Tehon & E.Y. Daniels,
Mycologia 19:111—112. 1927. Holotype: On
Plantago virginica L., Carthage, Hancock
County, Illinois, 24. VII.1922, leg. O.A.
Plunkett, ILLS 7652.

Stigmatophragmia sassafrasicola L.R. Tehon &
G.L. Stout, Mycologia 21:181. 1929.
Holotype: On Sassafras variifolium Kuntze,
Seymour, Champaign County, Illinois,
15.X.1925, leg. L.R. Tehon, ILLS 21698.

Taeniolella americana J.L. Crane & J.D.
Schoknecht, Canadian Journal of Botany
60:372-374. 1982. Holotype: On submerged,
decayed ament of Liquidambar styraciflua L.,
Goose Pond, Johnson County, Illinois,
28.1.1974, leg. J.L. Crane, ILLS 41016.

Taeniolina deightonii J.L. Crane & J.D. Schoknecht,
Mycologia 73:81—82. 1981. Isotype: On
Vismia guineensis J.D. Choisy, Njala (Kori)
Sierra Leone, 22.1.1951, leg. F.C. Deighton,
{as Torula herbarum], ex type material M
3806, ILLS 42505. Holotype: IMI 45404.

Triadelphia heterospora C.A. Shearer & J.L. Crane,
Mycologia 63:247—249. 1971. Holotype: On
balsa wood [Ochroma pyramidale (A.J.
Cavanilles) I. Urban] block submerged in the
Patuxent River, Lower Marlboro, Maryland,
8.XII.1967, leg. C.A. Shearer & J.L. Crane
A-33, ILLS 34793. Isotypes: DAOM
126798, ILLS 34793a, IMI 144033, NY.
Paratype: Brighton Dam, Triadelphia
Reservoir, Maryland, 5.1.1968, leg. C.A.
Shearer and J.L. Crane, A-204, ILLS 34894,
culture from the type: ATCC 22772.

Trichocladium linderi J.L. Crane & C.A. Shearer,
Mycologia 70:866—-869. 1978. Holotype: On
balsa wood [Ochroma pyramidale (A.J.
Cavanilles) I. Urban] blocks submerged in the
Rhode River Estuary, Edgewater, Anne Arundel
County, Maryland, 23.X1.1971, leg. C.A.
Shearer CS-186-9, ILLS 37019. Isotype: NY.
Paratypes: ILLS 37042, ILLS 37043.

Trichocladium moenitum J.L. Crane & C.A. Shearer,
Mycologia 70:871. 1978. Holotype: On
submerged decayed wood and filter paper in
petri dish, Elvira Cypress Swamp (Deer
Pond), Johnson County, Illinois, 21.11.1975,
leg. J.L. Crane & C.A. Shearer 588-1, ILLS
36986. Isotype: NY. Paratype: ILLS 37028.

May 1992 Catalog of Types

Triposporium batistae J.L. Crane & K.P. Dumont,
Canadian Journal of Botany 53:844-845.
1975. Isotype: Saprophytic on a decayed pod
(?), along the Rio Nueve Pasos, Dr. Luis
Roure’s property near Rosario, Puerto Rico,
elevation 140 m, 17.VI.1970, leg. R.P. Korf et
al., ILLS 34960. Holotype: NY. CUP-PR-
4179, ILLS 4179.

Tripterospora latipes N. Lundqvist, Botaniska
Notiser 122:592-593. 1969. =Zopfiella
latipes (N. Lundqvist) D. Malloch & R.F.
Cain, Canadian Journal of Botany 49:876.
1971. Isotype: Denmark, Sjaelland:
Kjobenhavn, Botanical Garden, isolated from
compost soil from a greenhouse, 26.11.1968,
leg. A. Kjoller, ILLS 35139. Paratype: ILLS
34539. Holotype: UPS.

Tubercularia ulmea J.C. Carter, Phytopathology
37:246. 1947. Holotype: On branches and
trunks of Ulmus pumila L., Onarga, Iroquois
County, Illinois, 16. VIII.1939, leg. J.C.
Carter, ILLS 29559.

Vanbeverwijkia spirospora V. Agnihothrudu,
Transactions of the British Mycological
Society 44:53. 1961. Isotype: On decaying
wood, Jorhat, Assam, India, 18.VIII.1958, leg.
H.K. Phukan 168, ILLS 34901 as a micro-
scopic preparation from HCIO. Holotype:
Mycological Herbarium, Tocklai Experiment
Station, Assam, India.

Varicosporium giganteum J.L. Crane, American
Journal of Botany 55:999. 1968. Isotypes:
From a foam sample, Androscoggin River,
Errol, Coos County, New Hampshire,
5.1X.1966, leg. J.L. Crane 190C-66, ILLS
34469, ILLS 39803, ILLS 39940. Holotype:
NY, culture from the type: ATCC 18147.

Verticillium rhizophagum L.R. Tehon & H.L.
Jacobs, Bulletin of the Davey Tree Expert
Company 6:16. 1936. Holotype: On U/mus
americana L., Dayton, Ohio, [X.1934, leg.
P.R. Grimes, ILLS 28858.

Zopfiella lundqvistii C.A. Shearer & J.L. Crane,
Transactions of the British Mycological
Society 70:456. 1978. Holotype: A dried
culture isolated from balsa wood and blocks
(Ochroma pyramidale (A.J. Cavanilles) I.
Urban), submerged in Elvira Cypress Swamp
(Deer Pond), Johnson County, Illinois,
28.V1.1974, leg. C.A. Shearer & J.L. Crane
CS-460-1, ILLS 36931. Isotype: NY, culture
from the type: ATCC 34976..

Zygorrhynchus verruculosus L.R. Tehon, Transac-
tions of the Illinois State Academy of Science
36:109-110. 1943. Holotype: Isolated from
dead elm roots (U/mus americana L.) , Peoria,
Peoria County, Illinois, Summer 1941, leg.
E.P. Metcalfe, ILLS 28927.

549

Literature Cited

Bailey, L.H., and E.Z. Bailey. 1976. Hortus third. A Little, E.L. 1979. Checklist of United States trees

concise dictionary of plants cultivated in the (native and naturalized). Agricultural Hand-
United States and Canada. MacMillan book No. 541. Washington, DC. 375 pp.
Publishing Company, New York. 1,290 pp. Porter, K.I., and C.J. Koster, eds. 1970. World list
Brown, P., and G.B. Stratton, eds. 1963. World list of scientific periodicals. New periodical titles
of scientific periodicals published in the years 1960-1968. Butterworths, London. 603 pp. +
1900-1960. William Clowes and Sons, indexes.
London and Beccles. Three volumes. Stafleu, F.A., and R.S. Cowan. 1976-1988.
Farr, E.R., J.A. Leussink, and F.A. Stafleu. 1979. Taxonomic literature. A selective guide to
Index Nominum Genericorum (Plantarum). botanical publications and collections with
Bohn, Scheltema & Holkema, Utrecht dr. W. dates, commentaries, and types. Second
Junk b.v. Publishers, The Hague. Three edition. Bohn, Scheltema & Holkema,
volumes. Utrecht/Antwerpen dr. W. Junk b.v. Publish-
Fernald, M.L. 1950. Gray’s manual of botany. ers, The Hague, Boston. Seven volumes.
Eighth edition. American Book Company, Willis, J.C. 1973. A dictionary of the flowering
New York. 1,632 pp. plants and ferns. Eighth edition. Revised by
Greuter, W., H.M. Burdet, W.G. Chaloner, V. H.K. Airy Shaw, Cambridge University Press,
Demoulin, R. Grolle, D.L. Hawksworth, D.H. London and New York. 1,245 pp. + LXVI
Nicolson, P.C. Silva, F.A. Stafleu, E.G. Voss, pp.)

and J. McNeill. 1988. International code of
botanical nomenclature adopted by the
Fourteenth International Botanical Congress,
Berlin, July-August 1987. Koeltz Scientific
Books, Konigstein, Federal Republic of
Germany. (Regnun Vegetabile Volume 118).
328 pp.

Appendix. Type Specimens Missing from ILLS Herbarium

Holotype Paratype
Name (Accession number) (Accession number)
Actinothyrium gloesporioides L.R. Tehon 2972 3671
Cryptostictis inaequalis L.R. Tehow & G.L. Stout 13698

Macrophoma smilacinae L.R. Tehon & G.L. Stout 20001
Viacrophoma zeae L.R. Tehon & E.Y. Daniels 1247

*

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