ILLUSTRATED

0
Rust Byunai
Genera
 Illustrated =

Genera of
Rust Fungi
Third Edition

George B. Cummins and Yasuyuki Hiratsuka

> II
CAB INTERNATIONAL

PRESS
The American Phytopathological Society
St. Paul, Minnesota
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Cover: Blastospora smilacis Dietel (Caeoma makinoi Kusano) on Prunus mume Sieb. & Zucc.
(reprinted with permission, from Kusano 1911)

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Copyright © 1983, 2003 by The American Phytopathological Society

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Cummins, G.B.; Hiratsuka, Y. 2003. Illustrated genera of rust fungi. Third ed. American Phyto-
pathological Society, St. Paul, MN. APS Press, St. Paul, MN.

ABSTRACT
0

he comprehensive introductory chapters include general information

onrust fungi: economic importance, symptoms, collection and identifi-
cation, host-rust relationships, phylogeny and relationship to other groups
of fungi, morphology of spore states and vegetative structures, life cycles,
cytology, host-rust relationships, and descriptions as well as a key to 13
families. The main body of the book consists of descriptions, illustrations,
and keys to 120 holomorph genera and 13 anamorph genera of rust fungi
worldwide. Each genus description includes information on the pertinent
microscopic features, type species, number of included species, hosts, dis-
tribution, relevant literature, and drawings or micrographs of diagnostic
features. Other new additions are eight full-page color plates containing
more than 60 photographs that illustrate various signs and symptoms
of rust fungi, an extensive list of general references on rust fungi, a list
of descriptive manuals and regional flora, a list of authors’ names and
recommended abbreviations, and a glossary of mycological terms.

RESUME
So

lee premiers chapitres présentent une introduction complete aux

champignons responsables de rouilles — importance économique,
symptomes, échantillonnage et identification, relations héte-rouille,
phylogénie, relations avec d’autres groupes de champignons, morphologie
des types de spores et des structures végétatives, cycles biologiques,
cytologie et descriptions —, ainsi qu’une clé d’identification couvrant 13
familles. La partie principale de l’ouvrage se compose de descriptions,
d’illustrations et de clés pour 120 genres d’holomorphes et 13 genres
d’anamorphes de champignons responsables de rouilles répartis dans le
monde entier. Les descriptions des différents genres fournissent des
précisions sur les caractéristiques microscopiques d’intérét, les especes-
types, le nombre d’especes, les hétes, la répartition et les publications
pertinentes, et s’accompagnent de dessins ou de micrographies des
caracteres diagnostiques. Huit planches couleur pleine page réunissant
plus de 60 photographies montrant différents signes et sympt6mes de
rouilles ont été ajoutées, ainsi qu’une liste de guides descriptifs et de publi-
cations sur des flores régionales, une liste de noms d’auteurs et de leurs
abréviations recommandeées et un glossaire de termes de mycologie.

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 CONTENTS

Morphology, Lite Gyclesrand Cy tolocy ofRust Fungi@ieon. 2 bo ee eee os

Definitions and morphology onspole staltccn mrtg ee eee tee et Joe Mowe ree eo:
SPETIMO SONU ea Na na ta Mea ea a i ata a WA dee Racer asa 0 bcc hs
PASO NUEOQWA RIG.$3 8O 88h Go oud CON aeohag (bhce ch.CRE Bo) conte c.CAeye ee aa aac ere ne
Drediniuint reese eee een er er es len ayeee A are aA aeuaa ame Ta
Telia re ened re nee ee eee es reno eee Oe Lace, oA adn Gas ce an) ead Do 2
Ba Sichuan ysat ieease Cee eee Ra ark ig goat Baad incin GL and ake id Mew 4).5
Sporemorpholopyegerarecs see tate ey ehcown Sweets, rete lain EUS I eG
DuUrface SeUl PIUle, OlSPOles aeata Tema ere eee, rhe, Ag ee ees ees eee ys
KSOUI POLES Jacga eee te Mee ete eee an Ne BASE esac goto eh tne Ge BeSytea 8, 9
Hyphaeand havistotiay awa ete ace es tertg socal RIRAemen ols fhe oe Cae se nase tien ele ass
Lifecyde Variations! maw). r tia TM Oe ee, Beene ches ice Er Poe Skee
HeWa)aVoyan avebgbl CSWeefe (Schr tte ds Boecn heSPT We er. Crm Peer at ar A a a ee
Theconcept elcorelated species andal raiZsciel silaWane sun yee neta, ee es es
Cytologyand nucteamey cles ait a tewcpbmtoniges ager Mee acral amaato es 2 ae
Literature: cited Saas eee eee ee ee I a a is ame aie me Se ke a

eee
EERE
EERE
SERRE
PEREPESERE
EPP
BEES
Anamorph Genera of Rust Fung <->. geo et ee ee eS ee en ee ee ee eee 36
Key to:anamorph:cenera: © os 515. swe meu peceencae satpro Sermons: fone 0+ cara ea yemriee, VoNeaee eee 36
Deschiptions of anamorph genera: o. 8. cee ane sored oie ee See cage nee 37
Materaturre cited teres ee rhs ps eee terme ee rea tare tan gc ee Re eee ee 40

Eolomorph. Genera Of Rust Funet Mae ge ys te ee erences elo ey ae a 42

Keys to holomorphieeneta, by family. veces es ny yee rusuey yee coker arcs ae 42
DESCRIPTIONS AND ILLUSTRATIONS OF FHOLOMORPH GENERA. 25 ~~ -aafineanererement 49

Appendix 1. Reference Materials (with someannotations). 2.3.46. 4.25.5... 1.5 7. eee Oi

Général references On. TUStLunGis fe ucts on bee cece ten wee eee 191
Descriptive manualsand regional flora. |. | Gace: cee mentees een en ner pede) es ener ee 191

Appendix 2. Authors of Names of Rust Fungi and Suggested Abbreviations ............. 194

Glossafy ¢ ¢< £5. 2 deh fede ae A OSBa Sah yS Deeg et, eh i ee ee 203

List-of Geniera:s: s..¢ 6... et8 Soke Gig ea each a gee ee tec 206

Tid @xG 85-5 oss gassing ES etal Sa a ne SG se ee eae ae 210

vi
 PREFACE
oo

his manual is designed primarily to be used in and keys to 120 holomorph genera and 13 ana-
the identification of rust fungi and to introduce morph genera of rust fungi worldwide. Each genus
students to the spore states, life cycles, and morpho- description includes information on the pertinent
logical diversity of the Uredinales. The arrangement microscopic features, type species, number of in-
of the genera suggests our concept of relationships, cluded species, hosts, distribution, relevant litera-
as does the assignment of the genera to 13 families. ture, and drawings or micrographs of diagnostic
An alphabetical arrangement of the genera could features. Entire articles of original descriptions are
also have been used, and this doubtless would have cited, and specific pages where the description
pleased some persons, especially those concerned appears are in parentheses. We attempted to clarify
with specific genera; but, as an aid to identification, the status of anamorph genera (form genera) of rust
grouping by morphological similarity is more use- fungi by defining them morphologically, disregard-
ful. Possibly, one will need only to look at the illus- ing their position in the life cycle. In addition to tra-
trations to reach at least a tentative identification. In ditionally used anamorph genera, such as Aecidium,
fact, much can be learned about the Uredinales just Caeoma, Peridermium, Roestelia, and Uredo,we added
by looking at the drawings and pictures. Calidion, Elateraecium, Lecythea, Malupa, Milesia,
There is not now, and probably never will be, Petersonia, Uredostilbe, and Wardia. These morpho-
unanimity of opinion concerning the numbers of logically defined anamorph types are included in
genera and the characters that delimit them. This is the description of each holomorph genus having
because different emphasis is accorded to the vari- anamorph state(s) and are designated as aecia
ous structures and to the importance of the relation- Aecidium-type, aecia Uredo-type, uredinia Milesia-
ship of the host plants parasitized. Although this type, etc.
situation may be confusing, it is not undesirable be- Eight full-page color plates containing more than
cause it stimulates discussion and additional study. 60 photographs that illustrate various signs and
Generic identity usually is only a first goal; conse- symptoms of rust fungi have been added. An exten-
quently, we have listed pertinent literature follow- sive list of general references on rust fungi, a list of
ing each generic description. This literature may descriptive manuals and regional flora, a list of
provide the means to identify species or may refer to authors’ names and recommended abbreviations,
microscopic illustrations or to electron micrographs and a glossary of mycological terms relevant to rust
of structures (TEM) or surface ornamentation fungi have been provided. Abbreviations of names
(SEM). There are, however, genera that have so few of authors of fungi are those suggested by Kirk and
species (sometimes only one) or that are so little Ansell (1992), with significant numbers of additions.
known that there is no special literature. There is still much to know about rust fungi and
The comprehensive introductory chapters include we sincerely hope that this publication will stimu-
general information on rust fungi: economic impor- late taxonomic and other studies of rust fungi by
tance, symptoms, collection and identification, present and future scientists.
host-rust relationships, phylogeny and relationship
to other groups of fungi, morphology of spore states LITERATURE CITED
and vegetative structures, life cycles, cytology, and Kirk, P.M.; Ansell, A.E., compilers. 1992. Authors of fungal
names. Int. Mycol. Inst., CAB Int., Kew, Surrey. 95 p.
descriptions and a key to 13 families. The main body
of the book consists of descriptions, illustrations,

vii
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 ACKNOWLEDGMENTS
oo

e acknowledge and thank former and present experience as a specialist in taxonomy of rust fungi;
Editor-in-Chief of APS Press, Dr. Sharon M. she performed a substantive technical editorial re-
Douglas and Dr. Randy C. Ploetz, for supporting the view, and her contributions significantly improved
publication of this book. The Publication Coordina- the quality of the end product. Special thanks are
tor of APS Press, Ms. Karen Cummings, coordinated due to Ms. Elaine Schiewe, whose considerable
activities at APS Press, and Dr. Wesley Chun acted talent in design and typesetting transformed our
as the senior editor of this book for APS Press. manuscript, photographs, and drawings into this
We especially thank Dr. R. Bandoni and Dr. Roger attractive book. The library staff, Ms. Edith Hopp,
S. Peterson for their detailed and constructive Ms. Debbie Oranchuk, and Ms. Denise Leroy, pro-
review comments. We are deeply grateful to them vided support, searching for and acquiring many
for their enthusiasm and encouragement through- reference materials.
out this process. Many scientists were consulted during the prepa-
We thank Northern Forestry Centre, Canadian ration of this book. Complete and thorough expert
Forest Service, for generously supporting this pro- reviews of selected chapters, provision of illustra-
ject from the beginning and providing substantial tions as well as critical review, advice and encour-
resources throughout the various stages of produc- agement were all given without hesitation. We
tion. We appreciate the encouragement and support thank and acknowledge the following individuals:
of the senior management, and we especially thank Dr. Eric Allen, Dr. Reinhard Berndt, Dr. Pablo
Mr. Boyd Case, Dr. Surj Malhotra, and Mr. Steve Buritica, Dr. Anibal Carvalho, Dr. P. (Chak)
Price. The publication staff are acknowledged for Chakravarty, Dr. José Dianese, Dr. Mario
their unwavering enthusiasm and support through- Figueiredo, Dr. Don Gardner, Dr. Joe Hennen, Dr.
out the many stages of this project. Ms. Brenda José Hernandez, Dr. Naohide Hiratsuka, Ms. Kim
Laishley was the managing editor for this project. Jakubec, Dr. Makoto Kakishima, Dr. Shigeru
She oversaw and contributed to the editorial Kaneko, Dr. Keizo Katsuya, Dr. Ken Mallett, Mr.
reviews and production of the publication. Her Paul Maruyama, Dr. Franz Oberwinkler, Dr.
superb skills, enthusiasm, and encouragement Yoshitaka Ono, Dr. Jack Parmelee, Dr. Denise
carried the project to its successful completion. Dr. Santiago, Dr. Shoji Sato, Dr. Marcus Scholler, Dr.
Patricia Crane is recognized for her extensive Helen Sotao, Dr. Aki Tsuneda, and Dr. Y. Yamaoka.
 eer
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 INTRODUCTION

“&

WHAT ARE THE RUST FUNGI? 2. many species need two unrelated groups of host
plants to complete their life cycle (heteroecious
The rust fungi are a unique and interesting group of life cycle), although others can complete their life
fungi traditionally classified in the order cycles on a single kind of host plant (autoecious
Uredinales of the class Basidiomycetes. The last life cycle); and
implies that, in their reproductive cycle, they pro- . species usually have narrow and specific host
duce basidia and basidiospores. More recent sug- ranges.
gestions regarding the taxonomy of rust fungi and
the phylogenetic position of rusts in relation to other
groups of Basidiomycetes are discussed in the next
RELATIONSHIP OF RUST FUNGI
section. The name rust fungi comes from the fact TO OTHER GROUPS OF FUNGI
that rust sori (sporulating structures) are often
Historically, many different classifications have
orange or reddish-orange. With some experience,
been proposed for fungi. These are summarized in
rust fungi are not difficult to recognize as rusts or
Table 4 of the 8" edition of the Dictionary of fungi
even to identify to the family and genus, even
(Hawksworth et al. 1995), including the relative
though they are not as conspicuous as mushrooms.
position of rust fungi (Uredinales). Traditionally,
In their natural habitat, rust fungi are obligate
rusts have often been classified together with smuts
biotrophic parasites of living plants, although a few
(Ustilaginales) under Teliomycetes (Talbot 1971;
species are now successfully cultured on artificial
Julich 1981), but recent molecular and ultra-
media (axenic culture). Rust fungi parasitize a wide
structural investigations have revealed that rusts
range of host plants, including ferns, conifers, and
are only distantly related to some of the smuts and
angiosperms (both mono- and dicotyledonous), and
are a part of a very old lineage of fungi that includes
recently a rust (Uredo vetus J.F. Hennen) has been dis-
pathogens of insects and plants such as
covered and described for the first time on a species
Eocronartium and Septobasidium (Berres et al. 1995;
of Lycopodiophyta (Selaginella sp.) (Hennen 1997).
Boehm and McLaughlin 1989; Frieders and
About 5000 species of rust fungi have been recog-
McLaughlin 2001; McLaughlin et al. 1995; Swann
nized, but the group is estimated to have more than
and Taylor 1993, 1995a, b). Ono and Hennen (1983)
7000 species (Hawksworth et al. 1995). About 300
suggested a strong relationship between rust genera
generic names have been proposed (Laundon 1965),
of the family Chaconiaceae and primitive
of which between 100 and 125 have been considered
auriculariaceous genera such as Herpobasidium,
as well-recognized genera (Cummins 1959;
Platycarpa, Jola, Eocronartium, and Kriegeria,
Cummins and Hiratsuka 1983; Thirumalachar and
although Hiratsuka (1990) was hesitant to include
Mundkur 1949a, b, 1950a, b; Hiratsuka 1955). In this
these genera in rusts for morphological reasons.
manual, 120 holomorph genera as well as 13
Some other genera considered to be related to rust
anamorph genera (form genera) are described and
fungi are Helicobasidium, Helicogloea, Rhodotorula,
illustrated.
Sporobolomyces, Sporidiomyces, Mixia, Bensingtonia,
The rust fungi of Europe, North America, Japan, Agaricostilbum, Erythrobasidium, and Sakaguchia.
New Zealand, Argentina, Brazil, and Australia have These fungi all have simple septal pores without
been reasonably well catalogued, but significant septal pore caps, and possess similar spindle pole
numbers of new species and perhaps genera are still body structures (Swann and Taylor 1995a).
expected, especially in tropical and subtropical With the advance of modern techniques such as
regions of Central and South America, Africa, and molecular analysis and ultrastructural examinations
southeast Asia. together with comparative morphology, many new
The rust fungi have three unique features: theories and suggestions have been proposed re-
1. up to five or six morphologically and function- garding the interrelationships among different groups
ally different spore states exist in the life cycle of a of fungi. Swann and Taylor (1995a, b) and Swann
single species; et al. (2001) separated the class Urediniomycetes,
including the rusts (Uredinales), from class 1995a) (Nishida et al. 1995). However, this fungus is
Ustilaginomyces and class Hymenomycetes under considered phylogenetically far from the fern rusts
division Basidiomycota. They placed the rusts in the (Sjamsuridzal et al. 1999). Sjamsuridzal et al. (1999)
class Urediniomycetes together with the studied rDNA sequence data of fern rusts and other
Septobasidiales, various species formerly placed in members of Urediniomycetes (sensu Swann and
the Ustilaginales (Microbotryum, Sphacelotheca, Taylor 1995a) and concluded that fern rusts are not
Rhodosporidium) or Auriculariales (Eocronartium, primitive among rust fungi.
Helicobasidium, Platygloea, Phleogena), and other
fungi of uncertain relationship (Pachnocybe,
Agaricostilbum, and Mixia). Within the class ECONOMIC IMPORTANCE OF
Urediniomycetes, true rusts (Uredinales) account THE RUST FUNGI
for over 95% of the species and more than 75% of the
genera. These three classes in the division Rust fungi as a group are among the most economi-
Basidiomycota (basidiomycetes) are based on ribo- cally important pathogens of many native and culti-
somal RNA analyses, and biochemical and _ultra- vated plants. Unlike other kinds of plant pathogens,
structural characters. Ina recent discussion, Bauer et which tend to attack weakened or poorly growing
al. (2001) and Swann et al. (2001) endorsed the topo- plants, rust fungi parasitize fresh tissues of vigor-
logical presentation showing the interrelationship ously growing plants. For this reason, problems
of the three groups of the Division Basidiomycota with rust fungi tend to increase with intensive and
originally proposed by Begerow et al. (1997), as extensive monoculture of many economically im-
shown in Figure 1. Most authors currently agree that portant agricultural, horticultural, and forest crops.
rusts (Uredinales) are a well-defined monophyletic Diseases caused by rust fungi are the major concerns
group of plant parasitic fungi with unique charac- and sometimes limiting factors for successful culti-
teristics such as possession of spermogonia, vation of such internationally important crops as
pleomorphic life histories, strict parasitism, lack of wheat, corn, coffee, and pine. It is interesting to note,
clamp connections, and similar ultrastructure of the however, that no rust fungus has been found on rice
septum and spindle pole body. or potato.
The following diseases caused by rust fungi are
considered to be economically important:
PHYLOGENY OF THE
On agricultural and horticultural crops
RUST FUNGI
Wheat stem rust (Puccinia graminis Pers.)
As obligate parasites of green plants, rusts are con- (Fig. 2A, B)
sidered to have coevolved with their hosts. Accord- Wheat leaf rust (Puccinia recondita Roberge ex
ingly, the species in three genera (Uredinopsis, Desm.) (Fig. 2C)
Milesina, and Hyalopsora) that parasitize ferns (II, Crown rust of oats (Puccinia coronata Corda var.
Ill') and Abies (0, I?) were thought to be the most avenae Fraser & Ledingham) (Fig. 2D)
primitive group of rusts because their two host Wheat stripe rust (Puccinia striiformis Westend.)
groups were considered relatively primitive (Ando Corn leaf rusts (Puccinia sorghi Schwein.,
and Katsuya 1984; Leppik 1953, 1956, 1959, 1965; P. polysora Underw.)
Savile 1955). However, Savile (1955, 1968) suggested Flax rust (Melampsora lini (Pers.) Lév.)
that most primitive rusts were derived from Coffee leaf rust (Hemileia vastatrix Berk. &
Taphrina-like ancestors on ancient tropical woody Broome) (Fig. 2E)
ferns, not the relatively recent herbaceous ferns that Mint rust (Puccinia menthae Pers.)
coexist with Abies species in the northern hemi- Sugarcane rusts (Puccinia kuehnii E.J. Butler,
sphere. It is interesting to note that a fern parasite, P. melanocephala Syd.)
Mixia osmundae (T. Nishida) Kramer, which was Chickpea rust (Uromyces ciceris-arietinus Jacz.)
originally placed in the Taphrinales, is now consid- Cotton rust (Puccinia cacabata Arthur & Holw.)
ered to be a primitive basidiomycete and is accom- Bean rust (Uromyces appendiculatus (Pers.) Unger)
modated within the simple septate basidiomycete Bamboo stem rust (Stereostratum corticioides
lineage (Urediniomycetes, sensu Swann and Taylor (Berk. & Broome) Magnus) (Fig. 2F)

' TI = uredina; III = telia.

> : ;
~ 0 =spermogonia; | = aecia.
 Soybean rust (Phakopsora pachyrhizi H. & P. Syd. Mulberry rust (Aecidium mori Barclay)
(Fig. 2G) (Fig. 3F)
Peanut rust (Puccinia arachidis Speg.) (Fig. 2H) Poplar and willow leaf rusts (Melampsora spp.)
Sunflower rust (Puccinia helianthi Schwein.) (Fig. 3G)
Onion rust (Puccinia allii Rudolphi) Eucalyptus rust (Puccinia psidii G. Winter) (Fig. 3H)
Peach and plum rust (Tranzschelia discolor
(Fuckel) Tranzschel & Litwinow) On forest trees
Birch—Thujopsis rust (Blastospora betulae White pine blister rust (Cronartium ribicola
S. Kaneko & Hirats. f. =Caeoma deformans J.C. Fisch. ex Rabenh.) (Fig. 4A)
(Berk. & Broome) Tubeuf) (Fig. 3A) Hard pine stem rusts (Cronartium flaccidum
Japanese apricot rust (Blastospora smilacis Dietel (Alb. & Schwein.) G. Winter, C. comandrae
=Caeoma makinoi Kusano) (Fig. 3B, 7) Peck (Fig. 4B), C. comptoniae Arthur, C.
Koa rusts (Aftelocauda koae (Arthur) Cummins coleosporioides Arthur (Fig. 4C), C. quercuum
and Y. Hirats., A. angustiphylloda Shirai ex Miyabe (incl. C. fusiforme Hedgc.
D.E. Gardner, Endoraecium acaciae Hodges & N.R. Hunt), C. orientale S. Kaneko,
& D.E. Gardner) (Fig. 3C) Endocronartium harknesstt (Moore) Y.
Rose rust (Phragmidium rosae-multiflorae Dietel) Hirats. (Fig. 4D), E. pini (Lév. emend
(Fig. 3D) Kleb.) Y. Hirats.)
Astilbe rust (Pucciniostele clarkiana (Barclay) Spruce needle, bud and cone rusts
Dietel (Fig. 3E) (Chrysomyxa spp.) (Fig. 5A—G)

Calocera viscosa

Tremella
mesenterica

Doassansia
epilobii

Ustilago hordei
Thecaphora
seminis-convolvuli

Urocystis ranunculi
Boletus rubinellus Entorrhiza casparyana
Cortinarius stuntzii
Entorrhiza aschersoniana
Marasmius delectans

Agaricus arvensis Kriegeria eriophori

Septobasidium Microbotryum violaceum

carestianum
Fulvisporium restifaciens
Aurantiosporium subnitens
sch ~ Ustilentyloma fluitans
mompa _ a
Eocronartium ie
muscicola eS en
Puccinia
K recondita graminis
Ep ea

UrediniomyceteS

FIGURE 1. Topology of division Basidiomycota. Obtained by neighbor-joining analysis of LSU rDNA sequences of
30 basidiomycetes (reprinted with permission, from Begerow et al. 1997). Shaded part indicates Uredinales.
FIGURE 2. Symptoms and signs of rust fungi of economically important plants. A. Puccinia graminis Pers. ex Pers.
(IL, III) on Triticum aestivum L. (wheat) (courtesy of Dr. J.F. Hennen). B. Puccinia graminis (0, 1) on Berberis
vulgaris L. (courtesy of Dr. M. Kakishima). C. Puccinia recondita Roberge ex Desm. (IL, IID) on Triticum
aestivum. D. Puccinia coronata Corda var. avenae Fraser & Ledingham (II, III) on Avena sativa L. (courtesy of
Dr. M. Kakishima). E. Hemileia vastatrix Berk. & Broome (II) on Coffea sp. (courtesy of Dr. J.F. Hennen).
F. Stereostratum corticioides (Berk. & Broome) Magnus (III) on Pleioblastus simonii (Carriere) Nakai (courtesy
of Dr. M. Kakishima). G. Phakopsora pachyrhizi H. & P. Syd. (II, III) on Glycine max Merr. (courtesy of Dr. M.
Kakishima). H. Puccinia arachidis Speg. (IL, 11) on Arachis hypogaea L.
 a betulae S. Kaneko & Hirats. f. (0, I) (Caeoma
FIGURE 3. Symptoms and signs of various rust fungi. A. Blastospor
& Zucc. (courtesy of Dr. M.
deformans (Berk. & Broome) Tubeuf) on Thujopsis dolabrata (Linn.f.) Sieb.
(courtesy of Dr. M. Kakishima).
Kakishima). B. Blastospora smilacis Dietel (II, III) on Smilax sieboldii Miq.
Acacia koa Gray (courtesy of Dr. D.E.
C. Atelocauda digitata (G. Winter) Cummins & Y. Hirats.(II, IM) on
Dietel (II, IIT) on Rosa multiflora Thunb. (courtesy of Dr. M.
Gardner). D. Phragmidium rosae-multiflorae
). E. Puccinioste le clarkiana (Barclay) Dietel (0, 1, II) on Astilbe microphylla Knoll (courtesy of Dr.
Kakishima
a occidentalis H.S. Jacks. (II, Il) on
M. Kakishima). F. Aecidium mori Barclay (1) on Morus alba L. G. Melampsor
sp.
Populus balsamifera L. H. Puccinia psidii G. Winter (II, II]) on Eucalyptus
FIGURE 4. Symptoms and signs of pine stem rusts. A. Cronartium ribicola J.C. Fisch. ex Rabenh.
(1) on Pinus albicaulis
Engelm. B. Cronartium comandrae Peck (1) on Pinus contorta Dougl. var. latifolia Engelm. C. Cronartium
coleosporioides Arthur (I) on Pinus contorta var. latifolia. D. Endocronartium harknessii J.P. Moore)
Y. Hirats.
(III) on Pinus contorta var. latifolia.
 Fir needle rusts (various genera) (Fig. 5H) 1. Hypertrophy and hyperplasia (galls)
Pine needle rusts (Coleosporium spp.) (Fig. 6A—C) Cronartium quercuum Miyabe ex Shirai—eastern
Cedar—apple rust (Gymnosporangium juniperi- gall rust of hard pines (0, D)
virginianae Schwein.) C. orientale S. Kaneko—Asian hard pine gall
Juniperus—Rosaceae rusts rust (0, 1)
(Gymnosporangium spp.) (Fig. 6D-H) C. conigenum Hedgc. & N.R. Hunt—southwestern
pine cone rust (0, I)
By using the narrow and specific host ranges of C. strobilinum (Arthur) Hedge. & G. Hahn—
rust fungi, attempts have been made to control certain southern pine cone rust (0, I)
noxious weeds. The following are examples of inter- Endocronartium harknessii (Moore) Y. Hirats.—
continental introduction of rusts to control weeds: western gall rust of hard pines (III) (Fig. 4D)
e rush skeletonweed (Chondrilla juncea L.) by
2. Fasciation (witches’ brooms)
Puccinia chondrillina Bubak & Syd., introduced
from the United States to Australia (Hasan and Chrysomyxa arctostaphyli Dietel—yellow witches’
broom of spruce (Picea) (0, I) (Fig. 5C)
Wapshere 1973);
Chrysomyxa woronini Tranzschel—bud rust of
blackberry (Rubus spp.) by Phragmidium
spruce (Picea) (0, I) (Fig. 5G)
violaceum (Schultz) G. Winter, introduced from
Melampsorella caryophyllacearum J. Schrot.—fir
Europe to Chile (Oehrens 1977);
(Abies) broom rust (0, D)
acacia (Acacia saligna (Labill.) Wendl.) by Uromy-
Atelocauda koae (Arthur) Cummins & Y. Hirats.—
cladium tepperianum (Sacc.) McAlpine, introduced
broom rust of Acacia koa Gray (II, II)
from Australia to South Africa (Morris 1989);
Blastospora betulae S. Kaneko & Hirats. f. (Caeoma
e water hyacinth (Eichhornia sp.) by Uredo
deformans (Berk. & Broome) Tubeuf) on
eichhorniae Gonz.-Frag. & Cif., introduced from
Thuja and Thujopsis (0, I) (Fig. 3A)
Argentina to the southern United States
(Charudattan and Conway 1975; Charudattan 3. Malformation
et al. 1978); Blastospora smilacis Dietel (Caeoma makinoi
e rubber-vine weed (Cryptostegia grandiflora Roxb. Kusano) on Prunus (0, I) (Fig. 7)
ex R. Br.) by Maravalia cryptostegiae (Cummins) Y. Pucciniastrum goeppertianum (J.G. Kuhn) Kleb.
Ono, potential introduction from Madagascar on Vaccinium (III) (Fig. 5H)
to Australia (Evans 1993; Evans and Fleureau
4. Pseudoflower formation
1993):
Puccinia monoica Arth. on Arabis (0, I) (Roy 1993)
Watson (1991) lists 17 species of rusts now under Uromyces pisi (Pers.) Wint. on Euphorbia (0, 1)
investigation to be introduced to non-native loca- (Pfunder and Roy 2000)
tions to control noxious weeds, mainly in Australia
but also in the United States and Canada. HOST-RUST RELATIONSHIP
Use of native species of rusts to control weeds has
been attempted; varying degrees of success have There are definite patterns of specificity between
been achieved with the following species: rust fungi and the plants they parasitize. Some
e nut sedge (Cyperus esculentus L.) by Puccinia genera of the rust fungi, especially Puccinia and
canaliculata (Schwein.) Lagerh., in the United Uromyces, comprise species that are capable of para-
States (Phatak et al. 1988), and sitizing plants of many families, but many genera
e Canada thistle (Cirsium arvense (L.) Scop.) by appear to be definitely restricted to certain plants.
Puccinia punctiformis (F. Strauss) Rohl. (Dyer et al. Host restriction may, in heteroecious species, apply
1982; Turner et al. 1981; Van Den Ende et al. to both phases of the life cycle or to only one phase.
1987). These restrictions may be taxonomically useful be-
cause of the shortcuts to identification that they pro-
vide. Whether or not exceptions are known, there is
SYMPTOMS CAUSED BY always the possibility that they occur. When trying
RUST FUNGI to separate genera on their own merits and without
regard to the host, it becomes obvious that too much
Plants infected with rust fungi often exhibit con- emphasis has sometimes been given to the host rela-
spicuous symptoms. The major types of symptoms tionship. Some examples of host-rust association
caused by rust fungi are listed below: are discussed below:
FIGURE 5. Symptoms and signs of spruce (Picea) and fir (Abies) rusts. A. Chrysomyxa ledicola (Arthur) H. & P. Syd.
(0, 1) on Picea glauca (Moench) Voss. B. Chrysomyxa ledicola (I, II) on Ledum groenlandicum Oeder.
C. Chrysomyxa arctostaphyli Dietel (0, 1) on Picea glauca. D. Chrysomyxa arctostaphyli (11) on Arctostaphylos
uva-ursi (L.) Spreng. E. Chrysomyxa pirolata G. Winter (I) on Picea glauca. F. Chrysomyxa pirolata (III) on Pyrola
asarifolia Michx. G. Chrysomyxa woroninii Tranzschel (0, I) on Picea glauca. H. Calyptospora goeppertiana J.G.
Kuhn (IL, 0-1) on Vaccinium vitis-idaea L. (left) and on Abies lasiocarpa (Hook.) Nutt. (right).
 (Dietel) Syd. (0, I) on Pinus contorta Dougl. var.
FIGURE 6. Symptoms and signs of tree rusts. A. Coleosporium asterum
latifolia Engelm. B. Coleospor ium clematidis Barclay (II, I) on Clematis sp. (courtesy of Dr. M. Kakishima).
asiaticum Miyabe ex Yamada
C. Coleosporium asterum (IIL, IIL) on Aster ciliolatus Lindl. D. Gymnosporangium
of Dr. M. Kakishima ) E. Gymnospo rangium clavipes (Cooke & Peck)
(0, D on Pyrus pyrifolia Nakai (courtesy
nelsoni Arthur (III) on
Cooke & Peck (0, I) on Amelanchier alnifolia (Nutt.) Nutt. F. Gymnosporangium
rangium gaeumannii Zogg ssp.
Juniperus horizontalis Moench (left hydrated, right unhydrated). G. Gymnospo
H. Gymnosporangium clavariiforme
albertensis Parmelee (IL, II1) on Juniperus communis L. var. depressa Pursh.
(Pers.) DC. (IID) on Juniperus communis var. depressa.
 FIGURE 7. Symptoms and signs of Blastospora smilacis Dietel (Caeoma makinoi Kusano) (0, 1) on Prunus
mume Sieb. & Zucc. (reprinted with permission, from Kusano 1911).

10
Host-restricted heteroecious rust fungi juniper or its relatives, the rust is a Gymno-
e There are three genera that alternate between sporangium, (b) if there is a rust of pine needles, it is a
ferns and fir (Abies spp.), without known species of Coleosporium, or (c) if there is a rust on
exceptions. These are Uredinopsis, Milesina, and stems or cones of pine, the genus is Cronartium or
Hyalopsora. Endocronartium. In temperate regions, rusts of the
e Aecial states of Coleosporium and Cronartium Rosaceae (excluding those with pomes or drupes)
occur on pines (Pinus spp.). Coleosporium pro- are likely to be species of Phragmidium, but in warmer
duces its aecia on the needles; Cronartium pro- regions, Hamaspora and Gerwasia are probable. Species
duces its aecia on stems or cones but never on of Uromyces are common on legumes, but Puccinia is
needles. Neither genus is restricted as to telial hosts. rare on this family. Species with two-celled telio-
¢ Most species in the genus Chrysomyxa produce spores on legumes mostly belong in Uropyxis or
their aecial state on needles, buds, or cones of
Sorataea. Although such shortcuts are useful, they
spruce (Picea spp.) and the telial state on plants need to be used with caution because exceptions
exist and one should not be led so far astray as to
of the Ericaceae and close relatives.
look upon the host as a characteristic of the fungus.
e Melampsora species are either heteroecious or
autoecious. Many of the heteroecious species
produce telia on Salix or Populus of the COLLECTION, IDENTIFICATION,
Salicaceae, and they produce aecia on Abies,
AND PRESERVATION
Larix, Picea, Pinus, or Pseudotsuga. Others para-
sitize Ribes, Saxifraga, or Allium. The autoecious
OF RUST FUNGI
species are not host-restricted. Proper procedures for collecting and preserving
¢ Most of the species of Gymnosporangium are specimens are important. It is essential to collect
heteroecious and have aecial states on dicots, enough plant parts, such as flowers, fruits, and
especially Pomoideae of the Rosaceae such as uninfected leaves, together with the infected parts,
Malus, Pyrus, Sorbus, Crataegus, and Cydonia, so that the host may be accurately identified. This is
and telial states on the conifer genera Calo- especially important for such plants as sedges and
cedrus, Chamaecyparis, Cupressus, and most on grasses. Without the inflorescence these plants are
Juniperus. These rusts are unique in that the nearly impossible to identify.
coniferous plants serve as the telial rather than A good hand lens is recommended for use in the
the aecial hosts. field to check for the presence of different spores or
to distinguish insects or other pathogens from rust
Host-restricted autoecious rust fungi fungi.
e Cumminsiella, a relatively small genus of All spore states can be preserved dry; therefore,
macrocyclic species, appears to be restricted to specimens can be handled in the same way as
species of Berberis and Mahonia. flowering plant specimens, using plant presses.
e Hamaspora is another small genus, whose Dried and pressed specimens are usually kept in
species occur in Africa, Asia, and Australia. specimen packets bearing labels with the essential
They parasitize members of the genera Rosa data, such as the name of the host, name of the
and Rubus, especially the latter. collector, the date and location where found, and
e Phragmidium, a relatively large genus, is re- preferably a collection number. Packet specimens
stricted to plants of the Rosaceae. Most species may be attached to standard herbarium sheets and
occur on Potentilla, Rosa, and Rubus. stored as are flowering plants. Some mycologists
e Ravenelia is one of the largest genera and is prefer to use a standardized 12.7 x 17.8 cm (5 x 7 in.)
primarily tropical or semitropical in distribu- packet and store them in boxes or filing cases. Either
tion. Most (perhaps all) of its species parasitize system works; it is a matter of personal choice.
legumes (Fabaceae). For the identification of these fungi, it is essential
e The species of Prospodium occur only on to have a compound microscope with a magnifica-
the Bignoniaceae (predominantly) and tion of at least 400x and equipped with a calibrated
Verbenaceae. micrometer. Spores are best mounted in lactophenol
or a similar non- or slow-drying medium with some
Some of these rust—host associations are useful in clearing action. Stains are generally not needed.
providing shortcuts to acquaintance with the com- Size, surface ornamentation, shape, and color
moner genera. For example, (a) if there is a rust on should be recorded reasonably promptly. If stored

fil
for a long time, some spore walls change consider- Evans, H.C.; Fleureau, L. 1993. Studies on the rust, Maravalia
cryptostegiae, a potential biological control agent of rubber-
ably, as does size. Thin sections made free hand or
vine weed, Cryptostegia grandiflora (Asclepiadaceae:
with a microtome may be useful or necessary to Periplocoideae), in Australia. II. Infection. Mycopathologia
determine the position of a sorus in the host tissues 124:175-184.
or to observe the structure of the fungus. Use of a Frieders, E.M.; McLaughlin, D.J. 2001. The heterobasidiomycete
scanning electron microscope, if available, can moss parasites Jola and Eocronartium in culture: cytology,
reveal detailed spore markings. ultrastructure, and anamorph. Mycol. Res. 105:734-744.
Hasan, S.; Wapshere, A.J. 1973. The biology of Puccinia
If a comprehensive regional descriptive manual
chondrillina, a potential biological control agent of
is available, the host index may be useful to narrow skeltonweed. Ann. Appl. Biol. 74:325-332.
the possibilities, and then each likely species can be Hawksworth, D.L.; Kirk, P.M.; Sutton, B.C.; Pegler, D.N. 1995.
compared with the specimen being identified. The Ainsworth & Bisby’s dictionary of the fungi. 8'" ed. Int.
importance of host identification, at least to generic Mycol. Inst., CAB Int., Egham, UK. 616 p.
level, cannot be overemphasized (see Host-rust rela- Hennen, J.F. 1997. Uredo vetus sp. nov., the first record of a rust on
tionship section). Then with the help of a rust man- Selaginella, and the use of the name Uredo. Mycologia 89:801—
803.
ual, one may key out the genus and then refer to the
Hiratsuka, N. 1955. [Uredinological studies.] Kasai Publ. Co.,
literature cited to determine specific identity. In Tokyo. 382 p. (In Japanese.)
some cases, identification may be confirmed by
Hiratsuka, Y. 1990. Auriculariaceous “rust”. Rep. Tottori Mycol.
comparing the specimen with authentically identi- Inst. 28:25-30.
fied specimens or by sending them to specialists. Julich, W. 1981. Higher taxa of basidiomycetes. Bibl. Mycol. 85:1—
485.
LITERATURE CITED Kusano, S. 1911. On the chloranthy of Prunus mume caused by
Ando, K.; Katsuya, K. 1984. Evaluation of taxonomic significance Caeoma makinoi. Bull. Coll. Agric., Tokyo Imp. Univ. 2:287-326.
of morphological characteristics of rust fungi based on axenic Laundon, G.F. 1965. The generic names of Uredinales. Mycol.
cultures. Rep. Tottori Mycol. Inst. 22:124-132. Pap., Commonw. Mycol. Inst. 99:1-24.
Bauer, R.; Begerow, D.; Oberwinkler, F.; Piepenbring, M.; Berbee, Leppik, E.E. 1953. Some viewpoints on the phylogeny of rust
M.L. 2001. Ustilaginomycetes. Pages 57-83 in D.J. fungi. I . Coniferous rusts. Mycologia 45:46-74.
McLaughlin and P. Lemke, eds. Mycota. Vol. VIIB. Springer- Leppik, E.E. 1956. Some viewpoints on the phylogeny of rust
Verlag, Berlin, Heidelberg. fungi. IL Gymnosporangium. Mycologia 48:637—-654.
Begerow, D.; Bauer, R.; Oberwinkler, F. 1997. Phylogenetic studies Leppik, E.E. 1959. Some viewpoints on the phylogeny of rust
on nuclear large subunit ribosomal DNA sequences of smut fungi. III. Origin of grass rusts. Mycologia 51:512-528.
fungi and related taxa. Can. J. Bot. 75:2045-2056.
Leppik, E. 1965. Some viewpoints on the phylogeny of rust fungi.
Berres, M.E.; Szabo, L.J.; McLaughlin, D.J. 1995. Phylogenetic V. Evolution of biological specialization. Mycologia 57:6—22.
relationships in auriculariaceous basidiomycetes based on McLaughlin, D.J.; Berres, M.E.; Szabo, L.J. 1995. Molecules and
25S ribosomal DNA sequences. Mycologia 87:821-840. morphology in basidiomycete phylogeny. Can. J. Bot.
Boehm, E.A.; McLaughlin, D.J. 1989. Phylogeny and 73(Suppl. 1):S684-S692.
ultrastructure in Eocronartium muscicola: meiosis and basidial Morris, M.J. 1989. The use of plant pathogens as control agents for
development. Mycologia 81:98-114. South African weeds. In E.S. Delfosse, ed. Proc. VII Int. Symp.
Charudattan, R.; Conway, K.E. 1975. Comparison of Uredo Biol. Control Weeds, 6-11 Mar. 1989, Rome, Italy.
eichhorniae, the waterhyacinth rust, and Uromyces pontederiae. Nishida, H.; Ando, K.; Ando, Y.; Hirata, A.; Sugiyama, J. 1995.
Mycologia 67:653-657. Mixia osmundae: transfer from the Ascomycota to the
Charudattan, R.; McKinney, D.E.; Cordo, H.A.; Silveira-Guido, Basidiomycota based on evidence from molecules and mor-
A. 1978. Uredo eichhorniae, a potential biocontrol agent for phology. Can. J. Bot. 73(Suppl. 1):S660-S666.
waterhyacinth. Pages 210-213 in Proc. VI Int. Symp. Biol. Oehrens, E. 1977. Biological control of the blackberry through the
Control Weeds, Gainesville, 30 Aug—2 Sept. 1976, Univ. introduction of rust Phragmidium violaceum in Chile. FAO
Florida, Gainesville, FL. Plant Prot. Bull. 25:26-28.
Cummins, G.B. 1959. Illustrated genera of rust fungi. Burgess Ono, Y.; Hennen, J.F. 1983. Taxonomy of Chaconiaceous genera
Publ. Co., Minneapolis, MN. 131 p. (Uredinales). Trans. Mycol. Soe. Jpn. 24:369-402.
Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust Pfunder, M.; Roy, B.A. 2000. Pollinator-mediated interactions be-
fungi. Rev. ed. APS Press, St. Paul, MN. 152 p. tween a pathogenic fungus, Uromyces pisi (Pucciniaceae), and
Dyer, W.E.; Turner, S.K.; Fay, P.K.; Sharp, E.L.; Sands, D.C. 1982.
its host plant, Euphorbia cyparissias (Euphorbiaceae). Am. J.
Bot. 87:48-55.
Control of Canada thistle by a rust, Puccinia obtegens. Pages
243-244 in R. Charudattan and H.L. Walker, eds. Biological Phatak, S.C.; Sumner, D.R.; Wells, H.D.; Bell, D.K.; Glaze, N.C.
control of weeds with plant pathogens. John Wiley & Sons, 1988. Method for controlling yellow nutsedge using Puccinia
New York. canaliculata. US Dep. Agric. Patent 4-731-104. 1 p.
Evans, H.C. 1993. Studies on the rust, Maravalia cryptostegiae, a Roy, B.A. 1993. Floral mimicry by a plant pathogen. Nature
potential biological control agent of rubber-vine weed (London) 362:56-58.
(Cryptostegia grandiflora, Asclepiadaceae: Periplocoideae) in Savile, D.B.O. 1955. A phylogeny of the basidiomycetes. Can. J.
Australia. I. Life-cycle. Mycopathologia 124:163-174. Bot. 33:60-104.

2
Savile, D.B.O. 1968. Possible interrelationship between fungal Thirumalachar, M.J.; Mundkur, M.J. 1949a. Genera of rusts. I.
groups. Pages 649-675 in G.C. Ainsworth and A.S. Sussman, Indian Phytopathol. 2:65-101.
eds. The fungi. III. Academic Press, New York. Thirumalachar, M.J.; Mundkur, M.J. 1949b. Genera of rusts. II.
Sjamsuridzal, W.; Nishida, H.; Ogawa, H.; Kakishima, M.; Indian Phytopathol. 2:193-244.
Sugiyama, J. 1999. Phylogenetic positions of rust fungi para- Thirumalachar, M.J.; Mundkur, M.J. 1950a. Genera of rusts. III.
sitic on ferns: evidence from 185 rDNA sequence analysis. Indian Phytopathol. 3:4—42.
Mycoscience 40:21—27. Thirumalachar, M.J.; Mundkur, M.J. 1950b. Genera of rusts. IV.
Swann, E.C.; Taylor, J.W. 1993. Higher taxa of basidiomycetes: an Indian Phytopathol. 3:203-204.
18S rRNA gene perspective. Mycologia 85:923-936. Turner, S.K.; Fay, P.K.; Sharp, E.L.; Sands, D.C.1981. Resistance of
Swann, E.C.; Taylor, J.W. 1995a. Phylogenetic perspectives on Canada thistle (Cirsium arvense) ecotypes to a rust pathogen
basidiomycete systematics: evidence from the 185 rRNA (Puccinia obtegens). Weed Sci. 29:623--624.
gene. Can. J. Bot .73(Suppl. 1): S862-S868. Van Den Ende, G.; Frantzen, J.; Timmers, T. 1987. Teliospores as
Swann, E.C.; Taylor, J.W. 1995b. Phylogenetic diversity of yeast- origin of systemic infection of Cirsium arvense by Puccinia
producing basidiomycetes. Mycol. Res. 99:1205-1210. punctiformis. Neth. J. Plant Pathol. 93:233-239.
Swann, E.C.; Frieders, E.M.; McLaughlin, D.J. 2001. 2. Watson, A.K. 1991. The classical approach with plant pathogens.
Urediniomycetes. Pages 37-54 in DJ. McLaughlin and P. Pages 3-23 in D.O. TeBeest, ed. Microbial control of weeds.
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Springer-Verlag, Berlin.
Talbot, P.H.B. 1971. Principles of fungal taxonomy. Macmillan,
London. 274 p.

13
 MORPHOLOGY, LIFE CYCLES, AND
CYTOLOGY OF RUST FUNGI

DEFINITIONS AND normally associated with spermogonia,

MORPHOLOGY OF SPORE STATES and giving rise to dikaryotic vegetative
mycelium.
Five kinds of spore-producing structures are gener- Urediniospores (sometimes called uredospores
ally recognized as basic spore states of the rust or urediospores): repeating dikaryotic
fungi: spermogonium (pl. spermogonia), aecium mitospores produced on a dikaryotic
(pl. aecia), uredinium (pl. uredinia), telium (pl. mycelium and usually giving rise toa
telia), and basidium (pl. basidia). They are often dikaryotic mycelium on the same host.
designated by Roman numerals as 0, I, II, IL, and IV,
respectively. Ina few species, such as Puccinia vexans Terminology based on either system is the same
Farl., thick-walled, darkly pigmented urediniospores for many well-known species such as Puccinia
called amphispores are produced besides regular graminis, P. helianthi, Cronartium ribicola, and
paler-colored urediniospores, and in these species Melampsora lini, but differs in other situations. For
six morphologically and functionally different example, urediniospores (by the ontogenic system)
spore stages exist. Also, many rust species are of Chrysomyxa and Coleosporium, which are produced
known to produce secondary basidiospores, and on nonconiferous alternate hosts, are produced in
they are often called ballistospores. chains and have verrucose ornamentation and
Two different systems have been applied in the should be called aeciospores by the morphologic
definition and terminology of spore states. They are system. These genera would, then, have two aecio-
the morphologic system and the ontogenic system. spore states, one on the aecial host and one on the
The morphologic system, as defined by Laundon (1967a, telial host. Also, it is impossible to define teliospores
b) and Holm (1973), emphasizes the morphology of by their morphology because so much variation ex-
spores as the basis for defining spore states. In this ists among the various genera (Fig. 8). Thus, the only
system, aeciospores are defined as being produced possible definition is the spore state that gives rise to
in chains and usually having ornamentation tradi- basidia, which is the ontogenic definition. In this
tionally known as verrucose; urediniospores are de- manual, we use terminology based on the ontogenic
fined as being always unicellular and borne singly system. For further discussion regarding these sys-
on pedicels, and usually having ornamentation tems of terminology refer to papers by Laundon
traditionally known as echinulate (Laundon 1972). (1967a, 1972), Hiratsuka (1973b, 1975), and Durrieu
On the other hand, the ontogenic system, used by (1979).
Cummins (1959), following Arthur (1905, 1925,
1929), and expanded by Hiratsuka (1973, 1975), SPERMOGONIUM (pl. spermogonia)
emphasizes the position of the spore states in the life (=pycnium, pl. pycnia) (0)
cycle rather than clearly recognizable morphologi- Spermogonia are produced ona haploid thallus that
cal entities as the basis for spore terminology. Defi- results from infection by a basidiospore. They are
nitions of spore states according to the ontogenic minute, but because they occur in groups, often on
system (Hiratsuka 1973b) are as follows: discolored spots or hypertrophied tissues, they may
be obvious macroscopically. Besides, the spermogo-
Teliospores: basidium-producing spores nia typically are associated with visible aecia or
(probasidia, hypobasidia). telia. The spermogonia produce spermatia in a sweet-
Basidiospores: haploid, often binucleate (but ish exudate and the spermatia function as dikaryo-
not dikaryotic) spores produced on tizing elements. Spermatia are small, one-celled,
basidia. hyaline spores that have little or no diagnostic
Spermatia: dikaryotizing elements. value, but morphological types of spermogonia have
Aeciospores: nonrepeating spores produced as been considered to be dependable as characteristics
a result of dikaryotization, thus in sori useful in suprageneric taxonomy (Hiratsuka and

14
 Wee Se Rg
2 ital

FIGURE 8. Morphological variations of teliospores. A. Pileolaria brevipes Berk. & Rabenh. B. Trachyspora intrusa
(Grev.) Arthur. C. Dasyspora gregaria (Kunze) Henn. D. Diorchidiella australis (Speg.) J.C. Lindq. E. Ravenelia
mera Cummins (=R. lonchocarpicola Speg. var. mera (Cummins) J.W. Baxter. F. Chrysella mikaniae Syd.
G. Kuehneola uredinis (Link) Arthur. H. Prospodium appendiculatum (G. Winter) Arthur. I. Sphaerophragmium
acaciae (Cooke) Magnus. J. Cumminsina clavispora Petr. K. Melampsorella symphyti Bubak. L. Chrysocelis lupini
Lagerh. & Dietel. M. Goplana dioscoreae Cummins. N. Lipocystis caesalpiniae (Arthur) Cummins.
O. Phragmidiella markhamiae Henn. P. Didymopsora africana Cummins. Q. Dietelia verruciformis (Henn.)
Henn. R. Dietelia portoricensis (Whetzel & Olive) Buritica & J.F. Hennen.
Cummins 1963). Twelve morphological types were AECIUM (pl. aecia) (1D
recognized and six groups designated (Figs. 9, 10, Aecial primordia (Fig. 12) as well as spermogonia
11) by Hiratsuka and Hiratsuka (1980). The types are are usually produced on a haploid thallus that
characterized by such features as position in the host results from infection by a basidiospore. Aecia are
tissue, shape of the hymenial layer, presence or preceded or accompanied by spermogonia, and
absence of bounding structures, and whether both structures are produced on the same thallus.
growth is determinate or indeterminate. Spermogo- The aecial initials receive nuclei of compatible
nia may not develop and sometimes are omitted mating types from spermatia through trichogynes
from life cycles. (flexuous or receptive hyphae) that extend through
Mims and Doudrick (1996) reported the ultra- spermogonia or stomata from the bases of aecial
structure of spermatial formation of Cronartium primordia. This process leads to dikaryotization of
quercuum (Berk.) Miyabe ex Shirai f.sp. fusiforme and the aecial primordia, then dikaryotic aeciospores
described annellidic formation of spermatia. develop. Aeciospores are unicellular. Upon germi-
A recent work by Anikster et al. (1999) showed nation, they produce dikaryotic mycelium, which,
that, in several species of Puccinia and Uromyces, in turn, produces either uredinia or telia, but not
when spermogonial nectar was transferred between aecia again.
spermatia of compatible mating types, a distinct cap
formed on one end of the spermatium. The signifi- UREDINIUM (pl. uredinia) (ID
cance of this mating-type-specific cap formation is Uredinia are produced on a dikaryotic mycelium,
not known, but it is likely an early significant event and the urediniospores produce dikaryotic myce-
in the processes leading to dikaryotization. lium when they germinate. Thus this spore state is
Historically, spermogonia were called pycnia or the state that is repeated in the life cycle of the rust
pycnidia because of the superficial resemblance to fungi. The first crop of uredinia to be produced in
pycnidia (often flask-shaped conidiomata), but these the life cycle develops on mycelium that results
names were suggested when the function of the from the germination of aeciospores. However, the
structure was not clearly understood and we think mycelium resulting from the germination of ure-
spermogonium is a more appropriate name for the diniospores may produce more uredinia, but this
structure. The name spermogonium was first used usually ultimately gives rise to telia and teliospores.
by Tulasne in 1851 (Arthur 1904), but Arthur (1905)
suggested the term pycnium instead of spermo- TELIUM (pl. telia) (111)
gonium because he argued that this structure is very Teliospores are the most important spore state in
inefficient and does not have an infectious or sexual generic distinctions. According to the International
function. After the discovery of the function of the code of botanical nomenclature (Greuter et al. 2000),
structure (Craigie 1927), Arthur himself, who teleomorphs of rust fungi are teliospores, and the
created the term pycnium, used the term spermogo- names based on teliospores are valid holomorphic
nium again (Arthur 1934). Many contemporary names covering all morphs. Therefore, telia repre-
mycologists prefer to use the term pycnium rather sent the indispensable spore form of the rust fungi
than spermogonium (Laundon 1974; Savile 1971, for nomenclature, and most genera are based
1976), because they recognize these structures as largely on the morphology of this sorus and its
spermatia-producing as well as receiving structures spores, the teliospores. Spermogonia, aecia, and
through flexuous hyphae. Craigie reported and uredinia may be omitted in the various types of life
emphasized the presence and possible function of cycle reduction; the telium (teleomorph) is never
so-called flexuous hyphae as receptive structures. omitted if the rust concerned can be considered as a
However, we agree with Cummins (1959) and Cain perfect (holomorphic) fungus. Teliospores produce
(1972) that aecial primordia (Fig. 12), not flexuous basidia and almost always basidiospores upon
hyphae, should be considered the receptive germination. The teliospore is equivalent to the
structures for dikaryotization, thus justifying the probasidium, and basidium to metabasidium in
use of the term spermogonium (Hiratsuka and general terminology is in other groups of lower
Hiratsuka 1980). Spermogonia may not develop and basidiomycetes, but we prefer to retain the terms
sometimes are absent from the life cycles, e.g., in telium and basidium for the rust fungi.
the case of many homothallic rusts and in those Telia may vary greatly in morphology (Figs. 8,
species in which dikaryotization occur through 13). The spores may be scattered in the mesophyll
anastomoses of compatible hyphae within host (Uredinopsis), within the epidermal cells (Milesina,
tissues. Thekopsora, Naohidemyces), grouped in subepidermal

16
 mec TTT ss

FIGURE 9. Morphological groups and types of spermogonia (reprinted with permission, from Hiratsuka and
Hiratsuka 1980).
 FIGURE 10. Spermogonia. A. Type 2 spermogonium of Coleosporium jonesii (Peck)
Arthur. B. Type 9 spermogonium of Cronartium ribicola J.C. Fisch.
C. Type 10 spermogonium of Phragmidium barnardi Plowr. & G. Winter.
D. Type 11 spermogonium of Phragmidium sanguisorbae (DC.) J. Schrot.
(reprinted with permission, from Hiratsuka and Cummins 1963).

18
FIGURE 11. Cross sections of rust-infected tissues showing spermogonia and aecia. A. A type 2 spermogonium of
Coleosporium asterum (Dietel) Syd. on Pinus contorta Dougl. var. latifolia Engelm. B. Type 3 spermogonia of
Melampsora albertensis Arthur on Larix sp. C. Type 4 spermogonia and aecia of Puccinia recondita Roberge.
ex Desm. on Thalictrum sp. D. A type 9 spermogonium of Cronartium ribicola J.C. Fischer ex Rabh. on Pinus
albicaulis Engelm. E. A type 7 spermogonium and an aecium of Phakopsora ampelopsidis (Dietel & P. Syd.)
Cummins & Ramachar on Meliosoma myriantha Sieb. & Zucc. F. A type 7 spermogonium of Pileolaria
brevipes Berk. & Rabenh. on Rhus sp. G. A type 12 spermogonium and an aecium of Caeoma peltatum C.G.
Shaw III & C.G. Shaw on Phyllocladus trichomanides D. Don. H. A type 12 spermogonium and an aecium of
Mikronegeria alba Oehrens & R.S. Peterson.

19
 Xv i iyf

NS ag S396 -\

wee

FIGURE 12. A semischematic drawing of spermogonia (S) and aecial primordia (AP) of Puccinia graminis Pers.
(reprinted with permission, from Buller 1950).

20
FIGURE 13. Teliospores and urediniospores of selected rust species. A. Teliospores of Puccinia graminis Pers. on
Triticum aestivum L. B. Teliospores of Puccinia coronata Corda. on Avena sativa L. C. Hyaline teliospores
and colored urediniospores of Porotenus memorae F.C. Albug. on Memora sp. D. Teliospores and
amphispores of Puccinia vexans Farl. on Bouteloua curtipendula (Michx.) Torr. E. Teliospores of Newinia
thaiana Kakish. & Y. Ono on Stereospermum cylindricum Pierre ex P. Don. F. Teliospores of Melampsora
albertensis Arthur on Populus tremuloides Michx. G. Urediniospores of Puccinia arachidis Speg. on Arachis
hypogaea L. H. Teliospores of Phragmidium rosae-multiflorae Dietel on Rosa multiflora Thunb. (A, D courtesy
of Dr. J.F. Hennen; B, E, H, of Dr. M. Kakishima; C, of Dr. M. Figueiredo; G, of Dr. S. Sato).

21
crusts that are one spore deep (Melampsora) or in them semi-internal basidia. Peterson and Oehrens
erumpent cushions (most species of Puccinia), or (1978) discuss this matter in relation to Mikronegeria
extruded as hairlike columns (Cronartium). Some alba Oehrens & R.S. Peterson.
telia simulate aeciospores or urediniospores and Anikster (1983) reported that due to a mitotic
their true nature can be determined only by germi- nuclear division in the basidiospores, binucleate
nating the spores, for example, Endocronartium basidiospores are common in rusts.
(Peridermium-type telia), Endophyllum and Mono-
sporidium (Aecidium-type telia), and Endoraecium
(Uredo-type telia). These are the endocyclic genera. SPORE MORPHOLOGY
Endocyclic genera are also theoretically possible as
derivatives of Roestelia, Elateraecium, Milesia, and all SURFACE SCULPTURE OF SPORES
other anamorphic genera. In general, the terminology and definitions to de-
Teliospores sometimes are called resting or scribe surface sculpture of rust spores have not been
winter spores. This is because many common rust very precise. In part, this is because the patterns
fungi, e.g., Puccinia graminis and Melampsora lini, tend to intergrade. For example, echinulation and
have teliospores capable of surviving unfavorable verrucosity become nearly indistinguishable when
periods such as extreme temperatures. In fact, many the cones and warts differ only slightly in the degree
teliospores will germinate only after surviving such of sharpness or roundness (Fig. 14A—C); the verru-
periods. However, there are rust fungi, perhaps half cae may be rounded or blocklike or have straight or
of all of them, whose teliospores germinate as soon annulate sides (Fig. 14D-G). Striate verrucosity
as fully formed. Teliospores of species of Coleosporium, merges into interrupted or complete ridges (Fig.
Cronartium, Chrysomyxa, Kuehneola, and Chaconia, 14H-J). Other terms used to describe spore surface
among others, have nonresting teliospores. It is ax1- sculpture are rugose (Fig. 14K), labyrinthiform or
omatic that rust fungi that lack resistant teliospores cerebroid (Fig. 14L), and reticulate (Fig. 14M, N).
compensate with some development elsewhere in Scanning electron microscopy doubtless will permit
the life cycle to permit survival in unfavorable the introduction of more precise definitions and
periods, e.g., resistant mycelium embedded in terms (Sato and Sato 1982; Lee and Kakishima
woody host tissue, as in Cronartium spp. or some 1999a, b).
species having pigmented resistant urediniospores. The surface sculpturing of spores contributes
little to generic definition, but it is used extensively
BASIDIUM (pl. basidia) (IV) to delimit species. Most species of Puccinia and
Basidia and basidiospores have received limited Uromyces on the monocots have smooth teliospores,
attention in classification, perhaps because taxo- but many other patterns occur in these genera. Telio-
nomic studies usually involve herbarium speci- spores of the Melampsoraceae, Coleosporiaceae,
mens with nonviable teliospores. Basidiospores do Cronartiaceae, Phakopsoraceae, Mikronegeriaceae,
differ in size and shape, however, and these differ- and Chaconiaceae have smooth walls. Most telio-
ences have been used in descriptions of species in spores of the Uropyxidaceae, Pileolariaceae, and
Coleosporium (Cummins 1978; Kaneko 1981) and in Sphaerophragmiaceae have some sort of ornamen-
Cronartium (Kaneko 2000). With more study, they tation. There is little consistency in the Pucciniaceae,
may prove useful in defining genera. Phragmidiaceae, and Raveneliaceae.
Several genera (Coleosporium, Chrysella, Classifying spore surfaces simply as echinulate
Chrysopsora, Goplana, and Ochropsora) have so-called or verrucose, as commonly practised, is misleading.
internal basidia. These basidia are not produced by In aeciospores and urediniospores of Chrysomyxa
germination of teliospores, but whole contents of and Coleosporium, the surface is generally called
the cells divide internally to produce usually four- verrucose, but the pattern of distribution as well as
celled basidia. Species having this type of basidium the structure of the verrucae varies greatly (Kaneko
formation can be considered as not having telio- 1981; Crane 2001). Sato and Sato (1985) have distin-
spores (probasidia), but in this manual we call them guished eight morphologically different types of
internal basidia. Another group of genera produce surface ornamentation of aeciospores within 30
basidia somewhat in between external and internal genera of rust fungi. Lee and Kakishima (1999a, b)
basidia formation. In these genera, the apical part of studied the surface structure of aeciospores and
the teliospore continues to grow and produce peridial cells of the genus Gymnosporangium and
basidia. They are Mikronegeria, Blastospora, recognized 12 types of spore surface structure and
Chrysocelis, and Chrysocyclus. In this book we call 10 types of peridial cell wall.

DZ
FIGURE 14. Surface sculpture of spores. A—C. Echinulate. D-G. Verrucose. H. Striately verrucose. I. Ridged.
J. Interruptedly ridged. K. Rugose, roughened but with no discrete pattern. L. Labyrinthiform or cerebroid.
M. Pseudoreticulate, irregular and incomplete netting. N. Reticulate, regular netting, ridges and pits.

Cee,
OWVOD
FIGURE 15. Some examples of germ pore arrangements and spore shapes (mainly urediniospores). A. Scattered
pores. B. Equatorial pores. C. Bizonate pores. D. Superequatorial, unizonate pores. E. Basal unizonate
pores. F. Subequatorial pores. G. Equatorial pores with pore caps (exaggerated). H. Equatorial unizonate
pores. I. Equatorial pores in lateral angles. J. Equatorial pores of depressed globoid or helmet-shaped
spores. K. Unizonate equatorial pores of transversely ellipsoid spores.

ZS
GERM PORES
Arrangement and number of germ pores are useful
and have constant characteristics within species
(Cummins 1936). Most, and possibly all, anamorph
spores have germ pores, but they may be indistinct
or indiscernible, especially in spores with colorless
or lightly pigmented walls. Pores may appear as
clear spots, or may be detected because of small
lens-like caps, or the wall may bulge inward slightly
where pores occur. The number of pores ranges
from 1 (uncommon) up to 15. The principal arrange-
ments are shown in Figure 15, along with shapes of
spores, but this does not necessarily imply a direct
relationship between these characters. Scattered
and equatorial arrangements are the commonest.
When pores are scattered, a range of 6 to 10 is
common; when they are zonate, 2 to 5 are common.
It is speculated that numerous germ pores represent
primitive forms, and equatorial and few germ pores
represent advanced forms (Cummins 1936).

HYPHAE AND HAUSTORIA

Typically, hyphae of rust fungi are intercellular in
both monokaryotic (0, I state) and dikaryotic states
(I, II states). Intercellular hyphae of all rusts are
hyaline and septate without clamp connections.
Detailed morphological studies have been carried FIGURE 16. D-haustorium of Melampsora lini (Ehrenb.)
out on septal pores of many rusts species (Coffey et Lév. HMC, haustorial mother cell; NR, neck
al. 1972; Littlefield and Bracker 1971; Harder and region; HC, host cytopalasm; N, two nuclei
Chong 1984, 1991). All reported that septa of rust (reprinted with permission, from Coffey et
al. 1972).
fungi are simple, i.e., the septum wall simply tapers
towards the central pore. This type is similar to that
in several other groups of Urediniomycetes. Haustoria of haploid infections (0, I states) are
Hymenomycetes have more complicated septal called M-haustoria. In general, they are more fila-
structures referred to as dolipore septa. mentous and irregular in shape than D-haustoria
Although direct evidence is still difficult to (Allen 1932; Pady 1935). Electron microscope exami-
obtain, it is generally assumed that haustoria are nations have revealed that M-haustoria lack clear
involved in the absorption of nutrients from living differentiation of neck regions (Kohno et al. 1977;
cells of the host plants. Haustoria of dikaryotic Harder 1978; Gold et al. 1979).
hyphae are called D-haustoria and have been
studied extensively. They are composed of a
haustorial mother cell (HMC), which adheres to the LIFE CYCLE VARIATIONS
host cell wall before penetration, a neck region,
usually with an electron-dense band called the Depending on the number of spores states, three
neckband, and the main body of the haustorium in basic types of life cycle are recognized: macrocyclic,
the living cell (Fig. 16). Good general reviews of this demicyclic, and microcyclic. The macrocyclic life
subject can be found in Harder and Chong (1984, cycle has all spore states, the demicyclic lacks the
1991) and Littlefield and Heath (1979). Shapes uredinial state, and the microcyclic lacks both the
(simple or branched) and sizes of haustoria as well aecial and the uredinial states, thus possessing only
as haustorial neck morphology are considered to be telia and sometimes spermogonia. Spermogonia may
useful for systematic considerations of some groups be absent from any type.
of rusts (Berndt 1996; Berndt et al. 1994; Berndt and In the macrocyclic and demicyclic life cycles, the
Oberwinkler 1995, 1997). rust may be either host alternating (heteroecious),

24
i.e., the aecial state is on one kind of plant, but the E. pini (Pers. emend Kleb.) Y. Hirats.—resin top
telial state is on a different and unrelated plant, or rust
non-host-alternating (autoecious), i.e., the aecial and Endoraecium acaciae Hodges & D.E. Gardner—
telial states are on the same kind of plant. Koa rust
Combining the number of spore states and presence A form of Gymnoconia peckiana (Howe in Peck)
or absence of host alternation, the following basic vari- Trotter—raspberry rust
ations are recognized. Examples are given. A hy-
phen between spore states indicates host alternation. The species with endocyclic life cycles have
spores that are morphologically like aeciospores or
1. Heteromacrocyclic: (0), I-IL, II, IV urediniospores, but they germinate to produce
Coleosporium asterum (Dietel) Syd.—pine basidia; thus they are considered to be teleomorphs
needle rust (teliospores). No consensus for dealing with the
Cronartium ribicola J.C. Fisch. ex Rabenh.—white nomenclature of endocyclic rusts exists at the pres-
pine blister rust ent time, and this causes difficulties. Obviously, the
Melampsorella caryophyllacearum J. Schrot.—fir end forms are derived from aecial or uredinial
broom rust anamorph states of parental species by the reduc-
Puccinia graminis Pers.—black stem rust of wheat tion of life cycles. Three options can be considered to
Uromyces striatus J. Schrot.—alfalfa rust deal with the nomenclature and taxonomy of
endocyclic genera and species:
2. Automacrocyclic: (0), I, I, HL, IV
Melampsora lini (Pers.) Lév.—flax rust 1. They are obviously morphologically anamorphs;
Phragmidium mucronatum (Pers.) Schtdl—rose therefore, we should use anamorphic generic
rust names. This practice is widely used by forest
Pileolaria terebinthi (DC.) Castagne—Pistacia rust pathologists, who call endocyclic pine stem rusts
Puccinia helianthi Schwein.—sunflower rust by their anamorph names (e.g., Peridermium
Uromyces appendiculatus (Pers.) Unger—common harknessti and P. pini), but this is obviously unsat-
bean rust isfactory in view of the International code of botani-
cal nomenclature (Greuter et al. 2000). If spores of
3. Heterodemicyclic: (0), I-III, IV endocyclic rusts are considered as teleomorphic,
Chrysomyxa arctostaphyli Dietel—spruce broom they cannot be called by anamorphic names.
rust
2. After all, they are derivatives of parental genera;
Gymnosporangium juniperi-virginianae
therefore we could use parental generic names.
Schwein.—cedar-—apple rust
Theoretically this seems to be the most logical so-
Puccinia interveniens Bethel—Malvaceae-Stipa lution. However, we cannot predict the parental
rust genera of many endocyclic species, and this creates
Calyptospora goeppertiana Kuhn—fir needle rust room for much speculation and causes confusion.
4. Autodemicyclic: (0), I, IL, IV 3. We could create endocyclic genera and put
Gymnoconia peckiana (Howe in Peck) Trotter— endocyclic species in these genera. These points
orange rust of Rubus are discussed in Hiratsuka (1968) to deal with
endocyclic pine stem rusts, and he concluded
5. Microcyclic: (0), HL, IV that the best solution is to establish endo-genera
Coleosporium pinicola (Arthur) Arthur—pine corresponding to morphological types of aecia.
needle rust To complicate the matter further, such genera as
Puccinia malvacearum Bert. ex Mont.—hollyhock Endophylloides, Dietelia and Pucciniosira are cer-
rust tainly derived from aecial states, but they are not
Ravenelia pringlei Cummins—Acacia broom rust currently recognized as endocyclic genera. It is
Tranzschelia thalictri (Chevall.) Dietel— interesting that Buritica and Hennen (1980) in-
Thalictrum rust cluded an endocyclic genus, Endophyllum, among
genera of Pucciniosireae (=Pucciniosiraceae) in
6. Endocyclic: (0), II, IV their treatment, but recently Buritica and Hennen
Endophyllum semperotvi (Alb. & Schwein.) (1994) attempted to put all endocyclic genera into
de Bary—Sempervivum rust a separate family, Endophyllaceae. In this manual,
Endocronartium harknessii (J.P. Moore) we placed endocyclic genera close to the families
Y. Hirats.—western gall rust of suspected parental genera or groups of genera.

25
 ABNORMAL LIFE CYCLES in all respects except that it produces one-celled
teliospores. Arthur (1934) recognized the existence
Although the life cycle of a species is usually stable, of correlated species in several genera. When a rust
a species may not produce certain spores, depending species with a longer life cycle has been reduced toa
on the conditions. For example, late in the season, microcyclic species, there is a definite rule or pattern
urediniospores of Cronartium species are not pro- as to the habit of its telial state. The pattern is called
duced, and only teliospore production is observed. Tranzschel’s Law.
Furtherfhore, the following rust species are known The essence of Tranzschel’s Law is this: the telia
to have variant life cycles that do not fit in the above of microcyclic species simulate the habit of the aecia
categories. There are two reports indicating that real of the parental macrocyclic species and occur on the
teliospores as well as spores morphologically like aecial host plants of the latter. For example, if the
aeciospores both germinate to produce basidia and aecial state of a macrocyclic species is systemic, the
basidiospores; they are in Puccinia pampeana Speg. derived microcyclic species also will have systemic
(Endophyllum pampeanum (Speg.) J.C. Lindq. = Aecid- mycelium. Puccinia monoica Arthur and P. holboellii
ium pampeanum Speg.) (Hennen et al. 1984) and (Hornem.) Rostr. (=P. thlaspeos C. Schub.) illustrate
Puccinia japonica Dietel (Kakishima et al. 1984). Vari- this situation. The heteroecious P. monoica, with
ation in the number of spore states in rust life cycles uredinia and telia on June grass (Koeleria cristata (L.)
is acommon phenomenon (Jackson 1931), but these Pers.), produces its systemic aecial state on species
rusts are unusual because two kinds of teleomorphs of Arabis and some other genera of the Cruciferae.
are produced in a species. Atelocauda koae (Arthur) Puccinia holboellii, a microcyclic species, produces its
Cummins & Y. Hirats. is known as a demicyclic systemic telia on Arabis and relatives. The telio-
species, but it produces nonfunctional aeciospores. spores of the macrocyclic and microcyclic species
Infections are initiated only by germinating telio- are morphologically similar, but the microtelia (telia
spores (basidiospores), indicating that the fungus is of microcyclic rusts) mimic the habit of the aecia of
in the process of becoming endocyclic (Chen et al. the macrocyclic parent. The microtelia, in this
1996). example, do not have a peridium, but the aecial
appearance is sufficiently obvious that the species
was once named Aecidium holboellii Hornem.
THE CONCEPT OF CORRELATED Another interesting pair of species that illustrates
SPECIES AND TRANZSCHEL’S LAW Tranzschel’s Law is Puccinia coronata Corda (crown
rust), a heteroecious macrocyclic species with aecia
It is generally believed that the heteroecious on Rhamnus and uredinia and telia on oats, and
macrocyclic life cycle is the primitive existing one Puccinia mesnieriana Thum., a microcyclic species
and that the shortened cycles are derivative types with telia on Rhamnus. In both these species the
(Jackson 1931). Species of differing cycles but obvi- teliospores have apical digitate projections (the
ously close relationships often are called correlated corona, which gives crown rust both its common
species. It is assumed that the species with reduced name and its Latin epithet). Many more examples
life cycles are descendants of macrocyclic species. are known, most of them involving telioid
Arthur (1934), in the Manual of the rusts in the United microcyclic species, but endo-type microcyclic
States and Canada, made extensive use of the concept species also occur. Examples are the heteroecious
of correlated species. Puccinia interveniens Bethel, P. demicyclic Puccinia graminella Dietel & Holw. and
graminella Dietel & Holw., P. sherardiana Korn., and Endophyllum tuberculatum Arthur & Fromme, and
Endophyllum tuberculatum (Ellis & Kellerm.) Arthur Gymnoconia peckiana (Howe in Peck) Trotter and its
& Fromme represent a group of correlated species, endocyclic form.
including a heterodemicyclic, an autodemicyclic, a
telioid microcyclic, and an endo- or aecidioid micro-
cyclic form. In this series the presumed macrocyclic CYTOLOGY AND
parent is unknown but, if extant, it should produce NUCLEAR CYCLES
aecia on a malvaceous host and telia on a Stipa
(Poaceae). In the case of Puccinia recondita Roberge The basic nuclear cycle associated with the life cycle
ex Desm. (P. rubigo-vera G. Winter), Uromyces of a macrocyclic rust is illustrated in Figure 17.
dactylidis G.H. Otth is also included as a correlated Spermogonia are produced on haploid mycelia, and
species, in addition to microcyclic species of haploid spermatia are produced. Spermatia are
Puccinia. It corresponds to the macrocyclic Puccinia responsible for the dikaryotization of haploid aecial

26
 PINE ALTERNATE HOST

Aeciospores
Urediniospores

see 6 Teliospores

NUCLEAR
FUSION

FIGURE 17. Nuclear cycle of a heteroecious rust (Cronartium spp.).

primordia of the compatible mating type. In Miyabe ex Shirai (now considered to be C. orientale S.
addition to dikaryotization by spermatia, several Kaneko (Kaneko 2000)) is the bifactorial tetrapolar
other ways of dikaryotization are observed, e.g., (a) type, based on crossing experiments on segmented
fusion of compatible haploid mycelia within the infected gall tissues. Based on mating experiments
host tissue in cases of multiple infection; (b) with haploid axenic cultures, Narisawa et al. (1994)
fusion of a dikaryotic mycelium (with unilateral also concluded that Puccinia coronata Corda var.
transfer of one nuclear type) with a haploid myce- coronata has a bifactorial tetrapolar mating system.
lium in autoecious macrocyclic species such as After dikaryotization, dikaryotic aeciospores are
Puccinia helianthi Schwein.; (c) spontaneous produced in aecia. Germinating aeciospores pro-
dikaryotization not involving compatible mating duce a dikaryotic mycelium on which uredinia and
types in the case of homothallic species such as telia are formed. Urediniospores are the dikaryotic
Puccinia adoxae DC., P. buxi DC., P. malvacearum repeating spore state (conidia) and they initiate
Mont., and P. xanthii Schwein. Homothallic species dikaryotic mycelia. Teliospores (probasidia) are ini-
usually lack spermogonia. Since the first report of tially dikaryotic, but nuclear fusion (karyogamy)
heterothallism in a rust, shown with Puccinia occurs in the spore to produce diploid nuclei. Upon
helianthi Schwein. by Craigie (1927) and reviewed germination, teliospores produce basidia
by Buller (1950), mating systems of rust fungi were (metabasidia). Meiosis takes place in the basidium
considered to be bipolar. However, Yamazaki and to produce usually four haploid cells, each of which
Katsuya (1988) suggested that the mating system of germinates to produce haploid basidiospores.
a hard pine gall rust, Cronartium quercuum (Berk.) Anikster (1983) reported that basidiospores having

idl
two nuclei per spore due to mitotic nuclear divisions Coffey, M.D.; Palevitz, B.A.; Allen, P.J. 1972. The fine structure of
two rust fungi, Puccinia helianthi and Melampsora lini. Can. J.
are common among rust fungi; they occur in many
Bot. 50:231—240.
basidiomycetes. The basidiospores germinate and
Craigie, J.H. 1927. Discovery of the function of the pycnia of the
initiate haploid mycelia, which produce spermogo- rust fungi. Nature (London) 120:765-767.
nia and aecial primordia. Many variations in the
Crane, P.E. 2001. Morphology, taxonomy, and nomenclature of
nuclear condition of teliospores and basidia have the Chrysomyxa ledi complex and related rust fungi on spruce
been reported among microcyclic species, especially and Ericaceae in North America and Europe. Can. J. Bot.
endocyclic species (Jackson 1935; Olive 1953; 79:957-982.
Rajendren 1967; Hiratsuka 1968; Imazu et al. 1991; Cummins, G.B. 1936. Phylogenetic significance of the pores in
Gardner 1981, 1988, 1994; Chen et al. 1996). urediniospores. Mycologia 28:103-132.
Hiratsuka and Sato (1982) list at least eight different Cummins, G.B. 1959. Illustrated genera of rust fungi. Burgess
Publ. Co., Minneapolis, MN. 131 p.
variations in the nuclear condition of teliospores
and basidia, including the most common one de- Cummins, G.B. 1978. Rust fungi on legumes and composites in
North America. Univ. Arizona Press, Tucson, AZ. 424 p.
scribed above.
Durrieu, G. 1979. Problems of terminology concerning the cycle
patterns of Uredinales (Fungi). Bull. Trimest. Soc. Mycol. Fr.
LITERATURE CITED 953/939De
Allen, R-F. 1932. A cytological study of heterothallism in Puccinia
Gardner, D.E. 1981. Nuclear behavior and clarification of the
coronata. J. Agric. Res. 45:513-541.
spore stages of Uromyces koae. Can. J. Bot. 59:939-946.
Anikster, Y. 1983. Binucleate basidiospores—a general rule in rust
fungi. Trans. Br. Mycol. Soc. 81:624-626. Gardner, D.E. 1988. Teliospore germination of Puccinia vitata, an
endemic Hawaiian rust. Mycologia 80:590-593.
Anikster, Y.; Eilam, T.; Mittelman, L.; Szabo, L.J.; Bushnell, W.R.
1999. Pycnial nectar of rust fungi induces cap formation on Gardner, D.E. 1994. Teliospore germination and nuclear behavior
pycniospores of opposite mating type. Mycologia 91:858-870. of Puccinia rutainsulara, a microcyclic Hawaiian rust.
Mycologia 86:486-493.
Arthur, J.C. 1904. Taxonomic importance of the spermogonium.
Bull. Torrey Bot. Club 31:113-123. Gold, R.E.; Littlefield, L.J.; Statler, G.D. 1979. Ultrastructure of the
pycnial and aecial stages of Puccinia recondita. Can. J. Bot.
Arthur, J.C. 1905. Terminology of the spore structures in the
Uredinales. Bot. Gaz. 39:219-222.
57:74-86.
Arthur, J.C. 1925. Terminology of the Uredinales. Bot. Gaz. Greuter, W.; McNeill, J.; Barrie, FR.; Burdet, H.M.; Demoulin, V.;
80:219-223. Filgueiras, T.S.; Nicolson, D.H.; Silva, P.C.; Skog, J.E.; Trehane,
P.; Turland, N.J.; Hawksworth, D.L. 2000. International code
Arthur, J.C. 1929. The plant rusts (Uredinales). John Wiley and
of botanical nomenclature (Saint Louis code). Koeltz Scien-
Sons, New York.
tific Books, K6nigstein, Germany. 474 p.
Arthur, J.C. 1934. Manual of the rusts in United States and
Harder, D.E. 1978. Comparative ultrastructure of haustoria in
Canada. Purdue Res. Found., Lafayette, IN. Reprinted witha
uredinial and pycnial infections of Puccinia coronata avenae.
supplement by G.B. Cummins (1962). Hafner Publ. Co., New
Can. J. Bot. 56:214-224.
York. 438 p.
Berndt, R. 1996. Ultrastructure of the D-haustoria of Coleosporium Harder, D.E.; Chong, J. 1984. Structure and physiology of
spp. (rust fungi, Uredinales). Sydowia 48:263-272. haustoria. Pages 431-507 in W.R. Bushnell and A.P. Roelfs,
eds. The cereal rusts. Vol. I. Academic Press, New York.
Berndt, R.; Oberwinkler, F. 1995. Ultrastructure of the parasitic
interface of Pucciniastrum, Thekopsora, Naohidemyces, and Harder, D.E.; Chong, J. 1991. Rust haustoria. Pages 235-250 in K.
Calyptospora (Uredinales, Pucciniastraceae) in the dikaryotic Mendgen and D.E. Lesemann, eds. Electron microscopy of
stage. Mycoscience 36:51-59. plant pathogens. Springer-Verlag, Berlin.
Berndt, R.; Oberwinkler, F. 1997. Haustorial ultrastructure and Hennen, J.F.; Figueiredo, M.B.; Pimental, C.P.; Russomanno,
morphology of Melampsorella and Thekopsora areolata. O.M.R. 1984. The life cycle and taxonomy of Puccinia
Mycologia 89:698-705. pampeana Speg. and Endophyllum pampeanum (Speg.) Lindq.
on Capsicum spp. and other Solanaceae. Rep. Tottori Mycol.
Berndt, R.; Bauer, R.; Oberwinkler, F. 1994. Ultrastructure of the
Inst. 22:209-220.
host-parasite interface in the fern rusts Milesia, Uredinopsis,
and Hyalopsora (Pucciniastraceae, Uredinales). Can. J. Bot. Hiratsuka, Y. 1968. Morphology and cytology of aeciospores and
72:1084-1094. aeciospore germ tubes of host-alternating and pine-to-pine
Buller, A.H.R. 1950. Researches on fungi. Vol. VIL. The sexual pro- races of Cronartium flaccidum in northern Europe. Can. J. Bot.
cess in the Uredinales. Univ. Toronto Press, Toronto. 458 p. 46:1119-1122.

Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales, Hiratsuka, Y. 1973. The nuclear cycle and the terminology of
Pucciniaceae). Flora Neotrop. Monogr. 24:1-50. spore states in Uredinales. Mycologia 65:432—-443.
Buritica, P.; Hennen, J.F. 1994. Familia Phakopsoraceae Hiratsuka, Y. 1975. Recent controversies on the terminology of
(Uredinales) 1. Géneros anamorficos y teliomdrficos. Rev. rust fungi. Rep. Tottori Mycol. Inst. 12:99-104.
Acad. Colomb. Cienc. Exactas Fis. Nat. 19:47-62. Hiratsuka, Y.; Cummins, G.B. 1963. Morphology of the spermo-
Cain, R.F. 1972. Evolution of the fungi. Mycologia 64:1-14. gonia of the rust fungi. Mycologia 55:487-507.
Chen, W.Q.; Gardner, D.E.; Webb, D.T. 1996. Biology and life cycle Hiratsuka, Y.; Hiratsuka, N. 1980. Morphology of spermogonia
of Atelocauda koae, an unusual demicyclic rust. Mycoscience and taxonomy of rust fungi. Rep. Tottori Mycol. Inst. 18:257—
36:91-98. 268.

28
Hiratsuka, Y.; Sato, S. 1982. Morphology and taxonomy of rust Lee, S.K.; Kakishima, M. 1999b. Surface structures of peridial cells
fungi. Pages 1-36 in K.J. Scott and A.K. Chakravorty, eds. The of Gymnosporangium and Roestelia (Uredinales). Mycoscience
rust fungi. Academic Press, London, New York. 40:121-131.
Holm, L. 1973. Some notes on rust terminology. Rep. Tottori Littlefield, L.J.; Bracker, C.E. 1971. Ultrastructure of septa in
Mycol. Inst.10:183-187. Melampsora lini. Trans. Br. Mycol. Soc. 56:181-188.
Imazu, M.; Kakishima, M.; Katsuya, K. 1991. Morphology and Littlefield, L.J.; Heath, M.C. 1979. Ultrastructure of rust fungi.
nuclear cycle of Endocronartium yamabense. Trans. Mycol. Soc. Academic Press, New York. 277 p.
Jpn. 32:371-379. Mims, C.W.; Doudrick, R.L.1996. Ultrastructure of spermatia and
Jackson, H.S. 1931. Present evolutionary tendencies and the ori- spermatium ontogeny in the rust fungus Cronartiwm quercuum
gin of life cycles in the Uredinales. Mem. Torrey Bot. Club f.sp. fusiforme. Can. J. Bot. 74:1050-1057.
18:1-108. Narisawa, K.; Yamaoka, Y.; Katsuya, K. 1994. Mating type of iso-
Jackson, H.S. 1935. The nuclear cycle in Herpobasidium filicinum lates derived from the spermogonial state of Puccinia coronata
with a discussion of the significance of homothallism in var. coronata. Mycoscience 35:131-135.
Basidiomycetes. Mycologia 27:553-572. Olive, L.S. 1953.The structure and behavior of fungus nuclei. Bot.
Kakishima, M.; Sato, T.; Sato, S. 1984. Life cycle, morphology and Rev. 19:439-458.
cytology of Puccinia japonica on Anemone flaccida. Trans. Br. Pady,S.M. 1935. The role of intracellular mycelium in systemic in-
Mycol. Soc. 82:469-475. fection of Rubus with the orange-rust. Mycologia 27:618-637.
Kaneko, S. 1981. The species of Coleosporium, the causes of pine Petersen, R.H. 1974. The rust fungus life cycle. Bot. Rev. 40:453—
needle rusts, in the Japanese Archipelago. Rep. Tottori Mycol. 513.
Inst. 19:1-159. Peterson, R.S.; Oehrens, E. 1978. Mikronegeria alba (Uredinales).
Kaneko, S. 2000. Cronartium orientale, sp. nov., segregation of the Mycologia 70:321-331.
pine gall rust in eastern Asia from Cronartium quercuum. Rajendren, R.B. 1967. A new type of nuclear life cycle in Hemileia
Mycoscience 41:115-122. vastatrix. Mycologia 59:279-285.
Kohno, M.; Ishizaki, H.; Kunoh, H. 1977. Cytological studies on Sato, T.; Sato, S. 1982. Aeciospore surface structure of the
rust fungi. VI. Fine structure of infection process of Kuehneola Uredinales. Trans. Mycol. Soc. Jpn. 23:51-63.
japonica (Diet.) Dietel. Mycopathologia 61:35-42.
Sato, T.; Sato, S. 1985. Morphology ofaecia of the rust fungi. Trans.
Laundon, G.F. 1967a. Terminology in the rust fungi. Trans. Br. Br. Mycol. Soc. 85:223-238.
Mycol. Soc. 50:189-194.
Savile, D.B.O. 1955. A phylogeny of the basidiomycetes. Can. J.
Laundon, G.F. 1967b. The taxonomy of the imperfect rusts. Trans. Bot. 33:60-104.
Br. Mycol. Soc. 50:349-353.
Savile, D.B.O. 1971. Generic disposition and pycnium type in
Laundon, G.F. 1972. Delineation of aecial from uredinial states. Uredinales. Mycologia 63:1089-1091.
Trans. Br. Mycol. Soc. 50:189-194.
Savile, D.B.O. 1976. Evolution of the rust fungi (Uredinales) as
Laundon, G.F. 1974. Uredinales. Pages 247-279 in G.C.
reflected by their ecological problems. Evol. Biol. 9:137-207.
Ainsworth, E.K. Sparrow, and A.S. Sussman, eds. The fungi,
Yamazaki, S.; Katsuya, K. 1988. Experiments on selfing and recip-
an advanced treatise. Vol. IVB. Academic Press, New York,
rocal crossings in the pine gall rust fungus, Cronartium
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quercuum. Trans. Mycol. Soc. Jpn. 29:93-96.
Lee, S.K.; Kakishima, M. 1999a. Aeciospore surface structure of
Gymnosporangium and Roestelia (Uredinales). Mycoscience
40:109-120.

20,
 CLASSIFICATION OF RUST FUNGI
are

FAMILIES OF RUST FUNGI and Cummins (1963) questioned placing too much
emphasis on the morphology of the telial state. They
Two families, Melampsoraceae and Pucciniaceae, pointed to the importance of morphological types of
were once recognized by several authors, including spermogonia and recognized 11 types. Hiratsuka
Dietel (1928), Cunningham (1931), Arthur (1934), and Hiratsuka (1980) added type 12, and based ona
Bessey (1950), Hiratsuka (1955), and Azbukina study of 224 species in 73 genera, placed genera into
(1974), and were widely accepted for a long time. six groups. With emphasis on spermogonial type as
Gatimann (1949) accepted these two and added also well as other characteristics, Cummins and
the Pucciniastraceae, Cronartiaceae, Chryso- Hiratsuka (1984) proposed the following 13 families
myxaceae, and Coleosporiaceae. Wilson and (originally 14 but reduced to 13 by uniting Sphaero-
Henderson (1966) used Coleosporiaceae, phragmiaceae with Raveneliaceae) as associating
Melampsoraceae, and Pucciniaceae. Families have most of the genera in relatively natural groups. They
also been divided into subfamilies or tribes. The considered these as probably reflecting phylogen-
most recent such treatment is that of Azbukina etic relationship better than previous systems,
(1974), who divided the Melampsoraceae into eight which divided the rusts into two rather heteroge-
subfamilies and nine tribes and the Pucciniaceae neous groups. However, family-level treatments of
into five subfamilies and nine tribes. Most classifica- rust fungi have not been subjected to critical modern
tions emphasize teliospore morphology. Hiratsuka phylogenetic evaluation (Swann et al. 2001).

KEY TO FAMILIES
A key to the 13 families is as follows.

1. Telia eriampents o.oo ois ds a nist saeco ae nice ces a ba ten genet ee eS ee 3
1, Teliatembedded in the host tissue aya. se oa © oeosgeauet mre oe 3
2; Teliospores' Sessiles.. pci bm. 215 4 aoe Uk es a.encase ec 5
2, Teliospores pedicellate. .ac.0es ie cae mace ed ec or ee eetee 9
3. Teliaccomposed ofa crust of teliospores usualliyal (celldeepiey meena
eae tai en 4
3. Telia composed of a crust of teliospores 2 or more cells deep. .............. Phakopsoraceae
4. Aecia CacoimndgAypess css) o tar ree en a ne oe a Melampsoraceae
4. Aecia. Peridermium-typaor Milesia-tye ise oc eee Pucciniastraceae
»: Uredimospores:catenulate,spermogonia Group lyase eee
een te Coleosporiaceae
5. Urediniospores usually pedicellate, spermogonia not Group lene
ane ee 6
6." Teliospores catenulatet” fea Sate spore aire eet Oh, es Go re oa ice ee ne ee ee 7
6...Teliospores not catenulate:j.0pf wc. sos se aso atch eee ee om en see 8
7. leliospores 1-celled; spermogonia Grovpitl ay ae me ae eee Cronartiaceae
7. Teliospores 1- or 2-celled with intercalary cells, spermogonia Group V......... Pucciniosiraceae

8. Basidia semi-internal, aecia Caeoma-type, spermogonia Group III. .......... Mikronegeriaceae

8. Basidia external, teliospores laterally free, spermogonia Group VI... .......... Chaconiaceae

30
 DO PSPermosonlaGrotiprv sewage wane ERD. cs GEIR E NR A IS 2 Pucciniaceae
2 SPETIUNOS Oa OG Gai OU Oy aurae mae mre aeaeRO RC Der oe eco PO ee aa (at6 GE Pn we sswy 10
LOMSpermoe onial Groupny lala weiner Meh MMERMEROMORD ce Ri ie Sar ae A wens, oF cee aloni ae 11
LO] Spermoconia: GLounp, Vi eval ee aoe aa ot eet Nn ware a Nicene bs Phragmidiaceae
TIP Velfosporessiinicell ilar Mayne verte ee I eal sah SO oe Bet suey. a Pileolariaceae
Ll! eliospores: 2, onmore cells various)\natran edi.) martes sees ene te thee aettnaeyl.. INES, BUEN PE 12
12. Teliospores mostly 2 (occasionally more) cells divided by transverse septa. ...... Uropyxidaceae
12. Teliospores divided into 2 or more cells by vertical or muriform septation. ....... Raveneliaceae

DESCRIPTIONS OF FAMILIES Caeoma-type; urediniospores catenulate and mostly

verrucose; germ pores mostly obscure and scattered.
PUCCINIASTRACEAE Gaum. ex Leppik Telia erumpent, waxy or gelatinous, hard when dry,
Ann. Bot. Fenn. 9:135-148. 1972. pulvinate or columnar; teliospores unicellular,
sessile, catenulate, pseudocatenulate, or in a single
DESCRIPTION: layer, walls thin, germ pores not differentiated; ger-
Spermogonia Group I (type 1, 2, or 3). Aecia minating without dormancy, either by septation of
Peridermium-type, with cylindrical peridium (two the spore contents internally (internal basidium)
species of Naohidemyces have Milesia-type aecia); and the production of sterigmata (Coleosporium), or
aeciospores catenulate (except in Naohidemyces) and each teliospore cell germinates to produce a
mostly verrucose. Uredinia with cellular peridium, basidium externally (Chrysomyxa). Most species are
opening by a pore, often with differentiated ostiolar heteroecious with the aecial state on needles, buds,
cells, Milesia-type; urediniospores borne singly with or cones of conifers.
or without definite pedicels, mostly echinulate, germ
TYPE: Coleosporium Lev.
pores obscure, scattered or bizonate. Telia either
subepidermal or intradermal, not erumpent, com- GENERA: Ceropsora, Chrysomyxa, Coleosporium
posed of laterally adherent teliospores one spore
REMARKS:
deep, spores multicellular by vertical septa or uni-
cellular, sessile, germ pore obscure, mostly 1, perhaps
Ceropsora may logically be classified as a part of
Chrysomyxa or together with other species with
not always differentiated; basidia external. Hetero-
well-developed telial basal cells in Stilbechrysomyxa
ecious and mostly macrocyclic, with aecia on conifers.
(not recognized in this manual). Aecial states are all
TYPE: Pucciniastrum G.H. Otth on conifers, especially Picea (Chrysomyxa, Ceropsora)
and Pinus (Coleosporium).
GENERA: Calyptospora, Hyalopsora, Melampsorella,
Melampsoridium, Milesina, Naohidemyces,
Pucciniastrum, Thekopsora, Uredinopsis
CRONARTIACEAE Dietel, in Engler and Prantl
REMARKS: Nat. Pflanzenfam. 1(1) (Suppl.):546-553. 1900.
Aecial states of all known species are on conifers,
predominantly on Abies, but also on Picea and Tsuga. DESCRIPTION:
Spermogonia Group II (type 9). Aecia
Peridermium-type, the peridium large and blister-
COLEOSPORIACEAE Dietel, in Engler and Prantl like, strongly developed, of one or more layers of
Nat. Pflanzenfam. 1(1) (Suppl.):546-553. 1900. cells, rupturing widely; aeciospores catenulate,
coarsely verrucose with rods or platelets, often with
DESCRIPTION: annulate sides. Uredinia with peridium and differ-
Spermogonia Group I (type 2). Aecia entiated ostiolar cells, Milesia-type, sometimes also
Peridermium-type, the peridium mostly strongly with intrasoral paraphyses; urediniospores borne
developed; aeciospores catenulate and verrucose. singly, echinulate, germ pores bizonate. Telia
Uredinia with rudimentary peridium or none, columnar, firm; teliospores unicellular, embedded in

31
a common matrix, germ pores 1 to 3; germinating peridium, Uredo-type; urediniospores borne singly,
without dormancy; basidium external. echinulate, germ pores scattered or bizonate. Telia
subepidermal or rarely subcuticular, not erumpent,
TYPE: Cronartium Fr.
consisting of laterally adherent teliospores in crusts
GENERA: Cronartium, Endocronartium (endocyclic) one spore deep or some species also with subjacent
sporelike (sterile?) cells; teliospores unicellular,
REMARKS:
sessile, pigmented, germ pore 1; basidium external.
Species (except endocyclic Endocronartium) are
Heteroecious or autoecious; mostly macrocyclic.
heteroecious and macrocyclic with aecial states
exclusively on stems and cones of Pinus. TYPE: Melampsora Castagne
GENERA: Melampsora
MIKRONEGERIACEAE Cummins & Y. Hirats. REMARKS:
Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (13). 1983.
The genus Melampsora is large and seems to be
monophyletic. Many genera were once included in
DESCRIPTION:
Spermogonia Group III (type 12). Aecia this genus because of sessile teliospores, but we do
Petersonia-type, without precise peridium but with not think they are close phylogenetically.
hyphoid cellular layers over spore mass; aeciospores
catenulate, without intercalary cells, verrucose.
Uredinia without peridium or paraphyses, Uredo- PHAKOPSORACEAE
type; urediniospores borne singly, echinulate, germ (Arthur) Cummins & Y. Hirats.
pores not seen, perhaps not differentiated. Telia Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (13). 1983.
waxy in appearance, without peridium or
paraphyses; teliospores sessile, unicellular, laterally Phakopsoreae Arthur, Man. Rusts U.S. Can. p. 1-
free, thin-walled, germinating without dormancy 438 (1). 1934.
by continuing apical growth. Entire protoplast is
used in the process of development of septa, DESCRIPTION:
sterigmata, and basidiospores; the basal structure Spermogonia Group VI (type 5 or 7). Aecia
then collapses. Semi-internal basidium. Aecidium-, Caeoma-, or Uredo-type, thus with or
TYPE: Mikronegeria Dietel without peridium; aeciospores catenulate or borne
singly, verrucose or echinulate. Uredinia Malupa-
GENERA: Mikronegeria, Blastospora, Chrysocelis type, mostly with basally united, peripheral,
REMARKS: incurved, dorsally thick-walled paraphyses;
All known species are heteroecious, with the urediniospores borne singly except in Arthuria,
aecial state on conifers and the telial state on where they are catenulate, echinulate; germ pores
Fagaceae (Mikronegeria) or a variety of host groups obscure, mostly scattered. Telia erumpent or
(Blastospora). Some unconnected rusts, such as embedded in host tissue, 2 to several spores deep;
Caeoma peltatum C.G. Shaw III & C.G. Shaw, obvi- teliospores unicellular, sessile, catenulate or irregu-
ously belong to this family. Chrysocelis was classified larly arranged, germ pore 1 in each cell or perhaps
in the family Chaconiaceae (Ono and Hennen 1983; not differentiated in some; basidium external.
Cummins and Hiratsuka 1983), but the deep-seated Heteroecious; not host restricted.
Group III (type 12) spermogonia, Petersonia-type TYPE: Phakopsora Dietel
aecia, and hyaline teliospores producing basidia by
apical elongation indicate possible placement here. GENERA: Arthuria, Batistopsora, Cerotelium,
Crossopsora, Dasturella, Kweilingia,
Nothoravenelia, Phakopsora, Phragmidiella,
MELAMPSORACEAE Dietel, i Engler and Prantl Pucciniostele, Scalarispora, Uredopeltis,
Nat. Pflanzenfam. 1(1):26-81.1897. Monosporidium (endocyclic)

DESCRIPTION: REMARKS:
Spermogonia Group I (type 2 or 3). Aecia Caeoma- Kweilingia is not considered a rust by some, but
type, with rudimentary or no peridium; aeciospores an auricularioid fungus similar to some genera
catenulate, verrucose. Uredinia with abundant capi- once placed in the Auriculariales sensu lato
tate paraphyses and sometimes also rudimentary (Thirumalachar and Narasimhan 1951).

ne
 CHACONIACEAE Cummins & Y. Hirats. Porotenus, Prospodium, Sorataea,
Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (14). 1983. Tranzschelia, Uropyxis
REMARKS:
DESCRIPTION:
Spermogonia Group VI (type 5 or 7). Aecia Uredo- This group of genera was once classified in the
Pucciniaceae because of the morphology of the telio-
or Aecidium-type, thus with or without peridium;
aeciospores borne singly or catenulately, mostly spores, but we do not think they are related to
echinulate, germ pores various. Uredinia Uredo-
genera of that family.
type, with or without paraphyses; urediniospores
borne singly, echinulate for most, germ pores vari- PILEOLARIACEAE
ous. Telia erumpent; teliospores unicellular, later- (Arthur) Cummins & Y. Hirats.
ally free, sessile or pedicellate, thin-walled, with or
Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (14). 1983.
without a poorly defined germ pore; germinating
without dormancy; basidium external or internal by Pileolariaceae Arthur, Rés. Sci. Congr. Int. Bot.
formation of septa within the spore. Autoecious or Vienne. p. 331-348 (340). 1906.
heteroecious; hosts various.

TYPE: Chaconia Juel DESCRIPTION:

Spermogonia Group VI (type 5 or 7). Aecia Uredo-
GENERA: Achrotelium, Aplopsora, Botryorhiza, type; aeciospores borne singly, usually reticulate,
Ceraceopsora, Chaconia, Goplana, ridged, verrucose, or spirally marked, germ pores
Hemileia, Maravalia, Ochropsora, Olivea zonate. Uredinia as the aecia, but not accompanied
by spermogonia. Telia erumpent; teliospores unicel-
REMARKS:
lular, with one or a few spores on each pedicel,
Ono and Hennen (1983) did not consider Achrote-
which may also have sterile cystlike cells laterally,
lium, Botryorhiza, Chaconia, and Maravalia as belonging
germ pore 1 per spore, germination by external
to this family because of the pedicellate teliospores.
basidium. Autoecious; hosts Anacardiaceae and
Fabaceae.
UROPYXIDACEAE TYPE: Pileolaria Castagne
(Arthur) Cummins & Y. Hirats.
Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (14). 1983. GENERA: Atelocauda, Pileolaria, Uromycladium,
Endoraecium (endocyclic)
Uropyxioideae Arthur, Rés. Sci. Congr. Int. Bot. REMARKS:
Vienne. p. 331-348 (341). 1906. All spore states of the genera of this family have
unique spore surface markings (see genus descrip-
DESCRIPTION:
tions).
Spermogonia Group VI (type 5). Aecia Aecidium-
or Caeoma-type, or more commonly Uredo-type, thus
with or without peridium, or with paraphyses; RAVENELIACEAE (Arthur) Leppik
aeciospores catenulate or borne singly, verrucose or Ann. Bot. Fenn. 9:135-148. 1972.
echinulate. Uredinia Calidion-, Malupa-, or Uredo-
type, with or without paraphyses, or Wardia-type Raveneliae Arthur, Rés. Sci. Congr. Int. Bot. Vienne.
(suprastomatal); urediniospores borne singly, p- 331-348 (341). 1906.
mostly echinulate, germ pores mostly scattered. Sphaerophragmiaceae Cummins & Y. Hirats., Illus.
Telia with or without paraphyses; teliospores 2- or Genera Rust Fungi. Rev. ed. p. 1-152 (13). 1983.
more celled by vertical septa, often with bilaminate
wall, the outer layer hygroscopic, germ pore 1 or DESCRIPTION:
more per cell; teliospores pedicellate, the pedicels Spermogonia Group IV, VI (type 5, 7, or 11). Aecia
often hygroscopic; basidium external. Autoecious Aecidium- or Caeoma-type, with or without
or heteroecious; hosts various. peridium, or typically Uredo-type, with or without
paraphyses; aeciospores borne singly on pedicels,
TYPE: Uropyxis J. Schrot.
echinulate in most, or catenulate and verrucose.
GENERA: Dasyspora, Desmella, Didymopsorella, Uredinia Malupa-, Calidion-, or Uredo-type; uredinio-
Dipyxis, Edythea, Kimuromyces, spores borne singly, mostly echinulate, germ pores
Macruropyxis, Newinia, Phragmopyxis, various, usually obvious. Telia erumpent, with or

31S,
without paraphyses; teliospores pedicellate, verti- without peridium and with aeciospores borne sin-
cally septate or vertically or radially arranged atop gly, or rarely without peridium but with catenulate
the pedicel, 2 or more spores on a pedicel, or 2 to spores. Uredinia Uredo-type, with or without
many cells of a spore on a pedicel, often subtended paraphyses, or Uredostilbe-type with palisadelike
by hygroscopic cysts or with pedicels having apical peridium; urediniospores borne singly, mostly echi-
cells; germ pores 1 or 2 in each cell or spore; nulate, germ pores various. Telia with or without
basidium external. Autoecious; hosts mostly paraphyses, but rarely with palisadelike peridium,
Fabaceae or Rosaceae, but also Euphorbiaceae, or telia may be separated into locules by stromatoid
Apiaceae, Tiliaceae, and Dioscoreaceae. paraphyses; teliospores borne singly, mostly pedi-
cellate, with 1 or 2 (rarely more) cells, septa trans-
TYPE: Ravenelia Berk.
verse or oblique, germ pore mostly 1 in each cell;
GENERA: Allotelium, Anthomyces, Anthomycetella, germination mostly by external basidium, rarely by
Apra, Cumminsina, Cystomyces, Diabole, internal basidium. Heteroecious or autoecious;
Dicheirinia, Diorchidiella, Diorchidium, hosts various.
Hapalophragmium, Kernkampella, Lipocystis,
TYPE: Puccinia Pers.
Nyssopsora, Ravenelia, Sphenospora,
Sphaerophragmium, Spumula, Triphrag- GENERA: Chirysella, Chrysocyclus, Chrysopsora,
miopsis, Triphragmium, Ypsilospora Cleptomyces, Corbulopsora, Cumminsiella,
Gymnosporangium, Kernella, Miyagia,
REMARKS:
Polioma, Puccinia, Stereostratum, Uromyces,
It may be justified to include Ypsilospora in the
Zaghouania, Endophyllum (endocyclic)
Chaconiaceae.
REMARKS:
This family includes the two largest rust genera,
PHRAGMIDIACEAE Corda
Puccinia and Uromyces; they are separated only by
Abfildungen Pilze Schwamme 1:1-32 (6). 1837.
the number of cells in the teliospores (2 for Puccinia
and 1 for Uromyces), which is sometimes not a stable
DESCRIPTION:
character.
Spermogonia Group IV (type 6,8, 10, or 11). Aecia
Caeoma- or Petersonia-type, with catenulate aecio-
spores, or Uredo-type, with spores borne singly on
PUCCINIOSIRACEAE
pedicels; aeciospores verrucose or echinulate. Ure-
(Dietel) Cummins & Y. Hirats.
dinia Calidion- or Uredo-type, mostly with thin-walled,
Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (15). 1983.
incurved, peripheral paraphyses; urediniospores borne
singly, mostly echinulate, germ pores scattered. Telia
Pucciniosireae Dietel, in Engler and Prantl, Nat.
erumpent, with or without paraphyses; teliospores
Pflanzenfam. 2 (6):24-98 (93). 1928.
with 1 to several cells by transverse septa, pedicellate,
germ pores | or more in each cell; basidium external.
DESCRIPTION:
TYPE: Phragmidium Link Spermogonia usually not produced, but known
species have Group V (type 4). Aecia and uredinia not
GENERA: Frommeélla, Gerwasia, Gymnoconia,
produced, all genera microcyclic. Telia in gross ap-
Hamaspora, Joerstadia, Kuehneola, Newinia,
pearance mostly resemble the aecial states of macro-
Phragmidium, Trachyspora, Xenodochus cyclic genera, with or without peridium; teliospores
REMARKS: sessile, of 1 or 2 cells, often catenulate with interca-
All known species are autoecious, mostly or lary cells, or extruded as columns or filaments, the
perhaps only on Rosaceae. columns often firm; basidia external or internal.

TYPE: Pucciniosira Lagerh.

PUCCINIACEAE Chevall. GENERA: Alveolaria, Baeodromus, Ceratocoma,
Flore Gén. Env. Paris 1:1-676. 1826. Chardoniella, Cionothrix, Didymopsora,
Dietelia, Pucciniosira, Trichopsora
DESCRIPTION:
Spermogonia Group V (type 4). Aecia Aecidium- REMARKS:
or Caeoma-type, with or without peridium, and cate- The species are all autoecious and presumably
nulate, mostly verrucose aeciospores, or Uredo-type, endocyclic derivatives of macrocyclic genera, but

34
the relationships are often obscure. This is a very Chevallier, RF. 1826. Flore générales des environs de Paris. Chez
Ferra jeune. Vol. 1. 676 p.
heterogeneous family. Buritica and Hennen (1980)
Corda, A.K.J. 1837. Icones fungorum hucusque cognitorum.
included the clearly endocyclic genus Endophyllum
Abfildungen der Pilze und Schwamme. 1:1-32.
in this family but not other endocyclic genera. Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
However, Buritica (1991) recognized the family fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
Endophyllaceae Dietel and included Endophyllum Cummins, G.B.; Hiratsuka, Y. 1984. Families of Uredinales. Rep.
and all other endocyclic genera in it. We think that Tottori Mycol. Inst. 22:191—208.
when possible, endocyclic genera should be classified Cunningham, G.H. 1931. The rust fungi of New Zealand. John
in the same family as suspected parental genera. McIndoe, Dunedin. 261 p.
Dietel, P. 1897. Uredinales. In A. Engler and K. Prantl. Nat.
Pflanzenfam. 1(1):24-81.
GENERA OF UNCERTAIN AFFINITY Dietel, P. 1900. Uredinales. In A. Engler and K. Prantl. Nat.
Pflanzenfam. 1(1) (Suppl.):546-553.
Hiratsukamyces Dietel, P. 1928. Hemibasidii (Ustilaginales und Uredinales). In A.
Engler and K. Prantl. Nat. Pflanzenfam. 2(6):24-98.
Masseeélla Gatimann, E. 1949. Die Pilze. Grundztige ihrer Entwicklungs-
geschichte und Morphologie. Birkhauser, Basel. 382 p.
Skierka
Hiratsuka, N. 1955. [Uredinological studies.] Kasai Publ. Co.,
Tokyo. 382 p. (In Japanese.)
Hiratsuka, Y.; Cummins, G.B. 1963. Morphology of the spermo-
LITERATURE CITED gonia of the rust fungi. Mycologia 55:487-507.
Arthur, J.C. 1906. Eine auf die Struktur und Entwicklungs- Hiratsuka, Y.; Hiratsuka, N. 1980. Morphology of spermogonia
geschichte begrtindete Klassifikation der Uredineen. Reés. Sci. and taxonomy of rust fungi. Rep. Tottori Mycol. Inst. 18:257—
Congr. Int. Vienne. p. 331-348. 268.
Arthur, J.C. 1934. Manual of the rusts in United States and Leppik, E.E. 1972. Evolutionary specialization of rust fungi
Canada. Purdue Res. Found., Lafayette, IN. Reprinted with a (Uredinales) on the Leguminosae. Ann. Bot. Fenn. 9:135-148.
supplement by G.B. Cummins (1962). Hafner Publ. Co., New Ono, Y.; Hennen, J.F. 1983. Taxonomy of Chaconiaceous genera
York. 438 p.
(Uredinales). Trans. Mycol. Soc. Jpn. 24:369-402.
Azbukina, Z.M. 1974. [Rust fungi of the Soviet Far East.] Nauka, Swann, E.C.; Frieders, E.M.; McLaughlin, D.J. 2001. 2. Uredinio-
Moscow. 527 p. (In Russian.)
mycetes. Pages 37-54 in D.J. McLaughlin and P. Lemke, eds.
Bessey, E.A. 1950. Morphology and taxonomy of fungi. Blakiston The Mycota. Vol. VIIB. Systematics and evolution. Springer-
Co., Philadelphia. 791 p. Verlag, Berlin.
Buritica, P. 1991. Familias del orden Uredinales con ciclo de vida Thirumalachar, M.J.; Narashimhan, M.J. 1951. Critical notes on
completamente reducio. Rev. Acad. Colomb. Cienc. Exactas some plant rusts. III. Sydowia 5:476-483.
Fis. Nat. 18:131-148. Wilson, M.; Henderson, D.M. 1966. British rust fungi. Cambridge
Buritica, P.; Hennen, J.-F. 1980. Pucciniosireae (Uredinales, Univ. Press, London. 384 p.
Pucciniaceae). Flora Neotrop. Monogr. 24:1—50.

35
 ANAMORPH GENERA OF RUST FUNGI
cog

[e this manual we define anamorph by morphol- Although this practice is allowed under Article 59.5
ogy only, without designating a life cycle stage of the International code of botanical nomenclature
(aecia or uredinia) for the morph, or discussing taxo- (Greuter et al. 2000), it adds confusion and unneces-
nomical or phylogenetic implications. For this sary names, and is specifically discouraged under
reason, presence or type of spermogonia are also Recommendation 59A-3. Although anamorphs are
disregarded, although the original descriptions of sometimes more characteristic and even more im-
certain genera might include them. portant in nature, especially in the tropics, the Code
We also discourage the use of terminology such clearly states that the name of the holomorph is that
as uredinoid aecia, aecioid uredia, or uraecia and the based on the teleomorph. The holomorph name
abbreviations such as II!, I", and II!, which are often includes the whole fungus in all its morphs and
used in textbooks and other publications. In species stages (teleomorph, anamorph, and even hyphae).
descriptions, instead of giving separate binomials A basic principle of our nomenclature (Principle IV
for anamorph states when the teleomorph is known, of the Code) is that one organism should have one
we suggest mentioning the morphological genus correct and legitimate name, and this principle
type (morph) of an anamorph together with life should be adhered to as much as possible in rust
cycle state, for example, aecia Aecidium-type, fungi.
uredinia Aecidium-type, or aecia Uredo-type, etc. In Based on the above considerations and several
recent years, some authors have assigned separate recent published studies (Buritica and Hennen 1994;
anamorph names, along with holomorph names, in Kenney 1970; Laundon 1967; Ono et al. 1992; Sato
new descriptions of rust fungi (Buritica 1998, 1999; and Sato 1985), we recognize 13 anamorphic genera
Buritica and Hennen 1994; Ono et al. 1992). as follows.

KEY TO ANAMORPH GENERA

1.. Spores pedicellate or'sesstle. | cote a crise, ths mha)osalee Oe Re elePare AS ea 2

1;. Spores catenulate ss oze.) sola ee ea aa a eta gen eae Vi
2. Sorus with no bounding structure (may have intermixed paraphyses)..............-.-. 3
2; OOrus With DOUNndING SEMUICHUITE: ce eae a oy cst ea eee ceecules ia) ead ee 4
3. worus subepidermal Or subcuticular ey ca. st ee Uredo
3. borus suprastomatal”s. [5 Sysea ase te lores so ae va Wardia
4. Sorus with peripheral paraphiyses< i. sue eale =<. Mireur ach ce ee ee
4) OOTUS With peridivm 5. civ gels, Siew. ee ee eg tS ea cmc real aa a 6
Db. porus with peripheral paraphyses united Dasallyen scm edicts eter cle el ane en oe ee Malupa
2, sors with peripheral paraphyses not-united! basally fe wee eve ee ne Calidion
6, Sorus covered byidome-shapedsnericnurrny ag foc ae eee eee ele ae ee Milesia
6. Sorus surrounded by laterally fused palisadelike compact peridium. ............. Uredostilbe
7s Interealary cells absent «3: cr.) hy aile ee ccs oo ade oem ered aan eant ee tae a 8
7, Intercalary tells present, 25. sun: sano cee ree ae epee eee neater ad 9
8. Spores intermixed with sterile elaterlike hyphae « 2. 6s14004844%+4500%05 Elateraecium

36
 37 No stemleelateriikehypnaesms saree. Petersonia (Caeoma IV and V of Sato and Sato 1985)

9.,Peridiumy pataphyses absent: a5" yan2/2). . .. . Caeoma (Caeoma I, I, Il of Sato and Sato 1985)
9.7 Peridium or paraphyses present \i).-02 28). ss .
10Peripheral paraphyses present.¢4 008.
5 ona!
10, BRenaiini presente | acs. e oRkeeneee
11. Sorus cup shaped, peridial cells usually short. .
11, Semis elongated and not cup shaped, peridialtcells usuiallyalonges. tat 4 2. 2 et 12
12. Sorus with single- to multilayered peridium. . . Fh, esate os Spe a omROR ag ae er, Smee Peridermium

12. Sorus with well-developed peridium, tending to shred longitudinally at maturity. ...... Roestelia

DESCRIPTIONS OF ANAMORPH GENERA simply as verrucose or echinulate, we consider

Aeciure as a synonym of Caeoma. Occurs as uredinia
AECIDIUM Pers. ex Pers. in Melampsora, Gymnoconia, Chrysocelis, Polioma,
Synop. Method. Fungorum. p. 1-708 (204). 1801. Arthuria, and others.

DESCRIPTION:
Cup-shaped sorus with well-developed CALIDION H. & P. Syd.
peridium. Spores catenulate with intercalary cells. Ann. Mycol. 16:240—248 (242-243). 1919.
Spore surface typically verrucose (Fig. 18C).
DESCRIPTION:
TYPE: Aecidium berberidis Pers. ex Pers. (lectotype, Sorus surrounded by thick-walled, strongly
Clemens and Shear 1931) incurved peripheral paraphyses. Spores produced
=Puccinia graminis Pers. ex Pers. singly on pedicels, or several spores develop
On: Berberis vulgaris L. (Berberidaceae) sympodially from a common basal cell (Fig. 181).
Type locality: Europe
TYPE: Calidion lindsaeae (Henn.) Syd.
REMARKS: =Uredo lindsaeae Henn.
Most species of Puccinia, Uromyces, and On: Lindsaea sp. (Polypodiaceae)
Tranzschelia have this type of anamorph generally as Type locality: Brazil
the aecial state, but also often as uredinia.
REMARKS:
This genus was established to accommodate the
CAEOMA Link uredinial state of a rust ona fern (Lindsaea) because it
differs from the uredinial states of other fern rusts.
Mag. Ges. Naturf. Freunde 3:5. 1809.
Occurs as uredinia of Prospodium, Olivea, Crossop-
DESCRIPTION: sora, and Phragmidium. Macabuna Buritica & Hennen
Sorus without bounding structure. Spores (Buritica and Hennen 1994) was established specifi-
catenulate, with intercalary cells (Fig. 18B). cally for spores produced sympodially instead of
the usual solitary terminal spores, but we will treat it
TYPE: Caeoma saxifragarum (DC.) Link (lectotype, here as a synonym of Calidion.
Clements and Shear 1931)
=Melampsora vernalis G. Winter
On: Saxifraga aizoides L. (Saxifragaceae) ELATERAECIUM Thirum., F. Kern, & B.V. Patil
Type locality: Europe Mycologia 58:391-396 (394-395). 1966.

REMARKS: DESCRIPTION:
Aeciure Buritica & J.F. Hennen (Buritica and Sori are like Petersonia, with catenulate spores
Hennen 1994) has echinulate rather than generally without intercalary cells, but with unique elaterlike
verrucose spore surface markings, but because spore hyphae that expand out of the sorus, carrying spores
surface markings are not clearly distinguished with them (Fig. 18Q).

37
 A Petersonia Aecidium D Peridermium

E Roestelia F
Lecythea G Uredo H Uredo

Calidion J Malupa K Milesia Ne Milesia

M Uredostilbe Wardia O Elateraecium

FIGURE 18. Anamorph genera. A. Petersonia. B. Caeoma. C. Aecidium. D. P.eridermium. E. Roestelia. F. Lecythea.
G, H. Uredo. 1. Calidion. J. Malupa. K , L. Milesia. M. Uredostilbe. N. Wardia. O. Elateraecium.
TYPE: Elateraecium salicicola Thirum., F. Kern & Type locality: UK
B.V. Patil
REMARKS:
On: Salacia prinoides DC. (Hippocrateaceae)
Occurs as uredinial states of Milesina, Hyalopsora,
Type locality: India
Uredinopsis, Melampsoridium, Cronartium,
REMARKS: Melampsorella, Pucciniastrum, and Thekopsora and
Known to occur only in the genus Hiratsukamyces. aecial and uredinial states of Naohidemyces. We con-
All known species have Type 12 spermogonia, as sider Peridipes Buritica & J.K. Hennen (Buritica and
with Petersonia. Hennen 1994), Peridiopsora K.V. Prasad et al. (Prasad
et al. 1993), and Uraecium Arthur as synonyms of
Milesia.
LECYTHEA Lév.
Ann. Sci. Nat. Ser. 38:373. 1847.
PERIDERMIUM (Link) J.C. Schmidt & Kunze
DESCRIPTION: Dtschl. Schwamme. p. 141. 1817.
Sorus with peripheral paraphyses and with
catenulate spores with intercalary cells (Fig. 18F). DESCRIPTION:
Sorus tublar or blisterlike with well-developed
TYPE: Uredo miniata Pers.
peridium. Spores catenulate with intercalary cells,
=Phragmidium mucronatum (Pers. ex Pers.)
generally verrucose, but many variations (Fig. 18D).
Schltdl.
On: Rosa eglanteria L. (Rosaceae) TYPE: Peridermium elatinum (Alb. & Schwein.) J.C.
Type locality: Europe Schmidt & Kunze (conserved type,
Hiratsuka 1974)
REMARKS:
=Melampsorella caryophyllacearum J. Schrot.
Corresponds to Caeoma III of Sato and Sato (1985).
On: Abies alba Mill. (Pinaceae)
Occurs as aecia of Phragmidium.
Type locality:: Europe
REMARKS:
MALUPA Y. Ono, Buritica & J.F. Hennen Occurs mainly as aecial states of Calyptospora,
Mycol. Res. 96:825-850 (828). 1992. Chrysomyxa, Coleosporium, Cronartium, Hyalopsora,
Melampsorella, Milesina, Pucciniastrum, and
DESCRIPTION: Thekopsora.
Sorus with basally united paraphyses, spores
produced singly, mostly sessile or pedicellate (Fig.
18)J). PETERSONIA Cummins & Y. Hirats.
TYPE: Malupa meibomiae (Arthur) Y. Ono, Buritica gen. anam. nov.

& J.F. Hennen

DESCRIPTION:
=Physopella meibomiae Arthur
Sori subepidermalibus, erumpentis; peridio et
On: Fabaceae
paraphysibus nullis; sporis catenatis, cellulis
Type locality: Brazil
intercalares nullis.
REMARKS: Sorus subepidermal, deep-seated. No peridium
Known to occur as uredinia of Phakopsora and or paraphyses. Spores catenulate without interca-
Olivea. lary cells (Fig. 18A).
TYPE: Petersonia sanctae-crucis (Espinosa)
MILESIA F.B. White Cummins & Y. Hirats.
Scott. Nat. 4:160-163 (162). 1878. =Caeoma sanctae-crucis Espinosa
=Mikronegeria fagi Dietel & Neger
DESCRIPTION: On: Araucaria araucana (Molina) K. Koch
Sorus with dome-shaped cellular peridium, with (Araucariaceae)
various modifications of ostiolar cells (Figs. 18K, L). Type locality: Chile

TYPE: Milesia polypodti F.B. White REMARKS:

=Miulesina dieteliana (P. & H. Syd.) Magnus This is Caeoma IV and V of Sato and Sato 1985.
On: Polypodium vulgare L. (Polypodiaceae) Represented by aecia of Mikronegeria, Blastospora,

Bo
and some unconnected aecia such as Caeoma TYPE: Uredostilbe pistila Buritica & J.F. Hennen
peltatum C.G. Shaw II & C.G. Shaw (Shaw and Shaw On: Annona nolosericea Safford (Annonaceae)
1976). Etymology: after noted uredinologist Dr. Type locality: Honduras
Roger S. Peterson.
REMARKS:
Occurs as uredinia of Batistopsora, Miyagia, and
ROESTELIA Rebent. Corbulopsora.
Prodr. Fl. Neomarch. p. 350. 1804.

DESCRIPTION: WARDIA J.F. Hennen & M.M. Hennen

Sorus with well-developed peridium that tends ined.
to shred longitudinally at maturity. Peridial cells
usually long and narrow (see figures under DESCRIPTION:
Gymnosporangium). Spores catenulate with interca- Sorus suprastomatal; spores borne singly on short
lary cells (Fig. 188). pedicels, typically strongly asymmetrical (Fig. 18N).

TYPE: Roestelia cancellatum Jacq.) Rebent. TYPE: Wardia vastatrix J.F Hennen & M.M. Hennen
On: Pyrus communis L. (Rosaceae) REMARKS:
Type locality: Europe Occurs as uredinia of Hemileia, Desmella, and
REMARKS: Edythea.
Typically applied to aecial states of Gymno-
sporangium, but aecia of some Gymnosporangium
species can be considered as Aecidium, and some EXCLUDED GENERA
Puccinia species have typically Roestelia-type aecia. Physopella has been used as an anamorph genus
by some authors (Ono et al. 1992), but the genus is
recognized as a holomorph generic name in this
UREDO Pers. ex Pers.
manual (see Phakopsora). The genus Uraecium was
Synop. Method. Fungorum, 1:1—708 (218). 1801.
established by Arthur (1933) to accommodate aecia
DESCRIPTION: on Tsuga (now recognized as the aecial state of Nao-
Sorus without definite bounding structure, but hidemyces) with uredinium-like morphology, but it
may have intermixed paraphyses. Spores, pedicel- is not a meaningful genus in our ontogenic system
late or nearly so, typically echinulate (Figs. 18G, H). and is placed as a synonym of Milesia. Intrapes has a
unique type of spore formation, but it is excluded
TYPE: Uredo euphorbiae-helioscopiae Pers. (lectotype) here because the fungus may not be a rust.
=Melampsora euphorbiae (C. Schub.)
Castagne
On: Euphorbia helioscopia L. (Euphorbiaceae) LITERATURE CITED
Type locality: Europe Arthur, J.C. 1933. New genera and species of Uredinales. Bull.
Torrey Bot. Club 60:475-476.
REMARKS:
Buritica, P. 1998. La familia Phakopsoraceae en el neotropico. II.
This definition is much narrower than is tradi- Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22:325-334.
tional for Uredo, but it still applies to many genera
Buritica, P. 1999. Familia Phakopsoraceae (Uredinales)
and species. It occurs mainly as uredinia and some generalidades y afinidades. Rev. Fac. Nac. Agron. Medellin
aecia of Melampsora, Puccinia, and Uromyces. We con- 52:467-505.
sider Uredendo Buritica & J.-F. Hennen (Buritica and Buritica, P.; Hennen, J.F. 1994. Familia Phakopsoraceae
Hennen 1994) a synonym of Uredo. (Uredinales). 1. Géneros anamorficos y teliomérficos. Rev.
Acad. Colomb. Cienc. Exactas Fis. Nat. 19:47-62.
Clemens, F.E.; Shear, C.L. 1931. The genera of fungi. H.W. Wilson,
UREDOSTILBE Buritica & J.F. Hennen New York. 496 p.
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. Greuter, W.; McNeill, J.; Barrie, RFR.; Burdet, H.M.; Demoulin, V.;
Filgueiras, T.S.; Nicholson, D.H.; Silva, P.C.; Skog, J.E.;
19:47-62 (49). 1994.
Trehane, P.; Turland, N.J.; Hawksworth, D.L. 2000. Interna-
tional code of botanical nomenclature (Saint Louis code).
DESCRIPTION: Koeltz Books, Konigstein, Germany. 474 p.
Sorus surrounded by laterally fused palisadelike Hiratsuka, Y. 1974. Proposal to conserve the generic name
cells (Fig. 18M). Peridermium (Link) Schmidt & Kunze with a conserved type

40
 species Aecidium elatinum Alb. & Schw. (Fungi Imperfecti Prasad, K.V.; Yadav, B.R.D.; Sullia, S.B. 1993. Taxonomic status of
Uredinearum). Taxon 23:428-429. rust on mulberry in India. Curr. Sci. (Bangalore) 65:424—426.

Kenney, M.J. 1970. Comparative morphology of the uredia of the Sato, T.; Sato. S. 1985. Morphology of aecia of the rust fungi. Trans.
rust fungi. Ph.D. thesis, Purdue Univ., West Lafayette, IN. Br. Mycol. Soc. 85:223-238.
76 p. Shaw, C.G., Ill; Shaw, C.G. 1976. Rust on Phyllocladus
trichomanoides—the first record on a member of the
Laundon, G.F. 1967. The taxonomy of imperfect rusts. Trans. Br.
Podocarpaceae. Trans. Br. Mycol. Soc. 67:506-509.
Mycol. Soc. 50:349-353.
Ono, Y.; Buritica, P.; Hennen, J.F. 1992. Delimitation of Phakopsora,
Physopella and Cerotelium and their species on Leguminosae.
Mycol. Res. 96:825-850.

41
 HOLOMORPH GENERA OF RUST FUNGI
“>

KEYS TO HOLOMORPH GENERA, BY FAMILY

PUCCINIASTRACEAE
I) Teliospore wall colorless <5 eer ne 7

1. Teliospore wall yellowish brown toxbrowmy: (72 ey cece 6

= «00. auiiaees te tenet kent a ceca 3

2. Urediniospores hyaline, devoid of pigment

2. Urediniospores with yellow to orange cytoplasmic pigment, wallicolorless =) sa. sn rence +

3. Teliospores in the mesophyll ce ms yeu. nn te erg Uredinopsis

3.. Teliospores within epidermalicells) 73. eee ee cet Milesina

4. Teliospores subepidermal, ostiolar cells of uredinia well developed. .....----- Melampsoridium

.......-....-++---: 5
4. Teliospores intraepidermal, ostiolar cells of uredinia not well developed

5. Uredinial peridium poorly developed, opening irregularly... -.. +--+ +222 eee Hyalopsora

5. Uredinial peridium well-developed, ostiole discrete... 6... ee +e eee Melampsorella

6. Aecial state Milesta-types 2...2 sss ete 0 ee Naohidemyces

6.. Aecial state Peridermium-type. 17 see = ce ee eee ee 7

7. Teliospores subepidermal = 2.2 cscs ce eet oi Pucciniastrum

7. Teliospores intraepidermal 29.5 ers ees a eee ec ee 8

8. Teliospores on stems, no uredinial state... 2. - 22-2 0 te eee ee Calyptospora

8. Teliospores on leaves, tiredinial state present.) an yemencg= nme cite eon nen ceca Thekopsora

COLEOSPORIACEAE
li. Basidia internal .4-..<-a05 oo 2 eee ce roi ae ee Denes Coleosporium

1.. Basidia-external.. {oj.cc 5)seca cere: eee aeeri ees Soc sec ce 2

2. Teliospores single on well-developed basal cells... .-- +--+ +e eee eee tts Ceropsora

2. Teliospores catenulate, with or without well-developed basal cells... ...-..-.---. Chrysomyxa

CRONARTIACEAE
y 4). 4c i2505 ac per wees
Heteromacrocclic: Cerea eae ne Cronartium

Endocyclic, teliospores similar to aeciospores of Cronartium .. 0.66 ee ee ee Endocronartium

MIKRONEGERIACEAE
1. “Teliospores pedicellatets s-5.oe- a.arairscnteatse- => okie es mea Blastospora
aneurin

1; Teliosporés sessile 4... 5 < Sink aie oe ee eh ec Cn ine 2 en wed gerne gre 2

42
 2, Oniavaceae ae acer hy Het enema ore ce Ste Sa ay eal oe & ROT Mikronegeria
27 On Fabaceae eh tatters oe eo hale, Olpes) were ce bee: bore acecge Chrysocelis

MELAMPSORACEAE
Melampsora

PHAKOPSORACEAE
Ie Leliosporesccaten
vilate om uw ieunn twee te mites Meter er Rete tee eon ee an ee Atos 2
i Tehospores noticatenUlate peammaa nr nt eee rae ete arrears aaclee ca he Se ogoe ce wi
Ph NCSIOS
B10) CEPA UO 0)
0)A ISISOF,oe a saad) acon eet oto Sinokceecerc ners lial tillers eco erel ftclra a aoe icra 3
23 helios pores more. thala cpOles COCep wae mre ei meme men encanta MM ter ete Ree 8s 4
o, leliospore wall: piemented, urediniaNeaidien
ty pens ese Ata comes et ee Dasturella
3. Teliospore wall thin and hyaline, uredinia Uredostilbe-type..........-..-++5--. Batistopsora
4*> Veliosporesacherentlaterallys warns er ear en eRe 2 Se sat 2 Ra hse ss 5
4~ eliosperes nouad Nereng laterallyanemme eee errs eee et ie ee so 8
5: Teliospores:ctronclyadheremtjam namiike columns 2teat ce hac gee | ee a le Crossopsora
ST illeliospores looseyea RClen ema ame meek ee Eee care roe erp ent se gee i) as 6
6. Producesitwokindsiot teliosporetocetheiis ayer weer eee eee es Pucciniostele
6. Prodncesioneskindsoftelioshotcener me rwetas cc Aor et i go ee ss 2 9
7, Reliaretumpentytelospotemvrall palesr: wm. jamal onotrag Seung re mean yin bert ay el yt nk Arthuria
7. Telia discoid-pulvinate, spread over host surface, teliospore wall brown ........... Kweilingia
6. Wredinianwithibasall vaunted arapliyces aria sucteace ag mite rdialee eine sectnss, Glas 2: Cerotelium
8 Wredinianwathoutmatap
hy ses. araet ace mak home rene orogens, Beene) Aan oak os Phragmidiella
9, Teliospores sessile withisubtendinecystoid cells Fa) cee ey pees 2 2 Nothoravenelia
DE Teliospores Without subrendmercystoid cells). ey cs eg ens eye pee ye ee cen oe > ASD iNest 10
10. Telia not erumpent, consisting of crusts of laterally adherent spores... ........... Phakopsora
10) Teliastrons lyaeruinpentemeenn eee ee ee ee ee he ee peg bole che eee yor Big oe id
11. Teliospores pointed at both ends, composed of 5-8 cells... -.- 5.2 ++2+++2++2-- Scalarispora
11. Telia consisting of cushions of laterally adherent spores... .-.---
2-2 +--+ +23. Uredopeltis
Endocyclic genus, telia Aecidium-type, spermogonia type7......-..-....-++-+4-. Monosporidium

CHACONIACEAE
dee lchawwanxy Oneciatnouss erect mn Men ane iau te eet a een eens ee Gebel Jes
PT elia net waxyoLcelatinOusme awe ee ee ees es Rok eg sees a fe]
2. Telia waxy, teliospores sessile on sporogenous basal cells. .......-.-.--+-++--. Ceraceopsora
2. Telia gelatinous, teliospores sessile, embedded ina gelatinous matrix ............--. Goplana
LISI
FeUPI ba\RasyOPEDIe oh oywriuel aire i Attlee abe sah polchashed ali Sc rte det ac A ie a i a a a a +
3. Basidia‘external... 02.0% de Aen © ee ee ee eee cee eeegro) 5

4. Basidiospores produced on long sterigmata ....-- eee teres Achrotelium

+--+ ese --.

4. Basidiospores produced without sterigmata, sessile... 2-66-1 ee ee 1ee Ochropsora

5. Teliospores pedicellate. . ..: ie sy eee t-te ee od eee 6

bi WTeliospores sessile Wo. ec we ce mere arr ere 8

6. Telia mostly suprastomatal, uredinia Wardia-type.....- +--+ eee ee es Hemileia

6. Telia not suprastomatal ... . 0 5 <1 6 Bees yee ee 7

7.. On Hippocrateaceae . . . 2.6 es ee eee ee a Botryorhiza

7. Mostly on Fabaceae and Sapotaceae 97 2025 2-502 oe =e ee one Maravalia

8. Teliospores produced on sporogenous basal cells... .. - 1-1 +e eee ee tes 8

8. Teliospores without sporogenous basal cells... 2... ee es Aplopsora

9. Telia with peripheral paraphyses and on well-developed sporogenous basal cells .....--. Olivea
9. Teliospores produced in groups on simple sporogenous basal cells... -. +. +++ +-+-- Chaconia

UROPYXIDACEAE
1. Teliospores pedicellate and:2-celled 72 =) wnt een er 2

1. Teliospores pedicellate and 3- to many celled... .. 2-22 - +s eee eee eae 1)

2. Teliospores 2-celled by transverse, oblique, or vertical septa, emerging from stomates.......-. 3

2. Teliospores 2-celled only by transverse septa, telia subepidermal, erumpent -.

++.
.... +--+: 4

3. Several teliospores produced on each sporogenous cell... . - - 0 0 ee ee ee Desmella

3. Single teliospores produced on each sporogenous cell... 26.1 2-2 ee ee Edythea

4. Telia with intermixed, hygroscopic paraphyses ... ..-

+--+ etree Didymopsorella
+ eee eee.

4. Telia without hygroscopie paraphySeSimans05 <u cence ee ee ose 5

5: Teliospore pedicels often hy roscOpic™ os.) eee ee

ee Uropyxis

5. Teliospore pedicels not hygroscopic. 2 ee se <p eee te 6

6. Teliospore walls smooth. <<, 2 veu ose ee oie ening ceeds ele Abeer ih

6. Teliospore walls ornamented): 7) 2g. ey jee een ee cc ice ne 2)

7. Basidia produced by elongation of apex of telialcells... 2... - 1 eee ee ee eee Sorataea

62 eee ees 8
7. Basidia produced from a discrete area of teliospore or from germ pores... 2...)
8. Leliospore germ pores | per cell 2 2a eyr0 o1g aerry eee stta eetee ean Porotenus

8. Teliospore germ: pores:3 to4 .-1 285 ese tee ee eh te cee Macruropyxis

9. Teliospores echinulate or verrucose) (5.22 = 5 2c ee 10

9. Teliospores with prominent simple or sometimes branched appendages. ......... Kimuromyces

10. Teliospore germ pores 2 in each cell, reniform urediniospores .... 6. 6-71 ee ee Dipyxis

10. \Teliospore germ pores: 1 in eachscell scascte, Bae sop? tye eine te eet neo gtd 11

11. Conspicuous rodlike, mostly branched verrucae on both ends of teliospores ........- Dasyspora

44
ii Veliosporeormamentation nods aby ewan a ae a erareine earcin mera iews ered Pei y ete mean ATs 12
12. Uredinia and telia erumpent or suprastomatal, often with basal peridial cups. ....... Prospodium
2a rediniawith oblongeto capitate, patap
iy Ses tray... pene terksbSik? le valent noite ettons gees? Tranzschelia
13. Teliospores mostly 3-celled, teliospore wall conspicuously bilaminate, echinulate . . . . Phragmopyxis
13. Teliospores 3- to many celled, teliospore wall pigmented and smooth .............. Newinia

PILEOLARIACEAE
Teliospores bormesl—3ion single pedicels, wallismoothine 2 en, os Gigs pos ee Uromycladium
Teliospores depressed-globoid, 1 on pedicel, verrucose or reticulate. ................ Pileolaria

Teliospores broadly ellipsoid or longer, smooth or with projections or brick-like verrucae . . . Afelocauda

Endocyclic genus, teliospores similar to aeciospores of Atelocauda ................ Endoraecium

RAVENELIACEAE
TeELLeMOSOOLesRWItlN Cy Sto, mums ne tem aterm re met sy aot wa Gar Cac Mar ek 8s SR ee Z
TEL CHGSPOreSaNit OLE CVSLS re treet ged hares Seah wet BULB PO tas ote Mee CORN cet eas Ge oPBe 5
ZL CIOS DOLE, PEGI) nnUNLI=
hy Dilal me mpee mer Meera rete een arrears tude cis Xe sn 3
2, PECICEL SIAeNO LOLLYane e eeaeae pees Pra nce te outs Veh Ba ids MernCch i eyele enw ee he sins +
3. Teliospore heads with a patelliform cellular layer between spores and cysts. ....... Kernkampella
2. Leliospore heads witiieystsattaCmedmO SPOLCS — sis see hg he ss Ravenelia
A Pedicel atiacnecut@ tNCiCy Gis aire tem aan ata. EMAL -eoe a Casta sew See, Se ee Cystomyces
4 eedicelrattachedsouune spore) cad mami an eral Lacon snare ua. Gite tee fw lA oss Spumula
Deel ClOS DOTES Heel eC amie tanta aerate fait rRaare alg sea es AL aan Bhat Osh RR dyOrne age Boe 6
Dt CHOSDOLES OI OPEICE IOC seam carrer. ae ure ded are eS ede nts Pa Saas fosates sats: wood a ables ef
6 Leliospores lateral yireciomspedicels eave = 5s sit ieee chen ee ete) ol Seta A 7,
6. Lellospokess eInlyaiiniteculalera leur ern ce gree pare ene Ones ome ea ts Anthomyces
J slelospore peice awathouta picaltcellG mnien war aismn ate ied Coe ca hy ety i eee Rates cus ie 8e Snps on 8
7alehosporespedicelavitirapicalice) sam meaame tis Sit ete ey eee a oy A aye ers cee bao 10
S eHOsSpOres.s io lysOMes Ole sDLOA PeCICele.. ea ierlnmuot seme cs, ole teco Sok ce a's ce els Lipocystis
8, Teliospores 2 onamore.on common pedicelsior basal cells. 2a ci 5. oe ah ed es Ree 9
9. Two to several spores on common basal cells firmly united to forma cluster ..... . Anthomycetella
OP Leliospotes Dome Inky -saped: pall ONicomimon Pedicels mea. cy 2 st = Sue ee ok Ypsilospora
10> Bach apical-cellavatinistn eles @lemr aa eter oar wei tee eee, Se aM) oa nn al8s oo) ayia ce Apra
10 aE achtapicalical limianpaliaO GpOLes met It cuh fern hus Ai oA cask ee Fee Diabole
1s) Relios pores werticalivere tale ae wee teeter ewes Ce utah te cla ane Aide hb sek “i 5)arFags 12
iil ebocpetesnomichiicallyiscM tater ek a aie A GOA ic setae Atcies tbar fice) ete) opGop ye sick gs os 16
12 eliospetes withe2 corm pores per celle i eisai te cs co veyip ojala seen yo Gea 8 A ney. @ Diorchidiella

45
12. Teliospores with 1 germ pore per cell or obscure ~ <9. i eve eS 13

13s Teltespore 2 celled Rtn sie Ae ee Meee Ba treodo etm erent een eget ae nena Sphenospora

13: ‘Teliospores’2- to 4-celled by verticaliseptas: x... Seta nans Ney E-Rate ee 14

14) Veliospores mostlyi2-celled) (esi teeeats eC wen ttetotesRett Mele etree a cnirteetea eee gremecr cee 19

14. Teliospores 3-celled, 1 basal, which is attached to the pedicel and 2 apicalicellsaen sat aie Esalque

14. Teliospores 4-celled, cells arranged rectangulary ...--- +++ +e esters Allotelium

15. Teliospores mostly with blocklike warts... . 6... es ee eet eee te eee teens Dicheirinia

15, Teliospores smooth or echinulate 517 cneg ea shee Diorchidium

16. Leliospores triquetrous) | ae. shaves oe cote en ck a 17.

16. Teliospores muriform ........ FS eeoS oo) OR ee a a es a 20

17. Two teliospore cells surmounted bya third... ... 1. +. eee eee eee eee Hapalophragmium

17. One teliospore cell surmounted by two others... ..- 6. eee ee ee 18

18. Teéliospores with’ germ pore per cell) 25 =e Triphragmium

18. Teliospores with 2. serm pores per cell 2) 0 en 19

19. Teliospores verrucose with irregular warts, germ pores in outer walls... ..-. +. | Triphragmiopsis

19. Teliospores with elongate, often apically divided projections, germ pores at inner angles . Nyssopsora

20. Cells of teliospores in a more or less globoid arrangement, not seriate .......-- Sphaerophragmium

20. Cells of teliospores seriately arranged to make an elongated spore head. .........-- Cumminsina

PHRAGMIDIACEAE
1. Teliospores’I=celleds 2 4741. Ge) ete ee ee 2

1- Teliospores 2- or more celled: eyey eet. See tegr 0 ane ees ea 3

2. Teliospore wall pale brown, telia suprastomatal .....- 6 6 eee ee es Gerwasia

2. Teliospore wall pigmented, telia subepidermal .... ....

+--+ -
eee eee: Trachyspora

3. Teliospores'2-celléd 2... ame 2 ire Gk eee ee ee ce +

8. Teliospores more:thanc2-celledi...5.) tm amet cece oe ee ee 5

4. Teliospore wall yellow-brown, spermogonia type8 .... 6. - 2 ee ee es Joerstadia

4. Teliospore wall chestnut-brown, spermogonia type6 eee eee

6-2 ---
... Gymnoconia

5. ‘Teliospore wall’ colorless) 4s -ake ey ei One ee erate es oon eer aes Reurea 6

5. Teliospore wall browits a= tie te 2 eye eneect oseeten ets ee es oe cre ce 7

6. Teliospores terete, tending to lanceolate, in felty extrusions... 61 eee eee ee ee Hamaspora

6. Teliospores cylindrical, appearing Articulated notetel thikcemememei mst mrirain iat ieee ene Kuehneola

7. All teliospore cells with ligerm pore, eee e

3.592 fe es oe es 8

7. Most teliospore cells with 2 or more germ pores... 6. - - ee ee 2

8. Spermogonia type 10, on Rosaceae. . - 6 1 ee ee et ee te ee Frommeélla

8. Spermogonia type 7, on Bignoniaceae. . . . . ee ee es Newinia

46
 9. Teliospore cells with 2 or more germ pores except the apical cells............... Xenodochus
9; Teliospore:cellsywith-2.or moreger pores inal Cells a. eyrate Fla berah. eeinranatt te Phragmidium
Endocyclic genus: telia similar to aecia of Gymnoconia...........+..- (No generic name available)

PUCCINIACEAE
dd eLeliospores:1- Cellec gemege mare rer aera Se ore Pein ce de: rele ye poche yay es peat: S
lileliospores- mone tate celled kane mea mee ate atmereee on lee Se etame ab) a0 llel sens ac ee dae 2 2
2. Leliospores.2-celled; Wwitiisomee Ce pUON smanta tanya nce ne eee ee 5

2.) leliospores more tna 2-celled Gren. merase ent cme trite, Renee eel oe aus. Chrysella
3. Uredinia and telia surrounded by pallisadelike peridium..............-.-..- Corbulopsora
Sp aUrediniavand tellaswvitiOub peril e meme alee mtn trie tie eee ie eels ae ene ae 4
AM Basidiospores sessile MOtMOnGlDly sejeCleCm tm cee ty otras 04 ea tre eer og ano Zaghouania
Am basidiospores DOlNe Ol stenleimata Orel oly Clete Cary a ue ha eer eee eee: Uromyces
DW DASICIA TTCTTalm weir eM are eer een aR ihn eyes ohOra seee ee ofa aha Chrysopsora
S Dasiciaascii linterna learrnme ie we@epCL net ye AG an face ae Fos a eae dc Chrysocyclus
SF DASIGIA EX CENTALM etre ueN etree ney feert Scere ear eee rey orceM TULA ch) SaMdase) vo,vnGna) ose aeSH 6
6: Uredinia.and telia:surrounded by. palisadelike peridium)n-payq4 6 ape suk ey Se Miyagia
Ga Wredinia aid teliawithoOut pckicivil mer sess teh Mewes cnr oecias ears tye eer get) Gc crac ie 7
Je, Veliospore pedicels mostly. cel abinous a mac mete okra ee Gymmnosporangium
Tem Lehospore pedicels note elatinOus armen nero eee cient Nene: or eR MOTs) = © aypnls 8
8, Teliospores sessile tre ae satis Mele es gs eed Eee Re eyes Polioma
8. Teliospores pedicellate se rustic ass aed) sis ae sear OA CAPER RRPe MCAT NGS 9
OS Teliospores produced anicOlU MNS gam snes neste ca ie Re ee ee ya eet Kernella
Or, Leliospores net produced amecOlumiis tse. gia a Re a eae Ree Ge 10
105 Germ: pores ofteliospores tiper cell tana Res earn) eee Adee ie 11
10<Germupores ofteliospores morethanel per celle s.5 5. eet ee ee ete ee nme nay os). 22
TIS eliospore
1 pedicelssnortand determinate. ..2 ashe) eee Ree carne Mee cs as: Puccinia
11. Teliospores produced on the tips of elongating and branching hyphal cells... ... . Ramakrishnania
123 Germ) porestortenos poles. er Cell ies: ey. eter tie ins ae eee ee tr Cumminsiella
12. Germ) pores of teliospores more than:2 per cell eee: CRM a area es 13
13) Germ pores of teliosporess3 per cell... eyes 2 + 5 esa na. ene ee eee he Stereostratum
13. Germ pores.of teliospores:4 ormore per cell ayn. aco ee oun te en ce eee Cleptomyces
Endocyclic genus; telia similar to aecia of Puccinia and others, Aecidium-type.......... Endophyllum

PUCCINIOSIRACEAE
ieeTenOspores (-cellcc mri rE a ether cer @ eet Oya Nari ot ony sone pasha epee ek a ae 2

47
 1 Teliospores 2-celled yc.) syse se eel ee ee re 8
2 *Teliospores without intercalary cells... 1. sets eite sseceeneres Geka tse eek ete 3)
24 Teliospores withinterealary cells). ceqee. << Geen ee tt Se te gee 8)
3, Telia-hairlike ‘or colummats soces(25 2 seen ch oe ee acts Se 4
3. Telia not hairlike or columnar, usually compact, with peridium...............-. Baeodromus

A. Teliashairlike i.de- > ada & 2 ieee ie ems, cet core a Coca ee Cionothrix
A. Telia colummat' a < z of& cha ee ees a ee 0 Cle eg en a Alveolaria
5.. Basidia internal 22ers. eee his WG cele es Soe ea re a Trichopsora
5 Basidia externals vn 2228 sskee ncn 4)eee Stee kes rr ee Oe a eea ee a a 6
6.. Intercalary cells‘clongated, pedicel-like 9 cass 2 female uth ree Chardoniella
6. Intercalary cells short). 2% 9 pois) doi ee a ee ee 7
7.. intercalary cells conspicuOUS=.cgens 0 se teny nn ee Ceratocoma
7, Intercalary cells present butmot-consPICluiOUs rn nat meee
eee ee Dietelia
8. Intercalary:cells:conspiGuOUs! sarees tet ee Pucciniosira
8. Intercalary cells present DutmoticOnspi1cuOu si mgamss aur ue tee ieee ee eer Didymopsora

GENERA OF UNCERTAIN AFFINITY

Hiratsukamyces
Maseeélla

Skierka

48
DESCRIPTIONS AND ILLUSTRATIONS
OF HOLOMORPH GENERA

49
MCI AATAGLY GAN 2a
‘RAIV40 HIROMDION I :
_
 CALYPTOSPORA J.G. Kuhn
Hedwigia 8:81-82 (81). 1869.

FAMILY: Pucciniastraceae American form, especially in the aecial state on

Abies spp. (Hiratsuka et al. 1967; Faull 1939; Krebill
DESCRIPTION: 1972). Comparative studies of North American
Spermogonia subcuticular, Group I (type 3). Aecia forms and European material (type locality) are
hypophyllous, subepidermal, cylindrical, peridium necessary.
colorless, firm, Peridermium-type; peridial cells Host alternation is believed to be obligatory, but
irregularly polygonal; aeciospores globose, sub- it has been suggested that Vaccinium-to-Vaccinium
globose to broadly ellipsoidal, contents orange-red. infection by basidiospores occurs. A few inoculation
Uredinia not produced. Telia caulicolous, forming a experiments have been done with negative results
continuous layer around the abnormally elongated (Ziller 1974).
and thickened stems; teliospores intracellular,
formed in the epidermal cells, subglobose, ellip- REFERENCES:
soidal, oblong or prismatic, 3 to 5 cells (mostly 4), Faull, J.H. 1939. A review and extension of our knowledge of
with vertical septa, walls golden-brown, germ pores Calyptospora goeppertianum Kuehn. J. Arnold Arbor. Harv.
Univ. 20:104-113.
1 on each cell close to the junction of cells, in the
Gaumann, E. 1959. Die Rostpilze Mitteleuropas. Buchdruckerei
center of spore balls; basidia external.
Buchler & Co., Bern. 1407 p.
Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.
TYPE: Calyptospora goeppertiana J.G. Kuhn Kasai Publ. Co., Tokyo. 167 p.
On: Vaccinium vitis-idaea L. (Ericaceae) Hiratsuka, Y.; McArthur, L.E.; Emond, FJ. 1967. A distinction
Type locality: Karpacz near Jelenia Gora, Poland between Pucciniastrum goeppertianum and P. epilobii with clari-
(Krummhtbel, Germany) fication of status of Peridermium holwayi and P. ornamentale.
Can. J. Bot. 45:1913-1915.
REMARKS: Krebill, R.G. 1972. Pucciniastrum goeppertianum in Abies lasiocarpa
in the Rocky Mountain states. Am. Midl. Nat. 87:570-576.
This genus, together with Thekopsora and
Kuhn, J.G. 1869. Calyptospora nov. gen. Uredinearum. Hedwigia
Pucciniastrum, has been included in the genus 8:81-82.
Pucciniastrum by some authors (Arthur and Sato, S.; Katsuya, K.; Hiratsuka, Y. 1993. Morphology, taxonomy
Cummins 1960), but others recognize them as sepa- and nomenclature of Tsuga-Ericaceae rusts. Trans. Mycol. Soc.
rate taxa (Gaumann 1959; Hiratsuka 1958; Sato et al. Jpn. 34:47—-62.
1993). Only one species, Calyptospora goeppertiana Ziller, W.G. 1974. The forest tree rusts of western Canada. Envi-
ron. Can., Can. For. Serv., Victoria, BC. Publ. 1329. 272 p.
J.G. Kuhn, is recognized, but the form in western
Canada seems to be different from the eastern North

FIGURE 19. Calyptospora. C. goeppertiana J.G. Kuhn. A. Aecia of the eastern North American form. B. Surface view of
teliospores.

Syl
 HYALOPSORA Magnus
Ber. Dtsch. Bot. Ges. 19:578-584 (582). 1901 (issued 1902).

FAMILY: Pucciniastraceae REMARKS:

Hiratsuka (1958) recognized seven species of
DESCRIPTION: Hyalopsora and five species of Uredo suspected to be-
Spermogonia subepidermal, Group I (type 2). long to this genus. Hyalopsora is generally similar to
Aecia subepidermal, erumpent, with peridium, Milesina except that there is pigment in the cyto-
Peridermium-type; aeciospores catenulate, plasm; the sori are yellow when fresh, but the color
verrucose. Uredinia subepidermal, with peridium, fades rapidly. The teliospores are not resting spores
Milesia-type, sometimes with rudimentary or but mature in the spring on overwintered fronds
without specialized ostiolar cells, thin-walled and then germinate. Amphispores are produced in
paraphyses sometimes also present; urediniospores addition to ordinary urediniospores and may pre-
borne singly, wall colorless, but pigment present in dominate. Insofar as the life cycles are known, the
the cytoplasm, wall verrucose or echinulate, pores species are heteroecious with aecia on Abies (firs)
mostly scattered, obscure. Telia scarcely organized; and uredinia and telia on ferns.
teliospores produced in the epidermal cells, 2- to
many-celled by vertical septa, sessile, wall thin, REFERENCES:
colorless, pore 1 in outer wall of each cell, obscure, Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.
Kasai Publ. Print. Co., Tokyo. 167 p.
germination occurs without dormancy; basidium
Magnus, P. 1901 (issued 1902). Weitere Mittheilung ber die auf
external. Farnkrautern auftretenden Uredineen. Ber. Dtsch. Bot. Ges.
19:578-584.
TYPE: Hyalopsora aspidiotus (Magnus) Magnus Ziller, W.G. 1974. The tree rusts of western Canada. Environ.
(Lectotype) Can., Can. For. Serv., Victoria, BC. Publ. 1329. 272 p.
=Melampsorella aspidiotus Magnus
On: Phegopteris dryopteris (L.) Fee
(Polypodiaceae)
Type locality: USA

FIGURE 20. Hyalopsora. A. An amphispore and a urediniospore of H. polypodii (Dietel) Magnus. B. Urediniospores of
H. chelianthis Arthur. C. Telia of H. aspidiotus (Magnus) Magnus. D. Amphispores of H. aspidiotus.

uw Nw
 MELAMPSORELLA J. Schrot.
Hedwigia 13:81-85 (85). 1874.

FAMILY: Pucciniastraceae REMARKS:

The two species are heteroecious and macrocyclic
DESCRIPTION: (Hiratsuka 1958). The type species produces con-
Spermogonia subcuticular, Group I (type 3). Aecia spicuous witches’ brooms on Abies and uredinia and
subepidermal, erumpent, Peridermium-type; aecio- telia on systemically infected Cerastium and Stellaria.
spores catenulate, verrucose. Uredinia subepidermal, Reports of aecia on Picea are erroneous. Melamp-
with peridium opening by a pore delimited by ostiolar sorella symphyti Bubak produces aecia on current-
cells, Milesia-type; urediniospores borne singly, wall season leaves of Abies without forming brooms, and
colorless, echinulate, pores obscure. Telia scarcely uredinia and telia on Symphytum (Boraginaceae).
organized, consisting of closely packed but loosely Neither species has resting teliospores.
adherent, 1-celled teliospores in the epidermal cells,
wall thin, colorless, germ pore obscure; germination REFERENCES:
occurs without dormancy through the outer wall of Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.
Kasai Publ. Print. Co., Tokyo. 167 p.
the spore; basidia external.
Pady, S.M. 1946. The development and germination of
intraepidermal teliospores of Melampsorella cerastii.
TYPE: Melampsorella caryophyllacearum J. Schrot. Mycologia 38:477-499.
On: Sfellaria uliginosa Murr. (Caryophyllaceae) Schroter, J. 1874. Melampsorella, eine neue Uredineen Gattung.
Type locality: Germany Hedwigia 13:81-85.

FIGURE 21. Melampsorella. M. caryophyllacearum J. Schrot. A. Urediniospores. B. Germinating teliospores.

53
 MELAMPSORIDIUM Kleb.
Z. Pflanzenkr. 9:14-26 (21-22). 1899.

FAMILY: Pucciniastraceae one at each end in M. alni (Thtim.) Dietel, bizonate in

M. betulinum and M. hiratsukanum S. Ito (2-3 + 2-3),
DESCRIPTION: equatorial in M. asiaticum Kaneko & Hirats. f. (2-5),
Spermogonia subcuticular, Group I (type 3). and subequatorial in M. carpini (Fuckel) Dietel (2-5).
Aecia subepidermal, erumpent, with peridium, Telia are not often collected because they develop
Peridermium-type; aeciospores catenulate, verru- late in autumn. Roll-Hansen and Roll-Hansen (1981)
cose. Uredinia subepidermal, with peridium open- suggested, from inoculation and morphology, that
ing by a discrete pore delimited by differentiated M. alni is synonymous with M. betulinum, but
ostiolar cells, Milesia-type; urediniospores borne Kaneko and Hiratsuka (1981, 1982) reported that
singly, echinulate, wall colorless, position and there are differences in urediniospore germ pores.
number of germ pores variable (see Remarks). Telia Kurkela et al. (1999) reported M. hiratsukanum for
subepidermal, not erumpent, consisting of crusts 1 the first time in Europe (Finland).
spore deep of closely packed spores; teliospores
sessile, 1-celled, with thin colorless walls, germ pore REFERENCES:
apical if differentiated; germination occurs after Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.
wintering on fallen leaves; basidia external, basidio- Kasai Publ. Print. Co., Tokyo. 167 p.
Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
spores globoid.
Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
Nakayama, K. 1992. The rust flora of Japan. Tsukuba
TYPE: Melampsoridium betulinum (Fr.) Kleb. Shuppankai, Ibaraki. 1205 + 159 p.
=Sclerotium betulinum Fr. Kaneko, S.; Hiratsuka, N. 1981. [Classification of the Melamp-
On: Betula verrucosa Ehrh. (Betulaceae) soridium species based on the position of urediniospore germ
Type locality: Sweden pores.] Trans. Mycol. Soc. Japan. 22:463-473. (In Japanese.)
Kaneko, S.; Hiratsuka, N. 1982. Taxonomic significance of the
urediniospore germ tubes in the pucciniastraceous and
REMARKS:
melampsoraceous rust fungi. Trans. Mycol. Soc. Jpn. 23:201—
Melampsoridium is a genus of six heteroecious and 210.
one presumably heteroecious species (Hiratsuka Kaneko, S.; Hiratsuka, N. 1983. A new species of Melampsoridium
1958; Hiratsuka et al. 1992; Singh and Pandy 1972). on Carpinus and Ostrya. Mycotaxon 18:14.
The aecia of M. betulinum (Fr.) Kleb. occur on needles Klebahn, H. 1899. Kulturversuche mit heterécischen Rostpilzen.
of Larix and the telia on leaves of Alnus and Betula. Z. Pflanzenkr. 9:14—26.
The ostiolar cells of the uredinial peridium have a Kurkela, T.; Hanso, M.; Hantula, J. 1999. Differentiating charac-

long spinelike apex in most species, making such teristics between Melampsoridium rusts infecting birch and
alder leaves. Mycologia 91:987-992.
species easy to recognize when uredinia are present;
Roll-Hansen, F.; Roll-Hansen, H. 1981. Melampsoridium on Alnus
this is the obvious spore state. Melampsoridium in Europe. M. alni conspecific with M. betulinwm. Eur. J. For.
inerme Suj. Singh & P.C. Pandey, on Magnolia from Pathol. 11:77-87.
India, is described as lacking the spinescent ostiolar Singh, S.; Pandy, P.C. 1972. Melampsoridium inerme on Magnolia.
cells (Singh and Pandey 1972). Position and number Trans. Br. Mycol. Soc. 58:342-344.
of urediniospore germ pores vary with the species:

uw +
FIGURE 22. Melampsoridium. A. Urediniospores. B. A telium of M. betulinum (Fr.) Kleb. (courtesy of Dr. S. Kaneko).

535)
 MILESINA Magnus
Ber. Dtsch. Bot. Ges. 27:320-327 (325). 1909.

FAMILY: Pucciniastraceae to this genus. As in Uredinopsis, all spore states lack

pigment and hence the sori are white. All species are
DESCRIPTION: presumably heteroecious, with aecia on the leaves
Spermogonia subepidermal, Group I (type 1). of Abies (firs) and the uredinia and telia on ferns. The
Aecia subepidermal, erumpent, with white perid- teliospores are not resting spores, but develop dur-
ium, Peridermium-type; aeciospores catenulate, ing the winter and germinate in the spring. Some
verrucose. Uredinia subepidermal, erumpent, species produce amphispores as well as ordinary
Milesia-type, dome shaped, with peridium, opening urediniospores. The name Milesia has often been
by an irregular pore without clearly differentiated used, especially in North American literature
ostiolar cells; urediniospores borne singly, mostly (Arthur and Cummins 1962; Faull 1932), but the
echinulate, but sometimes verrucose or smooth, name was based on a uredinial (anamorphic) state,
pores bizonate, obscure. Telia scarcely differenti- and therefore is used as a morphological type of
ated, consisting of spores in the epidermal cells; anamorphs in this manual.
teliospores few- to many-celled by vertical septa,
sessile, germ pores 1 in outer wall of each cell, REFERENCES:
obscure; the teliospores of most species developing Arthur, J.C.; Cummins, G.B. 1962. Manual of the rusts in United
in overwintered fronds and germinating without States and Canada with supplement. Hafner Publ. Co., New
York. 438 + 24 p.
dormancy; basidia external.
Faull, J.H. 1932. Taxonomy and geographical distribution of the
genus Milesia. Contrib. Arnold Arbor. Harvard Univ. 2. 138 p.;
TYPE: Milesina kriegeriana (Magnus) Magnus pl. 1-9.
(Lectotype) Henderson, D.M.; Prentice, H.T. 1977. The morphology of fungal
=Melampsorella kriegeriana Magnus spores: Milesina blechni. Notes R. Bot. Gard. Edinb. 35:415—
On: Dryopteris austriaca Jacq. (Polypodiaceae) 417; pl. 6-7.
Type locality: Germany Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.
Kasai Publ. Print. Co., Tokyo. 167 p.
Magnus, P. 1909. Bemerkungen tuber einige Gattungen der
REMARKS:
Melampsoreen. Ber. Dtsch. Bot. Ges. 27:320-327.
Hiratsuka (1958) recognized 34 species of
Milesina and 22 species of Uredo suspected to belong

FIGURE 23. Milesina. A. Urediniospores of M. vogesiae Syd. B. Urediniospores of M.

dieteliana (Syd.) Magn. C. Teliospores of M. pycnograndis (Arthur) Hirats. f.
 NAOHIDEMYCES S. Sato, Katsuya & Y. Hirats.
Trans. Mycol. Soc. Jpn. 34:47—-62 (48). 1993.

FAMILY: Pucciniastraceae 1993). Both alternate between Tsuga spp. (0, I on

Pinaceae) and Vaccinium spp. (I, HI on Ericaceae).
DESCRIPTION: The most distinctive feature of this genus is the pres-
Spermogonia subcuticular without bounding ence of Milesia-type aecia rather than Peridermium-
structure, Group I (type 3). Aecia subepidermal, type aecia. This is the only genus, so far recognized,
with domelike peridium with conspicuous ostiolar having Milesia-type aecia among heteroecious rusts.
cells, Milesia-type; aeciospores echinulate, borne singly The position of the teliospore germ pore is different
on inconspicuous pedicels. Uredinia with domelike from that in Thekopsora species; Naohidemyces has
peridium with ostiolar cells, Milesia-type; uredinio- germ pores at the center of each teliospore cell rather
spores echinulate, borne singly on inconspicuous than at the corner of each cell at the center of the
pedicels. Telia scarcely differentiated; teliospores spore balls as in Thekopsora species.
one cell deep, consisting of several laterally adher-
ent cells within epidermal cells of the host, one germ REFERENCES:
pore per cell located in the center of the cell, wall Hiratsuka, Y. 1965. The identification of Uraecium holwayi on hem-
pigmented; germination occurs after dormancy lock as the aecial state of Pucciniastrum vaccinii in Western
North America. Can. J. Bot. 43:475-478.
(overwintering) on dead leaves; basidia external.
Hiratsuka, Y.; Sato, S. 1976. [Species of Thekopsora on Tsuga.]
Trans. Mycol. Soc. Jpn. 17:543-548. (In Japanese.)
TYPE: Naohidemyces vaccini (G. Winter) S. Sato,
Sato, S.; Katsuya, K. 1979. Heteroecism of two rust fungi on
Katsuya & Y. Hirats. needles of Tsuga diversifolia and T. sieboldii. Trans. Mycol. Soc.
=Melampsora vaccini G. Winter Jpn. 20:1-4.
On: Vaccinium myrtillus L. (Ericaceae) Sato, S.; Katsuya, K.; Hiratsuka, Y. 1993. Morphology, taxonomy
Type locality: St. Gallen, Switzerland and nomenclature of Tsuga-Ericaceae rusts. Trans. Mycol. Soc.
Jpn. 34:47-62.
REMARKS:
The type species and another one, N. fujisanensis
S. Sato, Katsuya & Y. Hirats., are known (Sato et al.

FIGURE 24. Naohidemyces. A. Aecia of N. vaccinii (G. Winter) S. Sato, Katsuya & Y. Hirats.
B. Ostiolar cells of a uredinium of N. vaccinii. C. An aeciospore of N. fujisanensis
S. Sato, Katsuya & Y. Hirats. D. Teliospores of N. vaccinii. E. Surface view of
teliospores of N. fujisanensis. (Reprinted with permission, from Sato et al. 1993.)

Sy/f
 PUCCINIASTRUM G.H. Otth
Mitt. Naturforsch. Ges. Bern 1861:57-88 (71). 1861.

FAMILY: Pucciniastraceae and Tsuga, and the uredinia and telia on many
dicotyledonous plants, including the Aceraceae,
DESCRIPTION: Betulaceae, Ericaceae, Fagaceae, Onagraceae, and
Spermogonia subcuticular, Group I (type 3). Rosaceae. Two species of Uredo (Milesia) that will
Aecia subepidermal, erumpent, with peridium, prove to belong in Pucciniastrum are known on
Peridermium-type; aeciospores catenulate and Goodyera spp. (Orchidaceae). We follow Sato et al.
verrucose. Uredinia subepidermal, with peridium, (1993) and recognize Pucciniastrum, Thekopsora, and
opening by a pore delimited by differentiated Calyptospora as separate genera, and also recognize
ostiolar cells, Milesia-type; urediniospores borne Naohidemyces having Milesia-type aecia.
singly, wall colorless, echinulate, pores scattered,
obscure. Telia 1 spore deep, consisting of REFERENCES:
subepidermal crusts of laterally adherent telio- Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.
spores; teliospores sessile, 2- or more-celled by ver- Kasai Publ. Print. Co., Tokyo. 167 p.

tical septa, germ pore 1 in the outer wall of each cell, Hiratsuka, Y.; McArthur, L.E.; Emond, FJ. 1967. A distinction
between Pucciniastrum goeppertianum and P. epilobit with clari-
wall pigmented; germination occurs after dor-
fication of status of Peridermium holwayi and P. ornamentale.
mancy; basidia external. Can. J. Bot. 45:1913-1915.
Otth, G.H. 1862. Uber die Brand- und Rostpilze. Mitt.
TYPE: Pucciniastrum epilobii G.H. Otth Naturforsch. Ges. Bern 1861:57-88.
On: Epilobium angustifolium L. (Onagraceae) Pady, S.M. 1933. Teliospore development in the Pucciniastreae.
Type locality: Switzerland Can. J. Res. 9:458-485.
Sato, S.; Katsuya, K.; Hiratsuka, Y. 1993. Morphology, taxonomy
REMARKS: and nomenclature of Tsuga-Ericaceae rusts. Trans. Mycol. Soc.
Jpn. 34:47-62.
Hiratsuka (1958) listed 23 species in this genus
Ziller, W.G. 1974. The tree rusts of western Canada. Environ.
and 14 Uredo (Milesia) species suspected to belong to Can., Can. For. Serv., Victoria, BC. Publ. 1329. 272 p.
it. All species whose life cycles are known are
heteroecious, with aecia on needles of Abies, Picea,

FIGURE 25. Pucciniastrum. P. epilobii Otth. A. Cross section of teliospores. B. Surface view of teliospores.
 THEKOPSORA Magnus
Sitzungsber. Ges. Naturf. Freunde Berl. p. 57-60 (58). 1875.

FAMILY: Pucciniastraceae obviously belong to this genus were also reported.

After removal of two species with Milesia-type aecia
DESCRIPTION: to Naohidemyces, 11 species remain in Thekopsora.
Spermogonia subcuticular, Group I (type 3). Two additional uredinial states (Milesia-type)
Aecia cylindrical, peridium delicate, nearly color- probably belong in Thekopsora (Hiratsuka 1958; Sato
less or pale yellow, Peridermium-type, peridial cells et al. 1993). Telia of three Thekopsora species are on
irregularly polygonal; aeciospores globose, sub- Ericaceae, three are on Rubiaceae, and five are on
globose to broadly ellipsoidal. Uredinia subepi- other dicotyledonous families. Aecia of three
dermal, peridium applanate hemispherical to species are known: two infect Picea and one Tsuga.
conoidal, Milesia-type, peridial cells irregularly Of the 11 species, 10 occur in and near Japan; 8 of
polygonal, ostiolar cells developed. Telia amphi- them are also in Eurasia, and four in North America.
genous; teliospores forming within the epidermal
cells, 3- to 6-celled, with vertical septa, germ pores 1 REFERENCES:
for each cell at the center of spore balls; basidia Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
external. fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
Hiratsuka, N. 1958. Revision of taxonomy of the Pucciniastreae.

TYPE: Thekopsora areolata (Fr.) Magnus Kasai Publ. Print. Co., Tokyo. 168 p.
Magnus, P. 1875. Uber die Familie der Melampsoreae und ihre
=Xyloma areolatum Fr.
Gattungen. Sitzungsber. Ges. Naturf. Freunde Berl. p. 57-60.
On: Prunus padus L. (Rosaceae) Sato, S.; Katsuya, K.; Hiratsuka, Y. 1993. Morphology, taxonomy
Type locality: Smaland, Sweden and nomenclature of Tsga-Ericaceae rusts. Trans. Mycol. Soc.
Jpn. 34:47-62.
REMARKS:
Thirteen species were recognized by Hiratsuka
(1958) in this genus, and a few Uredo species that

ee

eae

FIGURE 26. Thekopsora. Thekopsora minima P. & H. Syd. A. Aecia on Tsuga diversifolia (Maxim.)
Mast. B. Cross section of teliospores. C. Surface view of teliospores.

SN)
 UREDINOPSIS Magnus
Atti Congr. Int. Bot. Genova, 1892. p. 163-168 (167). 1893.

FAMILY: Pucciniastraceae REMARKS:

Hiratsuka (1958) recognized 26 species of
DESCRIPTION: Uredinopsis and two Uredo species suspected to be-
Spermogonia Group I (type 1, 2, or 3). Aecia long to this genus. Uredinopsis is usually considered
subepidermal, erumpent, with white peridium, to be the most primitive existing genus of the
Peridermium-type; aeciospores catenulate, verru- Uredinales, in part because the aecial states are on
cose, colorless. Uredinia subepidermal, with leaves of the genus Abies (firs) and the simple telia
peridium, opening irregularly, ostiolar cells not on ferns, and in part because no pigment occurs in
clearly differentiated, Milesia-type; urediniospores the cytoplasm or the cell walls. Presumably all
borne singly, often extruded as white tendrils, typi- species are heteroecious and macrocyclic.
cally more or less lanceolate and apically
mucronate, colorless, smooth or with a few lines of REFERENCES:
coglike warts, germ pores near the ends; amphi- Faull, J.H. 1938. Taxonomy and geographical distribution of the
spores occur in some species. Telia as such not orga- genus Uredinopsis. Contrib. Arnold Arbor. Harv. Univ. XI.
120 p.
nized, the teliospores single-celled or 2- to 6-celled
Hiratsuka, N. 1958. Revision of taxonomy of Pucciniastreae.
with vertical or oblique septa, colorless, germ pore 1
Kasai Publ. Print. Co., Tokyo. 167 p.
in outer wall, obscure; germination occurs in Over- Magnus, P. 1893. Uber den Protomyces(?) filicinus Niessl. Atti
wintered fronds; basidia external. Congr. Int. Bot. Genova, 1892. p. 163-168.
Ziller, W.G. 1974. The tree rusts of western Canada. Environ.
TYPE: Uredinopsis filicina Magnus Can., Can. For. Serv., Victoria, BC. Publ. 1329. 272 p.
On: Phegopteris polypodioides Fée (Polypodiaceae)
Type locality: Bad Gastein, Austria

ronengr
<M
vse

FIGURE 27. Uredinopsis. A. Teliospores of U. longimucronata Faull. B. Urediniospores

of U. pteridis Dietel. C. Urediniospores of U. osmundae Magnus.

60
 CEROPSORA B.K. Bakshi & Suj. Singh
Can. J. Bot. 38:259-262 (260). 1960.

FAMILY: Coleosporiaceae vestigial peridium. The species doubtless is micro-

cyclic.
DESCRIPTION: This genus was considered within the family
Spermogonia, aecia, and uredinia unknown. Chaconiaceae (Cummins and Hiratsuka 1983), but
Telia subepidermal, becoming erumpent as waxy Ono and Hennen (1983) excluded it from that family
cushions; teliospores 1-celled, borne singly on com- and suggested a relationship with Chrysomyxa. The
pacted, parallel, multiseptate basal cells, the spores closest genus obviously is Chrysomyxa (microcyclic
laterally adherent when young but separating later, form) and it may be logical to classify it with Chry-
the wall colorless, without a differentiated germ somyxa species that have well-developed telial basal
pore, germination occurring without dormancy by cells, as a separate genus (e.g. Stilbechrysomyxa).
continued elongation of the apices of spores; basidia However, until a taxonomic study of Chrysomyxa
semi-external. and related genera is completed, we will retain this
as a Separate genus.
TYPE: Ceropsora piceae (Barclay) B.K. Bakshi & Suj.
Singh REFERENCES:
=Chrysomyxa piceae Barclay Bakshi, B.K.; Singh, S. 1960. A new genus in plant rusts. Can. J.
On: Picea smithiana (Wall.) Boiss (Pinaceae) Bot. 38:259-262.
Type locality: India Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
REMARKS: Ono, Y.; Hennen, J.F. 1983. Taxonomy of the Chaconiaceous
Only the type species is known. The telia have ill- genera (Uredinales). Trans. Mycol. Soc. Jpn. 24:369-402.
defined peripheral cells, which perhaps represent a

FIGURE 28. Ceropsora. C. piceae (Barclay) B.K. Bakshi & Suj. Singh. A. Telial sorus on Picea smithiana. B. Cross section
of a telium. C. Teliospores. D. A basidium with basidiospores.

61
 CHRYSOMYXA Unger
Beitr. Vgl. Pathol. [:1-25 (24-25). 1840.

FAMILY: Coleosporiaceae Crane et al. (2000b) discovered a unique dissemi-

nation mechanism in the microcyclic species C.
DESCRIPTION: weirii. The teliospores separate and disperse by
Spermogonia subepidermal, Group I (type 2). water to young spruce needles before they ger-
Aecia subepidermal, erumpent, with peridium, minate to produce a two-celled basidium and
Peridermium-type; aeciospores catenulate, basidiospores. In most species, teliospores germi-
verrucose. Uredinia subepidermal, erumpent, with nate in the telium and are dispersed by windborne
or without an inconspicuous peridium, Caeoma- basidiospores.
type; urediniospores catenulate, verrucose, the Thirumalachar (1951) considered Coleopucciniella
urediniospores and aeciospores quite similar within Hara as a synonym of Chrysomyxa and transferred
a species, pores scattered. Telia subepidermal, Coleopucciniella simplex (Dietel) Hara to Chrysomyxa
erumpent; teliospores 1-celled, catenulate, crowded (C. simplex (Dietel) Thirum.). However, Laundon
but loosely adherent, wall thin, pale; germination
(1975) clearly showed that the telial morphology is
occurs without dormancy; basidia external.
closer to Gymnosporangium than to Chrysomyxa.
Hiratsukaia, based on H. tsugae (Hirats. f.) Hara
TYPE: Chrysomyxa abietis (Wallr.) Unger
(Chrysomyxa tsugae Hirats. f.) may not even bea rust
=Blennoria abietis Wallr.
and is excluded here.
On: Picea abies Karst. (Pinaceae)
We presently retain Chrysomyxa piceae Barclay in
Type locality: Austria
the genus Ceropsora as Ceropsora piceae (Barclay) B.K.
REMARKS: Bakshi & Suj. Singh because it has a single row of te-
About 30 species are recognized. Except for five liospores in the telium. However, it may be related
microcyclic species, C. abietis (Wallr.) Unger, C. weirti to other Chrysomyxa species with well-developed
H.S. Jacks., C. keteleeriae (F.L. Tai) Y.C. Wang & R.S. telial basal cells (C. gilianensis Y.C. Wang et al.;
Peterson, C. deformans (Dietel) Jacz., and C. tsugae- C. tsugae-yunnanensis Teng; C. keteleeriae (F.L. Tai)
yunnanensis Teng (life cycles of the last two species Y.C. Wang & R.S. Peterson =Cronartium keteleeriae
are not proven), the species are heteroecious, with EL. Tai; C. succinea (Sacc.) Tranzschel, C. stilbae, and
uredinia and telia on Ericaceae sensu lato (Rhodo- C. himalensis Barclay). The last three species have
dendroideae, Pyroloideae, Empetraceae, Vac- been transferred to a new genus, Stilbechrysomyxa,
cinioideae) and Aquifoliaceae. The aecia occur on by Chen (1984). However, it may not be possible to
needles, buds, and cones of mostly Picea, but also on separate the two groups because it seems that there
Tsuga and Keteleeria species. No resting spores occur are various degrees of basal cell development. More
in the life cycles. Telia develop on overwintered, careful work is needed to clarify this problem.
persistent leaves and the teliospores germinate Crane (2001) clearly indicated that at least 7 mor-
without dormancy. Chrysomyxa arctostaphyli Dietel, phologically distinct taxa (species), including three
C. himalensis Barclay, C. menziesiae Dietel, C. perlaria new species, exist in the so-called Chrysomyxa ledi
Li Bin, C. gilianensis Y.C. Wang et al., C. stilbae Y.C. complex.
Wang et al., and C. woroninti Tranzschel are demi- Heavy infections in high-value spruce seed
cyclic. Perennial witches’ brooms on spruce bear the orchards and cone collection areas by C. pirolata can
aecia of C. arctostaphyli, and its telia are on Arcto- be economically significant. Very heavy infections
staphylos uva-ursi (L.) Spreng. Aecia are not known of needle rusts (C. ledicola and C. ledi complex) have
for all species and others are assigned with some often been reported. Heavy infection for 2 or more
doubt. Crane et al. (2000a) confirmed the life cycle of consecutive years could be economically significant
a spruce bud rust, C. woroninii Tranzschel, by inocu- in Christmas tree plantations or high-value com-
lation experiments; the connection with witches’ mercial stands, but the long-term impact on the
brooms on Ledum spp. had been speculated by many health of spruce is not known.
authors (Ziller 1974; Hiratsuka 1987). McBeath (1983)
suggested from field observations that this rust may REFERENCES:
be capable of reinfecting spruce without going Berndt, R. 1999. Chrysomyxa rust: morphology and ultrastructure
through alternate host states in arctic conditions. of D-haustoria, uredinia, and telia. Can. J. Bot. 77:1469-1484.

62
Chen, M.M. 1984. A new genus, Stilbechrysomyxa Chen gen. nov., McBeath, J.H. 1983. Symptomology on spruce trees and spore char-
of Chrysomyxaceae on Rhododendron. Sci. Silvae Sin. 20:267— acteristics of a bud rust pathogen. Phytopathology 74:456—461.
DIL. Savile, D.B.O. 1950. North American species of Chrysomyxa. Can.
Crane, P.E. 2001. Morphology, taxonomy, and nomenclature of J. Res. C, 28:318-330.
the Chrysomyxa ledi complex and related rust fungi on spruce Savile, D.B.O. 1955. Chrysomyxa in North America—additions
and Ericaceae in North America and Europe. Can. J. Bot. 79: and corrections. Can. J. Bot. 33:478-496.
957-982. Thirumalachar, M.J. 1951. Critical notes on some plant rusts III.
Crane, P.E.; Hiratsuka, Y.; Currah, R.S. 2000a. Clarification of the Mycologia 52:688-693.
life cycle of Chrysomyxa woroninii (Uredinales). Mycol. Res. Unger, F. 1840. Chrysomyxa abietis. Eine neue Krankheitsform der
104:581-586. Fichte. Beitr. Vgl. Pathol. I:1—25.
Crane, P.E.; Hiratsuka, Y.; Currah, R.S. 2000b. Reproductive Wang, Y.C.; Peterson, R.S. 1982. On Keteleeria needle rust. Acta
biology and evidence for water dispersal of teliospores in Mycol. Sin. 1:15-18.
Chrysomyxa weirii, a microcyclic spruce needle rust.
Wang, Y.C.; Wu, X.B.; Li, B. 1987. A new spruce needle rust fun-
Mycologia 92:754-763.
gus. Acta Mycol. Sin. 6:86-88.
Hiratsuka, Y. 1987. Forest tree diseases of the prairie provinces.
Ziller, W.G. 1974. The tree rusts of Western Canada. Environ.
Can. For. Serv., North. For. Cent., Edmonton, AB. Inf. Rep.
Can., Can. For. Serv., Victoria, BC. Publ. 1329. 272 p.
NOR-X-286. 142 p.

see

FIGURE 29. Chrysomyxa. A. Telium of C. ledicola (Arthur) P. & H. Syd. B. Stalked telium of
C. succinea (Sacc.) Tranzschel (reprinted with permission, from Hiratsuka and
Sato 1969). C, D. C. ledi de Bary sensu stricto. Urediniospore with groove (C)
and surface ornamentation of urediniospore (D). E. Aeciospores of C. cassandrae
Tranzschel with vertical cap (courtesy of Dr. P.E. Crane). F, G. C. rhododendri
de Bary. Inner surface (F) and outer surface (G) of aecial peridium. (A, C, D, F,G
reprinted with permission, from Crane 2001).

63
 COLEOSPORIUM Lév.
Ann. Sci. Nat. Bot. ILL, Ser. 8:369-376 (373). 1847.

FAMILY: Coleosporiaceae dicotyledonous plants. Uredinial and telial states of

more species occur on Asteraceae than on any other
DESCRIPTION: family. Most species are macrocyclic, but there are
Spermogonia subepidermal, Group I (type 2). life cycle variants. There are three demicyclic,
Aecia subepidermal, erumpent, with conspicuous autoecious species on Asteraceae, C. incompletum
peridium, Peridermium-type; aeciospores cate- Cummins, C. reichei Dietel, and C. viguierae Dietel &
nulate, verrucose with rodlike columns or knobs, Holw., and five microcyclic species on Pinus, C.
some with annulate sides, or broadly conical. crowellii Cummins, C. pinicola (Arthur) Arthur, C.
Uredinia subepidermal, erumpent, Caeoma-type, himalayense Durrieu, C. pini-densiflorae Zinno & S.
bright orange when fresh, fading to whitish; Kaneko, and C. pini-pumilae Azbukina. These species
urediniospores catenulate, verrucose, echinulate, or follow Tranzschel’s Law in that they produce
reticulate in some species, generally as the aecio- basidia on the aecial host of macrocyclic species.
spores of the species, pores scattered, obscure. Telia Arthur (1934) treated C. pinicola as a derivative of C.
subepidermal, erumpent as low or rarely columnar inconspicuum Arthur. Those who believe that micro-
cushions, gelatinous when wet; teliospores sessile, cyclic species should be segregated in a distinct
1-celled, in 1-layered crusts, or pseudocatenulate by genus because of catenulate basidia overlook the
intrusion of young spores among older spores, or occurrence of catenulate basidia in some macro-
catenulate, wall thick and gelatinizing above; ger- cyclic species. The relationship of the microcyclic
mination occurs without dormancy by division of species is obvious.
the protoplast into a 4-celled internal basidium, each Arthur (1906) chose C. rhinanthacearum (DC.) Kickx
cell producing a sterigma and one basidiospore, (=<Uredo rhinanthacearum DC.) as the designated
basidiospores ellipsoid to globoid. lectotype from five species listed in Léveillé’s original
description, but the species lacks a telial description.
TYPE: Coleosporium campanulae (F. Strauss) Tul. Laundon (1975) chose Uredo tremellosa var. campanulae
(Lectotype, Laundon 1975) F. Strauss (=C. campanulae (F. Strauss) Tul.) as the
=Uredo tremellosa F. Strauss var. new lectotype, stating that the description is clearly
campanulae F. Strauss telial despite its Uredo name.
On: Campanula sp. (Campanulaceae)
Type locality: Europe REFERENCES:
Arthur, J.C. 1906. Résultats Scientifiques du Congrés Interna-
REMARKS: tional de Botanique Vienne. 1905:331-348.
Coleosporium has many described species, some Arthur, J.C. 1934. Manual of the rusts in United States and
of which are doubtfully distinct morphologically, Canada. Purdue Res. Found., Lafayette, IN. Reprinted witha
but SEM studies (Hiratsuka and Kaneko 1975) of the supplement by G.B. Cummins (1962), Hafner Publ. Co., New
York. 438 + 24 p.
surface sculpture of aeciospores and urediniospores
Cummins, G.B. 1978. Rust fungi on legumes and composites in
show greater variability than previously recognized. North America. Univ. Arizona Press, Tucson, AZ. 424 p.
Further, Kaneko (1981) has recognized two types of Henderson, D.M.; Prentice, H.T. 1974. Spore morphogenesis of
basidial development. In type 1, the entire cell be- Coleosporium tussilaginis. Trans. Br. Mycol. Soc. 63:431—435.
comes a basidium without change in length; in type Hiratsuka, N.; Kaneko, S. 1975. Surface structure of Coleosporium
2, when the cell matures a septum forms that divides spores. Rep. Tottori Mycol. Inst.12:1-13.
the upper basidial part from a basal stalklike part. Kaneko, S. 1981. The species of Coleosporium, the causes of pine
needle rusts, in the Japanese Archipelago. Rep. Tottori Mycol.
Basidiospore shape varies also (Kaneko 1981;
Inst. 19:1—-159.
Cummins 1978), being more or less napiform-
Laundon, G.F. 1975. Taxonomy and nomenclature notes on
ellipsoid in some species but oblong-ellipsoid, nearly Uredinales. Mycotaxon 3:133-161.
allantoid, or globoid in some. Any monograph of Léveillé, J.-H. 1847. Sur la disposition méthodique des Urédinées.
the genus doubtless will use these characteristics Ann. Sci. Nat. Bot. IIL, Ser. 8:369-376.
(Kaneko 1981). Most species are heteroecious, with Saho, H. 1968. [Studies on the needle rusts of five-needle pines.]
the spermogonia and aecia on needles of Pinus and Bull. Tokyo Univ. For. 64:59-148. (Japanese with English
summary.)
uredinia and telia on monocotyledonous and

64
 ManatieFh th ax
Risa ra ne

a
AYES ER

FIGURE 30. Coleosporium. A. A spermogonium of C. bletiae Dietel. B. A urediniospore of C. neocacaliae Saho. C. An

aeciospore of C. phellodendri Kom. D. A urediniospore of C. phellodendri. E. Teliospores of C. pini-asteris
Orishimo. F. Teliospores of C. pulsatillae F. Strauss. G. Teliospores of C. asterum (Dietel) P. & H. Syd.
H. Basidiospores of C. asterum. (Reprinted with permission, from Kaneko 1981.)
 CRONARTIUM Fr.
Obs. Mycol. 1:1-230 (220). 1815.

FAMILY: Cronartiaceae Didymopsora, and Endophylloides (Peterson 1973;

Hiratsuka 1995).
DESCRIPTION: Recently, it has become clear that a gall rust that
Spermogonia intracortical, under the periderm, has been known as C. quercuum (Berk.) Miyabe ex
Group II (type 9). Aecia intracortical, erumpent, most Shirai, occurring in North America (eastern gall
with strongly developed peridium, Peridermium- rust) and Asia (Japan, China, Korea and Russian Far
type; infections perennial and commonly causing East), comprises two distinct species. The North
hypertrophy of stems or cones; aeciospores American species, represented by the type (lecto-
catenulate, verrucose with rodlike columns, mostly type), retains the name C. quercuum and the one
having annulate sides. Uredinia subepidermal, open- occurring in Asia was named C. orientale S. Kaneko
ing by a pore, with dome-shaped peridium and (Kaneko 2000). Basidiospore shape varies between
sometimes also with intrasoral paraphyses, Milesia- the two species and among biological forms within
type; urediniospores borne singly, echinulate, germ the C. quercuum complex (Kaneko et al. 1991;
pores scattered or bizonate, obscure. Telia subepi- Kuhlman and Kaneko 1991).
dermal, often arising in uredinia, becoming erum- The economic importance of Cronartium species
pent as hairlike columns of strongly adherent spores is determined partly by the value of the pine host
embedded ina common matrix; teliospores 1-celled, and partly by the type of injury inflicted. In northern
catenulate, germ pores 1-3, obscure, wall thick, pale areas where white pines are important timber trees,
to brown; germination occurs without dormancy; white pine blister rust (caused by C. ribicola) is of
basidium external, basidiospores ellipsoid to importance; in the southern states of the USA the
globoid. fusiform gall rust of southern hard pines (caused by
C. quercuum f.sp. fusiforme) is the important species.
TYPE: Cronartium asclepiadeum Fr. There are authenticated cases of pine-to-pine
=Cronartium flaccidum (Alb. & Schwein.) spread in about eight species and these have been
G. Winter referred to Endocronartium.
On: Unidentified species of Asclepiadaceae
Type locality: Europe REFERENCES:
Burdsall, H.H., Jr.; Snow, G.A. 1977. Taxonomy of Cronartium
REMARKS: quercuum and C. fusiforme. Mycologia 69:503-508.
Thirty-six validly described Cronartium species Fries, E.M. 1815. Observationes mycologicae. Vol. 1. 230 p.
are listed by Peterson (1973), but 19 of them are now Henderson, D.M.; Hiratsuka, Y. 1974. Ontogeny of spore
considered to belong to other genera. The species of markings on aeciospores of Cronartium comandrae and
Cronartium are heteroecious, producing spermogo- peridermioid teliospores of Endocronartium harknessit. Can. J.
Bot. 52:1919-1921.
nia and aecia on the stems or cones of Pinus spp. and
Hiratsuka, Y. 1971. Spore surface morphology of pine stem rusts
the uredinia and telia on various dicotyledonous
of Canada as observed under a scanning electron microscope.
plants. Because the teliospores are not resting Can. J. Bot. 49:371-372.
spores, a persistent aecial state is to be expected and Hiratsuka, Y. 1995. Pine stem rusts of the world—frame work for
occurs either in stems or cones. The period between a monograph. Pages 1-8 in S. Kaneko, K. Katsuya, M.
infection of the pine and the production of aecia Kakishima, Y. Ono, eds. Proc. 4'. TUFRO Rusts of Pines Work.
varies from 2 to 3 to many years. When cones are Party Conf., Oct. 1994, Tsukuba, Jpn.
Hiratsuka, Y.; Powell, J.M. 1976. Pine stem rusts of Canada. Envi-
infected, as in C. conigenum Hedge. & Hunt and C.
ron. Can., Can. For. Serv., Ottawa, ON. For. Tech. Rep. 4. 103 p.
strobilinum Hedge. & G. Hahn, infections terminate
Kaneko, S. 2000. Cronartium orientale, sp. nov., segregation of the
with sporulation because the affected cone dies. In pine gall rust in eastern Asia from Cronartium quercuum.
species that produce aecia on main stems and Mycoscience 41:115-122.
branches, e.g. C. ribicola J.C. Fisch., C. quercuum (Berk.) Kaneko, S.; Kuhlman, E.G.; Powers, H.R., Jr. 1991. Morphological
Miyabe ex Shirai, and C. coleosporioides Arthur, the and physiological differences in the Cronartium quercuum com-
aecial cankers may sporulate for years. plex. Pages 69-75 in Y. Hiratsuka, ].K. Samoil, P.V. Blenis, P.E.
Crane, B.L. Laishley, eds. Rusts of Pine. Proc. IUFRO Rusts of
Many autoecious rusts included in Cronartium Pine Work. Party Conf., 18-22 Sept. 1989, Banff, AB, Canada.
because of their columnar telia are now considered For. Can., Northwest Reg., North. For. Cent., Edmonton, AB.
to belong to such genera as Cionothrix, Crossopsora, Inf. Rep. NOR-X-317.

66
Kuhlman, E.G.; Kaneko, S. 1991. Comparisons of basidiospores Peterson, R.S. 1973. Studies of Cronartium (Uredinales). Rep.
and urediniospores of formae speciales of Cronartium Tottori Mycol. Inst. Jpn. 10:203-223.
quercuum. Mycologia 83:440-445.
Peterson, R.S. 1967. The Peridermium species on pine stems. Bull.
Torrey Bot. Club 94:511-542.

FIGURE 31. Cronartium. A. A spermogonium of C. ribicola J.C. Fisch. B. An aecium of C. coleosporioides Arthur.
C. An aeciospore of C. coleosporioides. D. An aeciospore of C. comandrae Peck. E. A telial horn of C. ribicola.
F. Cross section of a telium of C. ribicola.

67
 ENDOCRONARTIUM Y. Hirats.
Can. J. Bot. 47:1493-1495 (1493). 1969.

FAMILY: Cronartiaceae undergo nuclear fusion and meiosis. Then haploid

nuclei migrate into straight germ tubes produced
DESCRIPTION: beyond the vesicles, often separated by septa (E.
Spermogonia intracortical, Group II (type 9). yamabense, E. sahoanum, and a part of E. pini) (Fig.
Telia intracortical, becoming erumpent, indetermi- 32B). Type I basidia are described and illustrated in
nate, with peridium as in the aecia of Cronartium; Hiratsuka (1968) and Hiratsuka et al. (1966); Type II
teliospores 1-celled, catenulate, verrucose, wall basidia are described and illustrated in Imazu et al.
colorless; basidia external. (1989, 1991). The existence of both types in E. piniin
the United Kingdom was suggested in Gibbs et al.
TYPE: Endocronartium harknessii (Moore) Y. Hirats. (1988). Basidiospores have not been observed.
=Peridermium harknessii Moore
On: Pinus ponderosa P. Laws. ex C. Laws. REFERENCES:
(Pinaceae) Christenson, J.A. 1968. A cytological comparison of germinating
Type locality: Colfax, California aeciospores in the Cronartium coleosporium complex.
Mycologia 60:1169-1177.
Epstein, L.; Buurlage, M.B. 1988. Nuclear division in germinating
REMARKS:
aeciospores and its taxonomic significance for the western
This is the endocyclic form of the genus gall rust fungus, Peridermium harknessit. Mycologia 80: 235—
Cronartium. Besides the type species, there are three 240.
other species, E. pini (Pers.) Y. Hirats., E. yamabense Gibbs, J.N.; England, N.; Wolstenholme, K. 1988. Variation in the
(Saho & I. Takah.) Paclt, and E. sahoanum Imazu & pine stem rust fungus Peridermium pini in the United King-
Kakish. (with two varieties, var. sahoanum and var. dom. Plant Pathol. 37:45-53.
hokkaidoense Imazu & Kakish.). There are also two Hiratsuka, Y. 1968. Morphology and cytology of aeciospores and
aeciospore germ tubes of host-alternating and pine-to-pine
races of P. filamentosum Peck (Inyo Peridermium and races of Cronartium flaccidum in Northern Europe. Can. J. Bot.
Powell Peridermium) in North America, as well as a 46:1119-1122.
form on Pinus armandii in central China, that likely Hiratsuka, Y. 1969. Endocronartium, a new genus for autoecious
belong to Endocronartium. They can be distin- pine stem rusts. Can. J. Bot. 47:1493-1495.
guished from Cronartium only by the type of spore Hiratsuka, Y.; Morf, W.; Powell, J.M. 1966. Cytology of the aecio-
spores and aeciospore germ tubes of Peridermium harknesstt
germination. Hiratsuka and his co-workers
and P. stalactiforme of the Cronartium coleosporioides complex.
(Hiratsuka et al. 1966; Hiratsuka 1969) showed that Can. J. Bot. 44:1639-1643.
spores of E. harknessti and E. pini produce a basidium Imazu, M.; Kakishima, M. 1992. A new variety of Endocronartium
in which nuclear fusion followed by meiosis occurs. sahoanum found on Pinus pumila in Hokkaido, Japan. Trans.
However, instead of basidiospores, short, Mycol. Soc. Jpn. 33:167-176.
uninucleate branches form which presumably func- Imazu, M.; Kakishima, M.; Kaneko, S. 1989. Endocronartium
sahoanum, a new stem rust fungus on Pinus pumila in Japan.
tion as infection pegs. Laundon (1976), Christenson
Trans. Mycol. Soc. Jpn. 30:301-310.
(1968), Epstein and Buurlage (1988), and Vogler et al.
Imazu, M.; Kakishima, M.; Katsuya, K. 1991. Morphology and
(1997) question whether there is proof that nuclear nuclear cycle of Endocronartium yamabense. Trans. Mycol. Soc.
fusion and meiosis occur. Two types of basidia are Jpnes2:371-379;
observed. In the first type (Type I), nuclear fusion Laundon, G.F. 1976. Peridermium (Fungi). Taxon 25:186-187.
occurs in the spores, meiosis occurs in the young Saho, H. 1981. Notes on Japanese rust fungi. VII. Peridermium
straight germ tubes, and the haploid nuclei are sepa- yamabense sp. nov., a pine-to-pine stem rust of white pines.
rated by septa; germ tubes are with or without side
Vogler, D.R.; Epstein, L.; Cobb, F.W., Jr. 1997. Nuclear behaviour
branches (E. harknessii and a part of E. pint) (Fig. and evolution of two populations of the western gall rust fun-
32A). In the second type (Type II), two dikaryon gus. Mycol. Res. 101:791-797.
nuclei migrate into elongated vesicles that are pro-
duced at the end of short germ tubes, and they

68
FIGURE 32. Endocronartium. Types of teliospore germination (schematic). A. Type I. B. Type II.

69
 BLASTOSPORA Dietel
Ann. Mycol. 6:222-229 (222-223). 1908.

FAMILY: Mikronegeriaceae reported that Caeoma deformans (B. betulae) is capable

of reinfecting the same host by aeciospores, but the
DESCRIPTION: cytological details are not known. Harada et al.
Spermogonia subepidermal, Group III (type 12). (1993) also reported that the aeciospores of B. betulae
Aecia Petersonia-type, deep seated; aeciospores are capable of repeating on the same host in addition
catenulate without intercalary cells. Uredinia to infecting Betula spp. to initiate the uredinial and
subepidermal, but suprastomatal when mature, telial states.
Uredo-type, the urediniospores produced from
sporogenous cells that emerge through stomata; REFERENCES:
urediniospores borne singly, echinulate, pores Dietel, P. 1908. Uredineen aus Japan. II. Ann. Mycol. 6:222-229.
obscure. Telia suprastomatal, as the uredinia; telio- Hama, T. 1987. [Studies on the important rust diseases of some
spores 1-celled, borne singly on pedicels, wall thin, conifers in the central mountainous region of Japan.] Bull. For.
For. Prod. Res. Inst. 343:1-118. (In Japanese.)
except apically thickened in B. betulae, pale, without
Harada, Y.; Hoshino, T.; Minowa, K.; Iwama, T. 1993. On life cycle
a clearly differentiated germ pore; germination of Caeoma deformans, the witches’ broom fungus on Thujopsis
occurs without dormancy by growth of the apical dolabrata var. hondae. Ann. Phytopathol. Soc. Jpn. 59:759.
region of the spore; basidia external. (Abstract.)
Hiratsuka, Y.; Hiratsuka, N. 1980. Morphology of spermogonia
TYPE: Blastospora smilacis Dietel and taxonomy of rust fungi. Rep. Tottori Mycol. Inst. 18:257—
On: Smilax sieboldii Miq. (Smilacaceae) 268.
Kakishima, M.; Imazu, M.; Kaneko, S. 1993. Life cycle connection
Type locality: Kochi, Japan
between Caeoma deformans and Blastospora betulae. Trans.
Mycol. Soc. Jpn. 34:187-193.
REMARKS: Kaneko, S.; Hiratsuka, N. 1981. Blastospora betulae (Uredinales)
The complete life cycles of all three known spe- from Japan. Mycologia 73:577-580.
cies, B. smilacis Dietel, B. betulae S. Kaneko & Hirats. Kaneko, S.; Hiratsuka, N. 1984. [Newly found rusts of birches and
f., and B. itoana Togashi & Onuma, have been eluci- fir.] J. Jpn. For. Soc. 66:280-284. (In Japanese.)
dated recently by Japanese researchers (Kakishima Mains, E.B. 1938. The genus Blastospora. Am. J. Bot. 25:677-679.
et al. 1993; Ono et al. 1986, 1987). All aecial states of Ono, Y.; Kakishima, M.; Kudo, A.; Sato, S. 1986. Blastospora
these species have been known as species of Caeoma, smilacis, a teleomorph of Caeoma makinoi, and its sorus devel-
opment. Mycologia 78:253-262.
C. makinoi Kusanao, C. deformans (Berk. & Broome)
Ono, Y.; Kakishima, M.; Sato, S.; Harada, Y. 1987. Blastospora
Tubeuf, and C. radiatum Shirai, respectively. Sper- itoana on Smilax nipponica forms a Caeoma stage on Prunus
mogonia of all three species are Group III (type 12). grayana in Japan. Mycologia 79:668-673.
Although some authors consider some of these as Togashi, K.; Onuma, F. 1931. A new species of Blastospora. Bot.
type 4, the well-developed ostioles and large overall Mag. Tokyo 45:4-7.
size are characteristics of type 12. Hama (1987)
 anes
IIL

FIGURE 33. Blastospora. A. A deep-seated Group III, type 12, spermogonium of B. betulae S. Kaneko & Hirats. f.
(from Hiratsuka and Hiratsuka 1980). B. A suprastomatal uredinium of B. smilacis Dietel (reprinted with
permission, from Ono et al. 1986). C. A telium of B. smilacis Dietel (reprinted with permission, from Ono
et al. 1986). D. An aeciospore of B. itoana Togashi & Onuma (reprinted with permission, from Ono et al.
1987). E. A suprastomatal uredinium of B. itoana (reprinted with permission, from Ono et al. 1987).
F. Teliospores and a basidium of B. smilacis (reprinted with permission, from Dietel 1908). G. Teliospores
of B. itoana.

Tes
 CHRYSOCELIS Lagerh. & Dietel
in Mayor, Mem. Soc. Neuchatel. Sci. Nat. 5:442-599 (542). 1913.

FAMILY: Mikronegeriaceae Chrysocelis Iupini Lagerh. & Dietel apparently has

Petersonia-type uredinia, at least some sori occur
DESCRIPTION: that are similar to the aecia but are not accompanied
Spermogonia subepidermal, deeply seated, by spermogonia. The genus differs from Chaconia
Group III (type 12) or Group V (type 4). Aecia sub- mainly because of type 12 or type 4 spermogonia.
epidermal, without peridium, Petersonia-type, Chrysocelis geophilicola (J.M. Yen) Cummins & Y.
opening by a porelike rupture of the host epidermis; Hirats. (Stomatisora geophilicola J.M. Yen) does not
aeciospores catenulate, verrucose. Uredinia subepi- differ significantly. Hence we treat Stomatisora as a
dermal, erumpent, Uredo-type; urediniospores synonym of Chrysocelis. This genus may be consid-
borne singly on pedicels, echinulate (or in C. lupini ered in or close to Mikronegeriaceae. Thirumalachar
like the aecia and aeciospores?), pores obscure. Telia (1960) considered Desmotelium Syd. to beasynonym
subepidermal or substomatal, becoming erumpent, of Chrysocelis, whereas we and Ono and Hennen
waxy when moist; teliospores 1-celled, laterally (1983) consider Desmotelium to be a synonym of
free, sessile, wall thin and pale, germ pore not obvi- Chaconia.
ously differentiated; germination occurs without
dormancy, by elongation of the apex of the spore; REFERENCES:
basidia semi-external. Lagerheim, G. von; Dietel, P. 1914. 99. Genere Chrysocelis Lagerh.
& Dietel spec. nov. Pages 542-544 in E. Mayor, ed. Contri-
TYPE: Chrysocelis lupini Lagerh. & Dietel bution a l'étude des Urédinées de Colombie. Mem. Soc.
Neuchatel. Sci. Nat. 5:442-599.
On: Lupinus sp. (Fabaceae)
Ono, Y. 1980. Two new rusts from the Satsunan Islands, Japan.
Type locality: Colombia Trans. Br. Mycol. Soc. 75:501-504.
Ono, Y.; Hennen, J.F. 1983. Taxonomy of the Chaconiaceous
REMARKS: genera (Uredinales). Trans. Mycol. Soc. Jpn. 24:369-402.
Seven species are known on unrelated host fami- Thirumalachar, M.J. 1960. Critical notes on some plant rusts. III.
lies such as Acanthaceae, Cucurbitaceae, Fabaceae, Mycologia 52:688-693.
Polygonaceae, and Zingiberaceae (Ono and Hennen
1983). Spermogonia are known for two of these.

FIGURE 34. Chrysocelis. A. Aeciospores and germinating teliospores of C. lupini Lagerh. & Dietel. B. Aeciospores and
germinating teliospores of C. gymnostemmatis Y. Ono (reprinted with permission, from Ono 1980).
 MIKRONEGERIA Dictel
in Dietel and Neger, Engler Bot. Jahrb. 27:1—-16 (16). 1899.

FAMILY: Mikronegeriaceae Argentina. Mikronegeria fagi produces telia on

Nothofagus, and aecia on Araucaria. Mikronegeria alba
DESCRIPTION: also produces telia on Nothofagus, but aecia on
Spermogonia deep-seated, usually lobed or Austrocedrus (Peterson 1968; Peterson and Oehrens
multiloculate, Group III (type 12). Aecia subepi- 1978). Deep-seated spermogonia of type 12 also
dermal, erumpent, Petersonia-type; aeciospores occur in Blastospora, and both genera have telio-
catenulate, verrucose. Uredinia subepidermal, spores that germinate without dormancy by contin-
erumpent, Uredo-type; urediniospores borne singly ued growth of the spore apex (semi-internal basidia)
on pedicels, echinulate, pores obscure or not differ- and Petersonia-type aecia. Peterson and Oehrens
entiated. Telia subepidermal, erumpent, waxy; (1978) speculated that Mikronegeria, together with
teliospores 1-celled, sessile, laterally free, without a Ceropsora, Olivea, Goplana, and Chrysocelis, with
differentiated germ pore; germination occurs with- poorly defined teliospores and obscure germ pores,
out dormancy by continued growth of the apex of may represent more primitive forms than the fir—
the spore; basidia semi-internal. fern rusts such as Uredinopsis and Hyalopsora, which
were suggested by earlier workers as the most prim-
TYPE: Mikronegeria fagi (Dietel & Neger) Dietel itive rusts.
=Melampsora fagi Dietel & Neger
On: Nothofagus procera (Poepp. & Endl.) Oerst. REFERENCES:
(Fagaceae) Butin, H. 1969. Studien ziir Morphologie und Biologie von
=N. nervosa (Phil.) Krasser Mikronegeria fagi Diet. & Neger, Phytopathol. Z. 64:242—257.
Type locality: Chile Dietel, P.; Neger, F. 1899. Uredinaceae chilenses. III. Engler Bot.
Jahrb. 27:1-16.
REMARKS: Peterson, R.S. 1968. Rust fungi on Araucariaceae. Mycopathol.
Mycol. Appl. 34:17-26.
Two species have been described, M. fagi (Dietel
Peterson, R.S.; Oehrens, E. 1978. Mikronegeria alba (Uredinales).
& Neger) Dietel and M. alba Oehrens & R.S. Peter- Mycologia 70:321-331.
son. Both species occur in the Andes in Chile and

FIGURE 35. Mikronegeria. A. An aeciospore of M. alba Oehrens & R.S. Peterson. B. Urediniospores of M. alba.
C. Germinating teliospores of M. fagi (Dietel & Neger) Dietel.

TS
 MELAMPSORA Castagne
Obs. Plantes Acotylédonées Fam. Urédiné 2:1—24 (18). 1843.

FAMILY: Melampsoraceae on Hyperium, supposedly is demicyclic, but lacks

spermogonia. The sori (possibly aecia) have peridial
DESCRIPTION: cells adherent to the upturned epidermis, verru-
Spermogonia Group I (type 3, subcuticular; or cose, catenulate spores, and intercalary cells. This
type 2, subepidermal). Aecia subepidermal, erum- species has, at times, been segregated as Mesopsora
pent, generally considered to be Caeoma-type, but hypericorum (G. Winter) Dietel.
some species have peridial cells adherent to the host Melampsora lini (Ehrenb.) Lév. is probably the
epidermis; aeciospores catenulate, verrucose with most important pathogen of a cultivated crop (flax).
rodlike columns or blocks. Uredinia subepidermal, In the intensive short-rotation cultivation of poplar
erumpent, Uredo-type, bright yellow or orange when and willow, several species of Melampsora may
fresh, fading to nearly colorless, with abundant cause sufficient damage to warrant the develop-
capitate paraphyses and in some species a partial ment of resistant cultivars and to consider control
peridium also; urediniospores borne singly on pedi- measures.
cels, wall colorless, echinulate, pores scattered or
bizonate, obscure. Telia subepidermal or rarely sub- REFERENCES:
cuticular, remaining covered, consisting of laterally No comprehensive monograph of this genus is
adherent crusts 1 spore deep; teliospores 1-celled, available other than vol. 3 of Sydow’s Monographia
sessile (some species have sporelike, presumably Uredinearum.
sterile, cells below the spores), wall brown or
Bagyanarayana, G. 1998. The species of Melampsora on Populus
brownish, germ pore 1; basidia external.
(Salicaceae). In R. Jalkanen, P. E. Crane, J.A. Walla, eds. Proc.
First IUFRO Rusts of Forest Trees Working Party Conf., 2-7
TYPE: Melampsora euphorbiae (C. Schub.) Castagne Aug. 1998, Saariselka, Finland. Finn. For. Res. Inst. Res. Pap.
=Xyloma euphorbiae C. Schub. 7237-01.
On: Euphorbia exigua L. (Euphorbiaceae) Castagne, L. 1843. Observations sur quelques plantes

Type locality: Germany acotylédonées de la famille des Urédinées et dans les sous-
tribus des Nemasporées et des Aecidinées, recueillies dans le
Dep. des Bouches-du-Rhone. II, Aix, 8. 24 p.; 2 tab.
REMARKS:
Hiratsuka, N.; Kaneko, S. 1982. A taxonomic revision of
Numerous species have been described, but not Melampsora on willows in Japan. Rep. Tottori Mycol. Inst.
all are distinct morphologically. Both heteroecious 20:1-32.
and autoecious species occur. Most heteroecious Kuprevich, V.E,; Tranzschel, V.G. 1957. Cryptogamic plants of the
species produce aecia on the needles of conifers and USSR. Vol. 4. Rust fungi. 1. Israel Prog. Sci. Transl., Jerusalem,
telia on Populus and Salix, but some produce the 1970. 518 p. (English translation.)
Pei, M.H.; Royle, DJ.; Hunter, T. 1995. A comparative study of
aecia on Allium, Saxifraga, or Ribes. Some species are
stem- and leaf-infecting forms of Melampsora rust on Salix
not obligatorily heteroecious and persist far from viminalis in UK. Mycol. Res. 99:357—-363.
the putative aecial hosts. The autoecious species Spiers, A.G.; Hopcroft, D.H. 1996. Morphological and host range
occur on various dicotyledonous families, including studies of Melampsora rusts attacking Salix species in New
Euphorbiaceae and Linaceae. Melampsora farlowti Zealand. Mycol. Res. 100:1163-1175.
(Arthur) Davis is microcyclic on Tsuga canadensis in Ziller, W.G. 1974. The tree rusts of western Canada. Environ.
Can., Can. For. Serv., Victoria, BC. Publ. 1329. 272 p.
North America. Melampsora hypericorum G. Winter,
 eS
come
naecere
cee
mos

FIGURE 36. Melampsora. A. A telium, urediniospores, and a paraphysis of M. occidentalis H.S. Jacks. B. A uredinial
paraphysis and a urediniospore of M. monticola Mains. C. Teliospores of M. lini (Pers.) Lév. (courtesy of
Dr. S. Kaneko). D. Teliospores of M. chelidonii-pierotii Matsumoto (courtesy of Dr. S. Kaneko).

75
 CEROTELIUM Arthur
Bull. Torrey Bot. Club 33:27-34 (30). 1906.

FAMILY: Phakopsoraceae a peridium, but uredinia of C. fici (E.J. Butler) Arthur

have peripheral paraphyses. Cerotelium fici is the
DESCRIPTION: most common species and is circumglobal in warm
Spermogonia subcuticular, Group VI (type 7). regions. Ono et al. (1992) suggested a taxonomic
Aecia subepidermal, erumpent, with peridium, affinity of C. tanakae to Aplopsora, but we retain it
Aecidium-type; aeciospores catenulate, verrucose. in Cerotelium. Buritica (1998) transferred C. tanakae
Uredinia subepidermal, slightly erumpent, with dome- S. Ito to Aplopsora (A. tanakae (S. Ito) Buritica &
shaped peridium, Milesia-type; with peripheral, J.E Hennen). Ono (1995) showed that the teliospores
basally united paraphyses, Malupa-type; or neither, of C. asari are formed successively from basal spo-
Calidion-type; urediniospores borne singly, rogenous cells as in Chaconia and also suggested the
echinulate, pores scattered, obscure. Telia subepi- close relationship of this species to Aplopsora,
dermal, becoming erumpent; teliospores catenulate assuming that the teliospores of Aplopsora are
in short chains, scarcely adherent laterally, wall thin formed sympodially from sporogenous basal cells.
and pale, germ pore obscure if differentiated; germi- Five species of Cerotelium on Fabaceae have been
nation occurs without dormancy; basidia external. well described and illustrated, with a key to species,
in Ono et al. (1992).
TYPE: Cerotelium canavaliae Arthur (as Cerateliumc.)
On: Canavalia ensiformis (L.) DC. (Fabaceae) REFERENCES:
Type locality: Puerto Rico Arthur, J. C. 1906. New species of Uredineae. Bull. Torrey Bot.
Club 33:27-34.
REMARKS: Buritica, P. 1998. La familia Phakopsoraceae en el neotr6pico. Il.
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22:325-334.
About 20 species are known from warmer re-
Buritica, P. 1999. La familia Phakopsoraceae (Uredinales) en el
gions on various host families. Aecia are known for
neotropico. IV. Géneros: Crossopsora, Cerotelium, Phragmidiella
C. dicentrae Mains & H.W. Anderson, which pro- y Catenulopsora. Rev. Acad. Colomb. Cienc. Exactas Fis. Nat.
duces aecia on Dicentra cucullaria (L.) Berth. and telia 23:407-431.
on Laportea canadensis Gaud.; and C. asari S. Kaneko Laundon, G.F. 1975. Taxonomy and nomenclature notes on
et al., which produces aecia on Corydalis lineariloba Uredinales. Mycotaxon 3:133-161.
Sieb. & Zucc. and uredinia/telia on Asarum Laundon, G.F; Rainbow, A. F. 1971. Cerotelium fici. CMI Descrip-
tions of pathogenic fungi and bacteria. 281:1-2.
caulascens Maxim (Ono 1995). The aecial state of
Ono, Y. 1995. Life cycle of Cerotelium asari (Uredinales). Sydowia
both species is systemic, and because teliospores 47:54-64.
germinate without dormancy, it is probable that a Ono, Y.; Buritica, P.; Hennen, J.F. 1992. Delimitation of Phakopsora,
persistent aecial state of some sort occurs in most or Physopella and Cerotelium and their species on Leguminosae.
all species. The uredinia of C. canavaliae Arthur have Mycol. Res. 96:825-850.

FIGURE 39. Cerotelium. A. A paraphysis and two urediniospores of C. fici (EJ. Butler)
Arthur. B. Two urediniospores and a telium of C. canavaliae Arthur.

78
 BATISTOPSORA Dianese, Medeiros & Santos
in J.C. Dianese, R.B. Medeiros, L.T.P. Santos, C. Furlanetto, M. Sanchez and A.C. Dianese
Fitopatol. Bras. 18:436—450 (437). 1993.

FAMILY: Phakopsoraceae known. This genus belongs in the family Phakop-

soraceae but differs from other genera in the family
DESCRIPTION: because of the position of the spermogonia and the
Spermogonia subepidermal, Group VI (type 5). presence of laterally fused paraphyses surrounding
Uredinia subepidermal becoming erumpent, para- the uredinia. However, these characteristics may
physes fused to each other to form a palisade around not be sufficient to separate this genus from
the uredinia (Uredostilbe-type); urediniospores Phakopsora.
echinulate with a truncate base, sessile. Telia
subepidermal with 3-4 layers of spores; teliospores REFERENCES:
formed in parallel columns, thin walled and Buritica, P. 1999. La familia Phakopsoraceae en el neotropico. III.
hyaline. Géneros: Batistopsora y Phakopsora. Rev. Acad. Colomb. Cienc.
Exactas Fis. Nat. 23(87):271-305.
Dianese, J.C.; Medeiros, R.B.; Santos, L.T.P.; Furlanetto, C.;
TYPE: Batistopsora crucis-filii Dianese, Medeiros
Sanchez, M.; Dianese, A.C. 1993. Batistopsora gen. nov. and
& Santos new Phakopsora, Ravenelia, Cerotelium, and Skierka species
On: Annona tomentosa R.E. Fries (Annonaceae) from the Brazilian Cerrado. Fitopatol. Bras. 18:436-450.
Type locality: Brazil

REMARKS:
The type and another species on Annona spp., B.
pistila Buritica & J.R. Hennen (Buritica 1999), are

FIGURE 38. Batistopsora. A uredinium (left) and a telium (right) of B. crucis-filii Dianese, Medeiros & Santos
(reprinted with permission, from Dianese et al. 1993).

77
 CEROTELIUM Arthur
Bull. Torrey Bot. Club 33:27-34 (30). 1906.

FAMILY: Phakopsoraceae a peridium, but uredinia of C.fici (E.J. Butler) Arthur

have peripheral paraphyses. Cerotelium fici is the
DESCRIPTION: most common species and is circumglobal in warm
Spermogonia subcuticular, Group VI (type 7). regions. Ono et al. (1992) suggested a taxonomic
Aecia subepidermal, erumpent, with peridium, affinity of C. tanakae to Aplopsora, but we retain it
Aecidium-type; aeciospores catenulate, verrucose. in Cerotelium. Buritica (1998) transferred C. tanakae
Uredinia subepidermal, slightly erumpent, with dome- S. Ito to Aplopsora (A. tanakae (S. Ito) Buritica &
shaped peridium, Milesia-type; with peripheral, J.F Hennen). Ono (1995) showed that the teliospores
basally united paraphyses, Malupa-type; or neither, of C. asari are formed successively from basal spo-
Calidion-type; urediniospores borne singly, rogenous cells as in Chaconia and also suggested the
echinulate, pores scattered, obscure. Telia subepi- close relationship of this species to Aplopsora,
dermal, becoming erumpent; teliospores catenulate assuming that the teliospores of Aplopsora are
in short chains, scarcely adherent laterally, wall thin formed sympodially from sporogenous basal cells.
and pale, germ pore obscure if differentiated; germi- Five species of Cerotelium on Fabaceae have been
nation occurs without dormancy; basidia external. well described and illustrated, with a key to species,
in Ono et al. (1992).
TYPE: Cerotelium canavaliae Arthur (as Ceratelium c.)
On: Canavalia ensiformis (L.) DC. (Fabaceae) REFERENCES:
Type locality: Puerto Rico Arthur, J. C. 1906. New species of Uredineae. Bull. Torrey Bot.
Club 33:27-34.
REMARKS: Buritica, P. 1998. La familia Phakopsoraceae en el neotropico. II.
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22:325-334.
About 20 species are known from warmer re-
Buritica, P. 1999. La familia Phakopsoraceae (Uredinales) en el
gions on various host families. Aecia are known for neotropico. IV. Géneros: Crossopsora, Cerotelium, Phragmidiella
C. dicentrae Mains & H.W. Anderson, which pro- y Catenulopsora. Rev. Acad. Colomb. Cienc. Exactas Fis. Nat.
duces aecia on Dicentra cucullaria (L.) Berth. and telia 23:407-431.
on Laportea canadensis Gaud.; and C. asari S. Kaneko Laundon, G.F. 1975. Taxonomy and nomenclature notes on
et al., which produces aecia on Corydalis lineariloba Uredinales. Mycotaxon 3:133-161.
Sieb. & Zucc. and uredinia/telia on Asarum Laundon, G.F.; Rainbow, A. F. 1971. Cerotelium fici. CMI Descrip-
tions of pathogenic fungi and bacteria. 281:1—2.
caulascens Maxim (Ono 1995). The aecial state of
Ono, Y. 1995. Life cycle of Cerotelium asari (Uredinales). Sydowia
both species is systemic, and because teliospores 47:54-64.
germinate without dormancy, it is probable that a Ono, Y.; Buritica, P.; Hennen, J.F. 1992. Delimitation of Phakopsora,
persistent aecial state of some sort occurs in most or Physopella and Cerotelium and their species on Leguminosae.
all species. The uredinia of C. canavaliae Arthur have Mycol. Res. 96:825-850.

FIGURE 39. Cerotelium. A. A paraphysis and two urediniospores of C. fic! (E.J. Butler)
Arthur. B. Two urediniospores and a telium of C. canavaliae Arthur.

78
 CROSSOPSORA H. & P. Syd.
Ann. Mycol. 16:240-248 (243). 1918.

FAMILY: Phakopsoraceae spermogonia are different, the aecia of Crossopsora

are not Peridermium-type, the uredinia have pa-
DESCRIPTION: raphyses rather thana peridium, and the teliospores
Spermogonia subcuticular, Group VI (type 7). are not embedded in a common matrix. Mundkur
Aecia subepidermal, opening by a porelike rupture and Thirumalachar (1946) describe and illustrate
of the epidermis, without a peridium, Caeoma-type; suprastomatal uredinia in C. zizyphi (Syd. & EJ.
aeciospores catenulate, echinulate. Uredinia subepi- Butler) H. & P. Syd., but whether most species have
dermal, becoming erumpent, usually with septate such sori is doubtful.
and basally united paraphyses, Malupa-type; No general statement can be made about life
urediniospores borne singly, echinulate, pores scat- histories because spermogonia and aecia have been
tered, obscure. Telia subepidermal, erumpent as described only in C. sawadae (H. & P. Syd.) Arthur &
hairlike columns of 1-celled, catenulate, strongly ad- Cummins. All species are probably autoecious.
herent teliospores, germ pores not observed; germi-
nation occurs without dormancy; basidia external. REFERENCES:
Arthur, J.C.; Cummins, G.B. 1936. Philippine rusts in the Clemens
TYPE: Crossopsora zizyphi (Syd. & E.J. Butler) H. & Collection 1923-1926. II. Philipp. J. Sci. 61:463-488.
Pasyd. Dianese, J.C.; Buritica, P.; Hennen, J.F. 1994. The rust of “jatoba”:
=Cronartium zizyphi Syd. & E.J. Butler a new Crossopsora species from Neotropica on Hymenaea
(Leguminosae). Fitopatol. Bras. 19:588-591.
On: Ziziphus oenopolia Mill. (Rhamnaceae)
Peterson, R.S. 1973. Studies of Cronartium (Uredinales). Rep.
Type locality: India Tottori Mycol. Inst. 10:203-223.
Mundkur, B.B.; Thirumalachar, M.J. 1946. Revisions of and addi-
REMARKS: tions to Indian Fungi. I. Commonw. Mycol. Inst. Mycol. Pap.
The genus has not been monographed, but there 16:1-27.
are 12 to 15 species and all inhabit warm regions. Sydow, H.; Sydow, P. 1918. Mykologische Mitteilungen. Ann.
The telia superficially resemble those of Cronartium, Mycol. 16:240-248.
but it is doubtful that the two genera are related. The

=>

FIGURE 40. Crossopsora. A. Cross section of a telium of C. zizyphi (Syd. & E.J. Butler) H. & P. Syd. (courtesy of Dr. R.
Berndt). B. A telium of C. malloti (Racib.) Cummins.

72
 DASTURELLA Mundkur & Khesw.
Mycologia 35:201—206 (202-203). 1943.

FAMILY: Phakopsoraceae REMARKS:

Only two or three species are known. The type
DESCRIPTION: species is heteroecious with aecia on Randia
Spermogonia subcuticular, Group VI (type 7). (Rubiaceae) and the telia on Dendrocalamus
Aecia subepidermal, erumpent, with peridium, (Poaceae). Dasturella differs from Physopella in the
Aecidium-type; aeciospores catenulate, verrucose. strongly erumpent telia and from Uredopeltis in the
Uredinia subepidermal, erumpent, with abundant, catenulate teliospores.
incurved, peripheral paraphyses, Calidion-type;
urediniospores borne singly, echinulate, pores REFERENCE:
equatorial. Telia subepidermal, becoming erumpent Mundkur, B.B.; Kheswalla, K.F. 1943. Dasturella—a new genus of
in cushionlike crusts several spores deep; telio- Uredinales. Mycologia 35:201—206.
spores 1-celled, firmly adherent laterally and termi-
nally, catenulate, brown, germ pore not seen;
basidium external.

TYPE: Dasturella divina (Syd.) Mundkur & Khesw.

=Angiopsora divina Syd.
On: Dendrocalamus strictus Nees (Poaceae)
Type locality: India

FIGURE 41. Dasturella. D. divina (Syd.) Mundkur & Khesw. A. One paraphysis and one urediniospore. B. Teliospore
chains. C. A telium.

80
 KWEILINGIA Teng
Sinensia 11:105-130 (124). 1940.

FAMILY: Phakopsoraceae & P.C. Pandey) Buritica and K. divina (Syd.) Buritica)
(Buritica 1998). Kweilingia bagchii occurs on Dendro-
DESCRIPTION: calamus strictus Nees (Bambusoideae-Poaceae) in
Spermogonia and aecia unknown. Uredinia sub- India. Another species, K. americana Buritica & J.F.
epidermal, erumpent, with peripheral paraphyses, Hennen, is described on Costus pictus Don
Calidion-type; urediniospores borne singly, echinu- (Costaceae) from Mexico (Buritica 1998).
late, pores obscure. Telia subepidermal, erumpent, Thirumalachar and Narasimhan (1951) concluded
large, blackish, discoid-pulvinate, flabelliform in that the fungus is not a rust but a member of the
longitudinal section or spreading over the leaf and Auriculariales, based on a study of the type of
sheath as a mantle; teliospores 1-celled, catenulate, Kweilingia. However, uredinia and urediniospores
the spores or chains loosely adherent at first but occur in addition to telia, suggesting affinity with
separating later, wall brown, germ pores 2, when the rusts. Further investigation will be needed to
visible, apical or nearly in each cell; basidium doubt- decide the proper classification of Kweilingia.
less external.
REFERENCES:
TYPE: Kweilingia bambusae (Teng) Teng Buritica, P. 1998. La familia Phakopsoraceae en el neotropico. II.
=Chrysomyxa bambusae Teng Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22:325-334.

On: Bambusa sp. (Poaceae) Mundkur, B.B.; Kheswalla, K.F. 1943. Dasturella—a new genus of
Uredinales. Mycologia 35:201-206.
Type locality: Kwangsi, China
Singh, S.; Pandy, P.C. 1971. Tunicopsora, a new rust genus on
bamboo. Trans. Br. Mycol. Soc. 56:301-318.
REMARKS:
Teng, S.C. 1940. Supplement to higher fungi in China. Sinensia
Tunicopsora and Dasturella are considered syn- 11:105-130.
onyms of Kweilingia. Type species of these genera Thirumalachar, M.J.; Narasimhan, M.J. 1951. Critical notes on
were transferred to Kweilingia (K. bagchti (Suj. Singh some plant rusts. III. Sydowia 5:476-483.

FIGURE 42. Kweilingia. A. A telium of K. divina (Syd.) Buritica (reprinted with permission, from Mundkur and
Kheswalla 1943). B. A paraphysis and a urediniospore of K. divina. C. Teliospores of K. bambusae (Teng)
Teng. D. A telium of K. americana Buritica & J.F. Hennen (courtesy of Dr. J.F. Hennen.)

81
 MEHTAMYCES Mundkur & Thirum.
Mycologia 37:619-628 (620-621). 1945.

FAMILY: Phakopsoraceae REMARKS:

Thirumalachar and Mundkur (1949) considered
DESCRIPTION: Mehtamyces to be a synonym of Phragmidiella, but
Spermogonia and aecia unknown. Uredinia sub- Ramachar and Rao (1981) supported the recognition
epidermal, aparaphysate, Uredo-type; uredinio- of this genus, based on the morphological character-
spores borne in clusters on sporogenous basal cells, istics of Mehtamyces, Phragmidiella, and Physopella.
wall bilaminate. Telia subepidermal crusts, non- We follow their view, especially recognizing the lat-
erumpent; teliospores develop in chains, laterally erally united teliospores and nonerumpent telia of
united, forming a crust, basipetally produced; ger- Mehtamyces.
mination unknown.
REFERENCES:
TYPE: Mehtamyces stereospermi (Mundkur) Mundkur, B.B.; Thirumalachar, M.J. 1945. Two new genera of
Mundkur & Thirum. rusts on Bignoniaceae. Mycologia 37:619-628.
=Phakopsora stereospermi Mundkur Ramachar, P.; Rao, A.S. 1981. Mehtamyces (Uredinales) reinstated.
Mycologia 73:778-780.
On: Stereospermum suaveolens Wall.
Thirumalachar, M.J.; Mundkur, B.B. 1949. Genera of rusts. II.
(Bignoniaceae) Indian Phytopathol. 2:239-241.
Type locality: Nagapur, India

FIGURE 43. Mehtamyces. Teliospores and urediniospores of

M. stereospermi (Mundkur) Mundkur & Thirum.

82
 NOTHORAVENELIA Dietel
Ann. Mycol. 8:304-314 (310). 1910.

FAMILY: Phakopsoraceae REMARKS:

There are two species, the type on Euphor-
DESCRIPTION: biaceae, and N. commiphorae Cummins (Cummins
Spermogonia subcuticular, Group VI (type 7). 1952) on Burseraceae. The genus is superficially
Aecia Caeoma-type; aeciospores catenulate and similar to Ravenelia and Kernkampella, but it is
verrucose. Uredinia subepidermal, erumpent, with probably related to the phakopsoroid genera.
peripheral paraphyses, Calidion-type; uredinio-
spores echinulate, pores obscure. Telia subepi- REFERENCES:
dermal, erumpent, with basally united, peripheral Cummins, G.B. 1952. Uredinales from various regions. Bull.
paraphyses; teliospores laterally and terminally Torrey Bot. Club 79:212-234.

strongly adherent, forming discoid heads, the heads Dietel, P. 1910. Uredineen aus Japan. II]. Ann. Mycol. 8:304-314.
Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
1-3 spores thick, each chain of spores subtended by
Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
a cystlike cell, the cystoid cells remaining attached Nakayama, K. 1992. The rust flora of Japan. Tsukuba
to the spore heads but separating from the sorus; Shuppankai, Ibaraki. 1205 + 159 p.
successive heads may form and force the older ones
from the sorus; spore wall pigmented, cystoid cells
hyaline, germ pores not seen; germination
unknown, but basidia undoubtedly external.

TYPE: Nothoravenelia japonica Dietel

On: Securinega suffruticosa Rehd. (Euphorbiaceae)
=S. ramiflora Muell.
Type locality: Kochi-ken, Japan

FIGURE 44. Nothoravenelia. A teliospore head with paraphyses

and a urediniospore of N. commiphorae Cummins.

83
 PHAKOPSORA Dietel
Ber. Dtsch. Bot. Ges. 13:332—335 (333-334). 1895.

FAMILY: Phakopsoraceae treat Physopella as wellas Angiopsora as synonyms of

Phakopsora. However, we do not support the use of
DESCRIPTION: Physopella as anamorph genus name (Ono et al.
Spermogonia subcuticular, Group VI (type 7). 1992). Although a mistake was made in choosing the
Aecia subepidermal, erumpent, Calidion-type or type specimen as pointed out by Cummins and
Milesia-type; aeciospores borne singly and similar to Ramachar (1958), the mistake should be considered
the urediniospores. Uredinia subepidermal, erum- as a “formal error”, and Physopella should be recog-
pent, with peripheral, incurved, unusually dorsally nized as a holomorph name. The species that are
thick-walled paraphyses surmounting peridial often segregated into Bubakia are autoecious. Sper-
tissue, Malupa- and Calidion-type (Phakopsora sensu mogonia and aecia are not known for Phakopsora
stricto), or without paraphyses, Uredo-type (Bubakia sensu stricto, and it is possible that some or all of
when segregated); urediniospores borne singly, them are heteroecious.
wall echinulate, brownish or nearly colorless, pores Phakopsora pachyrhizi H. & P. Syd. (=Physopella
scattered or equatorial. Telia subepidermal, not pachyrhizae (H. & P. Syd.) Azbukina) on soybean,
erumpent, consisting of crusts of laterally adherent and P. gossypii (Lagerh.) Hirats. f. on cotton, are eco-
teliospores 2 or more cells deep; teliospores sessile, nomically important.
either catenulate or irregularly arranged, 1-celled,
the wall usually brown or brownish, with 1 apical REFERENCES:
germ pore; presumably germinating after dor- Azbukina, Z.M. 1970. O sistematicheskom polozhenii i
mancy in most or all species; basidia external. geneticheskikh rodov triby Phakopsoreae Arth. emend. Azb.
Nov. Sist. Nizshikh Rast. 7:208-232.
TYPE: Phakopsora punctiformis (Barclay & Dietel) Buritica, P. 1999. La familia Phakopsoraceae en el Neotrépico. III.
Generos: Batistopsora y Phakopsora. Rev. Acad. Colomb. Cienc.
Dietel Exactas Fis. Nat. 23:271-305.
=Melampsora punctiformis Barclay & Dietel Buritica, P. 1999. Familia Phakopsoraceae (Fungi: Uredinales)
On: Galium aparine L. (Rubiaceae) generalidades afinidades. Rev. Fac. Nac. Agron. Medellin
Type locality: India 52:467-505.
Cummins, G.B.; Ramachar, P. 1958. The genus Physopella
REMARKS: (Uredinales) replaces Angiopsora. Mycologia 42:741—744.
More than 90 species have been named, occur- Dietel, P. 1895. Drei neue Uredineen Gattungen, Maseeélla,

ring on over 150 genera and 30 families of monocots Phakopsora u. Schizospora. Ber. Dtsch. Bot. Ges. 13:332-335.
Ono, Y. 2000. Taxonomy of the Phakopsora ampelopsidis species
and dicots (Ono et al. 1992). Most are distributed in
complex on vitaceous hosts in Asia including a new species, P.
warm regions. Numerous Uredo species also may euvitis. Mycologia 92:154-173.
belong here. Phakopsora and Physopella have been Ono, Y.; Buritica, P.; Hennen, J.F. 1992. Delimitation of Phakopsora,
distinguished because the teliospores of Phakopsora Physopella and Cerotelium and their species on Leguminosae.
are irregularly arranged, whereas those of Physo- Mycol. Res. 96:825-850.
pella are in chains. However, we do not consider this Thirumalachar, M.J.; Kern, FD. 1949. Notes on some species of
Phakopsora and Angiopsora. Mycologia 41:283-290.
difference adequate to separate the two genera and

84
 Sead
Pare Ng >

FIGURE 45. Phakopsora. A. A uredinium of P. ampelopsidis Dietel & Syd. ex Dietel (reprinted with permission, from
Ono 2000). B. A urediniospore of P. ampelopsidis. C. Urediniospores of P. mexicana (Cummins) Cummins &
Y. Hirats. comb. nov. (@Physopella mexicana Cummins). D. A paraphysis and urediniospores of
P. compressa (Mains) Cummins & Y. Hirats. comb. nov. (=Angiopsora compressa Mains). E. Teliospores
of P. compressa. F. Teliospores of P. pachyrhizi H. & P. Syd. G. A paraphysis and urediniospores of
P. pachyrhizi.

85
 PHRAGMIDIELLA Henn.
Engler, Bot. Jahrb. 38:102-118 (104). 1905.

FAMILY: Phakopsoraceae plants of the Bignoniaceae or Anacardiaceae. Inso-

far as the life cycles are known, the species are
DESCRIPTION: autoecious and macrocyclic. The genus has telia
Spermogonia subcuticular, Group VI (type 7). similar to Cerotelium; both genera have type 7 sper-
Aecia subcuticular, erumpent, Uredo-type; aecio- mogonia; Cerofelium has Aecidium-type aecia, but in
spores borne singly, echinulate. Uredinia subepi- Phragmidiella the aecia are Uredo-type; and the
dermal, erumpent, Uredo-type; urediniospores urediniospores are quite different. It is doubtful that
borne singly, echinulate, pores equatorial. Telia sub- the two genera are closely related. Mehtamyces
epidermal, mostly erumpent, composed of crowded Mundkur & Thirum. (M. stereospermi (Mundkur)
but not adherent chains of 1-celled teliospores, germ Mundkur & Thirum., type) is treated here sepa-
pore 1, apical and indistinct, wall pale; germination rately, as suggested by Ramachar and Rao (1981),
occurs without dormancy; basidia external. who have proposed reinstatement of Mehtamyces.

TYPE: Phragmidiella markhamiae Henn. REFERENCES:

On: Markhamia sansibarensis Schumann Buritica, P.-E. 1999. La familia Phakopsoraceae (Uredinales) en el
(Bignoniaceae) Neotropico. IV. Géneros: Crossopsora, Cerotelium, Phragmidiella
Type locality: Tanzania, Africa y Catenulopsora. Rev. Acad. Colomb. Cienc. Exactas Fis. Nat.
23:407-431.
Hennings, P. 1905. Fungi Africae orientalis. IV. Engler, Bot. Jahrb.
REMARKS: 38:102-118.
Five species are known from the Caribbean, Af- Ramachar, P.; Rao, A.S. 1981. Mehtamyces (Uredinales) reinstated.
rica, Brazil, and India (Buritica 1999). They parasitize Mycologia 73:778-780.

FIGURE 46. Phragmidiella. A telium of P. markhamiae Henn.

86
 PUCCINIOSTELE Tranzschel & Kom.
in Komarov, Arb. Naturforsch. Ges. St. Petersb. 30:135-140 (138). 1899.

FAMILY: Phakopsoraceae All four known species parasitize Astilbe of the

Saxifragaceae. Pucciniostele hashiokai (Hirats. f.)
DESCRIPTION: Cummins on Ampelopsis (Vitaceae) has been trans-
Spermogonia subcuticular, Group VI (type 7). ferred to a new genus, Scalarispora (Buritica and
Aecia subepidermal, erumpent, Caeoma-type, with- Hennen 1994).
out peridium; aeciospores catenulate, verrucose. Dietel (1904) pointed out that Komarov and
Uredinia subepidermal, erumpent, with peridium Tranzschel (1899) mistook the specimen they had for
and catenulate spores, Aecidium-type, or with a separate species from India (Xenodochus clarkiana
peripheral paraphyses and spores borne singly on Barclay). Dietel renamed a Manchurian (Jilin,
pedicels, Calidion-type. Telia subepidermal, only China) species that Komarov and Tranzschel
slowly exposed; teliospores catenulate, of two (Komarov 1899) had designated as the type as P.
kinds: (1) the primary spores developing in aecia mandschurica Dietel and retained the Indian species
or closely associated with aecia and consisting of as P. clarkiana (Barclay) Dietel (non Tranzschel &
catenulate, 4-celled, tetradlike disks, and (2) second- Kom.).
ary spores developing in separate sori, 1-celled,
catenulate, germ pores not seen; germination un- REFERENCES:
known but basidia doubtless external. Buritica, P.; Hennen, J.F. 1994. Familia Phakopsoraceae
(Uredinales) 1. Géneros anamorficos y teliomérficos. Rev.
TYPE: Pucciniostele clarkiana (Barclay) Tranzschel Acad. Colomb. Cienc. Exactas Fis. Nat. 19:47-62.
Cummins, G.B. 1950. Uredinales of continental China collected
& Kom.
by S.Y. Cheo I. Mycologia 42:779-797.
=Xenodochus clarkiana Barclay
Cummins, G.B.; Thirumalachar, M.J. 1953. Pucciniostele a genus of
=Pucciniostele mandschurica Dietel rust fungi. Mycologia 45:572-578.
On: Astilbe chinensis Franchet & Savatier Dietel, P. 1904. Uber die Uredineengattung Pucciniostele
(Saxifragaceae) Tranzschel et Komarov. Ann. Mycol. 2:20-26.
Type locality: Jilin, China Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
REMARKS: Nakayama, K. 1992. The rust flora of Japan. Tsukuba
Shuppankai, Tsukuba. 1205 + 159 p.
Pucciniostele is a poorly understood genus and
Komaroy, V.L. 1899. [A new genus of rust fungi—Pucciniostele
anomalous in that it apparently produces two kinds Tranzschel & Komar.] Arb. Naturforsch. Ges. St. Petersb.
of teliospores, but germination has not been studied 30:135-140. (In Russian.)
in either kind (Cummins and Thirumalachar 1953).

FIGURE 47. Pucciniostele. P. mandschurica Dietel.

A. Tetradlike teliospores. B. Catenulate
teliospores.

87
 SCALARISPORA Buritica & J.F. Hennen
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 19:47-62 (61). 1994.

FAMILY: Phakopsoraceae pedicels. Hiratsuka (1937) originally described this

species in Cerotelium, but Hiratsuka et al. (1992)
DESCRIPTION: recognized it in Pucciniostele, as did Cummins (1950)
Spermogonia and aecia unknown. Uredinia and Cummins and Thirumalachar (1953), with some
minute, with club-shaped peripheral paraphyses, reservation. Buritica and Hennen (1994) established
Calidion-type; urediniospores pedicellate and a new genus for this species.
echinulate. Telia erumpent, powdery; teliospores
pointed at both ends, consisting of 5 to 8 cells pro- REFERENCES:
duced in chains. Buritica, P.; Hennen, J.F. 1994. Family Phakopsoraceae
(Uredinales). 1. Géneros anamorficos y teliomorficos. Rev.
TYPE: Scalarispora hashiokai (Hirats. f.) Buritica & Acad. Colomb. Cienc. Exactas Fis. Nat. 19:47-62.
Cummins, G.B. 1950. Uredinales of continental China collected
J.F. Hennen
by S.Y. Cheo. I. Mycologia 42:779-797.
=Cerotelium hashiokai Hirats. f.
Cummins, G.B.; Thirumalachar, M.J. 1953. Pucciniostele agenus of
=Pucciniostele hashiokai (Hirats. f.) rust fungi. Mycologia 45:572-578.
Cummins Hiratsuka, N. 1937. Miscellaneous notes on the East-Asiatic
On: Ampelopsis cantoniensis Planch. (Vitaceae) Uredinales with special reference to the Japanese species. I. J.
Type locality: Taiwan Jpn. Bot.13:422-251.
Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
REMARKS: Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
Nakayama, K. 1992. The rust flora of Japan. Tsukuba
Only the type species is known. The teliospores Shuppankai, Tsukuba. 1205 + 159 p.
resemble those of Phragmidium but without

FIGURE 48. Scalarispora. Teliospores of S. hashiokai (Hirats. f.)

Buritica & J.F. Hennen.

88
 UREDOPELTIS Henn.
Ann. Mus. Congo Belge (Ser. 5) 2:223. 1908.

FAMILY: Phakopsoraceae telia. Laundon (1963) studied the type specimen

from the Congo and specimens from Northern
DESCRIPTION: Rhodesia. Besides the type species, two others, U.
Spermogonia and aecia unknown. Uredinia sub- guettardae Buritica & J.F. Hennen on Guettarda sp.
epidermal, becoming erumpent, small, with (Rubiaceae) from Brazil and U. dominicana (F. Kern)
incurved, basally united, peripheral paraphyses, Buritica & J.F. Hennen on Croton spp. (Euphor-
Calidion-type; urediniospores borne singly, biaceae) from Mexico and the Dominican Republic,
echinulate, pores scattered. Telia subepidermal, are known (Buritica 1998; Buritica and Hennen 1994).
becoming strongly erumpent, consisting of cush-
ions of laterally adherent spores several spores REFERENCES:
deep; teliospores sessile and irregularly arranged, Buritica, P. 1998. La familia Phakopsoraceae en el neotropico. II.
not catenulate, 1-celled, germ pore probably 1; Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22:325-334.
basidia not seen but doubtless external. Buritica, P.; Hennen, J.F. 1994. Familia Phakopsoraceae
(Uredinales). 1. Géneros anamorficos y teliomorficos. Rev.
Acad. Colomb. Cienc. Exactas Fis. Nat. 19:47-62.
TYPE: Uredopeltis congensis Henn.
Hennings, P. 1908. Fungi. Ann. Mus. Congo Belge (Ser. 5) 2:223.
On: Markhamia sp. (Bignoniaceae) Laundon, G.F. 1963. Uredopeltis (Uredinales). Trans. Br. Mycol.
Type locality: Congo Soc. 46:503-504.

REMARKS:
Uredopeltis appears to be similar to Phakopsora
and Physopella, but the latter have nonerumpent

FIGURE 49. Uredopeltis. A telium and a urediniospore of

U. congensis Henn.

89
 ACHROTELIUM Syd.
in Sydow and Petrak, Ann. Mycol. 26:414—446 (425). 1928.

FAMILY: Chaconiaceae Periplocaceae (one species), Sapotaceae (one

species), Thymeleaceae (one species) and
DESCRIPTION: Urticaceae (one species). They occur in Africa, India,
Spermogonia Group VI (type 7). Aecia Philippines, West Indies, and southern United
subepidermal, erumpent, Uredo-type, similar to States (Ono 1978). Together with Botryorhiza and
uredinia; aeciospores borne singly, echinulate. Maravalia, Ono and Hennen (1983) hesitated to put
Uredinia subepidermal, erumpent, Uredo-type; Achrotelium in the family Chaconiaceae because of
urediniospores borne singly on pedicels, echinulate, the pedicellate nature of the teliospores, but we
pores obscure or scattered. Telia subepidermal, retain it in Chaconiaceae.
erumpent; teliospores produced in groups on basal
cells, pedicellate, wall pale, spores 1-celled at first, REFERENCES:
becoming 4-celled by division of the protoplast (in- Arthur, J.C.; Cummins, G.B. 1936. Philippine rusts in the Clemens
ternal basidia), each cell producing 1 basidiospore; collection 1923-26. II. Phillipp. J. Sci. 61:463-488; plates 1+.
teliospore germination occurs without dormancy. Ono, Y. 1978. Taxonomic revisions of the Tribe Oliveae and
morphologically related genera (Uredinales). Ph.D. thesis,
Purdue Univ., Lafayette, IN.
TYPE: Achrotelium ichnocarpi Syd.
Ono, Y.; Hennen, J.F. 1983. Taxonomy of the Chaconiaceous
On: Ichnocarpus volubilis Merr. (Apocynaceae) genera (Uredinales). Trans. Mycol. Soc. Jpn. 24:369-402.
Type locality: Philippines Sydow, H.; Petrak, F. 1928. Micromycetae philippinenses. Ann.
Mycol. 29:414-446.
REMARKS:
At least seven species are known on Apocy-
naceae (two species), Asclepiadaceae (one species),

FIGURE 50. Achrotelium. A. ichnocarpi Syd. A. Teliospores with internal

basidia. B. Urediniospores.

90
 APLOPSORA Mains
Am. J. Bot. 8:442-451 (442). 1921.

FAMILY: Chaconiaceae species are A. lonicerae Tranzschel and A. corni Y.

Ono & Harada on Cornus controversa Hemsley
DESCRIPTION: (Cornaceae) (Ono and Harada 1994) and A. hennenti
Spermogonia and aecia unknown. Uredinia sub- Dianese & Santos on Qualea multiflora Mart.
epidermal, only slightly erumpent, Malupa-type, (Vochysiaceae). Buritica (1998) placed Cerotelium
with peripheral paraphyses, united basally; tanakae S. Ito in Aplopsora (A. tanakae (S. Ito) Buritica
urediniospores borne singly on pedicels, pores ob- & J.F. Hennen) (see Cerotelium).
scure. Telia subepidermal but soon exposed, con- Aplopsora has uredinia similar to those of
sisting of a single layer of crowded, 1-celled, sessile, Phakopsora, but the teliospores are borne singly.
thin-walled, pale teliospores; germ pore obscure if Although the teliospores resemble those of Melamp-
differentiated; germination occurs without dor- sora the paraphyses are unlike those of Melampsora.
mancy; basidia external. Ochropsora, also in Chaconiaceae, has internal
basidia.
TYPE: Aplopsora nyssae Mains
On: Nyssa aquatica L. (Nyssaceae) REFERENCES:
Type locality: Mississippi, USA Buritica, P. 1998. La familia Phakopsoraceae en el neotropico. II.
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22:325-334.
Dianese, J.C.; Santos, L.T.P. 1995. Aplopsora hennenii sp. nov., the
REMARKS:
first fungus recorded from Vochysiaceae. Mycol. Res. 98:914—
Because the teliospores germinate without 916.
dormancy and the hosts are deciduous, it is Mains, E.B. 1921. Unusual rusts on Nyssa and Urticastrum. Am. J.
probable that the aecial state has perennial or at least Bot. 8:442-451.
overwintering mycelium. The species are probably Ono, Y.; Harada, Y. 1994. Aplopsora corni sp. nov. on Cornus
heteroecious. In addition to the type, other known controversa from Hokkaido, Japan. Mycoscience 35:179-181.

FIGURE 51. Aplopsora. A. nyssae Mains. A. Urediniospores. B. Paraphyses. C. Basidiospores. D. Teliospores.

on
 BOTRYORHIZA Whetzel & Olive
in Olive and Whetzel, Am. J. Bot. 4:44-52 (47). 1917.

FAMILY: Chaconiaceae genus as a synonym of Maravalia, but Botryorhiza has

nomenclatural priority (Ono 1984). Moreover, Bot-
DESCRIPTION: ryorhiza seems to have a different type of aecia.
Spermogonia, aecia, and uredinia unknown. Hennen (personal communication) thinks that
Telia subepidermal, becoming erumpent, pulveru- several species of Aecidium described on plants of
lent, tightly packed, yellow orange; teliospores 1- Hippocrateaceae are likely the aecial state of the
celled, borne singly on long pedicels, wall pale and type species.
uniformly thin, without a differentiated germ pore;
germination occurs without dormancy by con- REFERENCES:
tinued growth of the apex of the spore; basidia Olive, E.W.; Whetzel, H.H. 1917. Endophyllum-like rusts of Puerto
external. Rico. Am. J. Bot. 4:44-52.
Ono, Y. 1978. Taxonomic revisions of the Tribe Oliveae and
TYPE: Botryorhiza hippocrateae Whetzel & Olive morphologically related genera (Uredinales). Ph.D. thesis,
Purdue Univ., West Lafayette, IN. 254 p.
On: Hippocratea volubilis L. (Hippocrateaceae)
Ono, Y. 1984. A monograph of Maravalia (Uredinales). Mycologia
Type locality: Puerto Rico 76:892-911.

REMARKS:
Botryorhiza is a monotypic genus that occurs in
the West Indies and Brazil. Ono (1978) treats the

FIGURE 52. Botryorhiza. B. hippocrateae Whetzel

& Olive. A. Teliospores.
B. Germinating teliospore with
basidium and basidiospores.

92
 CERACEOPSORA Kakish., T. Sato & S. Sato
Mycologia 76:969-974 (969). 1984.

FAMILY: Chaconiaceae flaccida Fr. Schm. (Ranunculaceae) and Elaeagnus

species (E. umbellata Thunb. and E. macrophylla
DESCRIPTION:
Thunb.) (Elaeagnaceae) (Kakishima et al. 1984;
Spermogonia subcuticular, lenticular, hymenium Kakishima and Sato 1984). The morphology of the
flat to slightly concave, Group VI (type 7). Aecia spermogonia (type 7) and the aecia (Aecidium-type)
erumpent, cupulate, Aecidium-type; aeciospores of C. elaeagni is similar to that in species of Ochrop-
catenulate, verrucose. Uredinia erumpent, with sora and Tranzschelia, but the morphology of the
paraphyses, Uredo-type; urediniospores pedicellate, uredinia and telia is distinctly different. The
echinulate. Telia erumpent, white, waxy; teliospores uredinia and telia are similar to those of some
sessile, unicellular, laterally free, grouped on
species of Aplopsora, Chrysocelis, and Chaconia, but
sporogenous basal cells; walls hyaline, smooth;
the telia of Aplopsora are produced under the host
germination occurs without dormancy; basidia
epidermis, and the teliospores are produced in a
external.
single layer without developed sporogenous basal
cells; Chrysocelis has a different type of spermogonia
TYPE: Ceraceopsora elaeagni Kakish., T. Sato &
(Group V, type 4); and Chaconia is autoecious and
S. Sato
has Uredo-type aecia.
On: Elaeagnus umbellata Thunb. (Elaeagnaceae)
Type locality: [baraki, Japan
REFERENCES:
Kakishima, M.; Sato, S. 1984. Inoculation and morphology of
REMARKS:
Ceraceopsora elaeagni. Rep. Tottori Mycol. Inst. 22:171-178.
Only the type species is known. It is heteroecious Kakishima, M.; Sato, T.; Sato, S. 1984. Ceraceopsora, anew genus of
and has been proven to alternate between Anemone Uredinales from Japan. Mycologia 76:969-974.

FIGURE 53. Ceraceopsora. C. elaeagni Kakish., T. Sato & S. Sato. A. A spermogonium. B. An aecium.
C. Teliospores. D. Germinating teliospores. (Reprinted with permission, from Kakishima et al. 1984.)

95
 CHACONIA Juel
Bih. K. Sven. Vetensk. Akad. Handl. 23:1—30 (12). 1897.

FAMILY: Chaconiaceae (Ono and Hennen 1983), and one on Bignoniaceae

(Ono et al. 1988). Complete life cycles are known for
DESCRIPTION: three species, and these are macrocyclic and autoe-
Spermogonia subepidermal or subcuticular, cious. Unlike Cummins and Hiratsuka (1983) and
Group VI (type 5 or type 7). Aecia subepidermal, Ono and Hennen (1983), Santiago (1999) described
erumpent, paraphyses lacking, Uredo-type, or the aeciospores and urediniospores of C. ingae (Syd.)
peripheral and incurved, Calidion-type; aeciospores Cummins as having a reticulate or honeycombed
borne singly on pedicels, echinulate, verrucose in lower surface and prominent ridges at the top (Fig.
lines, or more or less reticulate, pores equatorial or 54B).
obscure. Uredinia similar to the aecia but not
accompanied by spermogonia; urediniospores like REFERENCES:
the aeciospores. Telia subepidermal, erumpent; Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
teliospores 1-celled, laterally free, sessile, and fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
grouped on sporogenous basal cells, germ pore Juel, H.O. 1897. Die Ustilagineen und Uredineen der ersten
probably not differentiated, wall thin and pale; Regnell’schen Expedition. Bih. K. Sven. Vetensk. Akad.
Handl. 23:1-30.
germination occurs without dormancy; basidia
Mains, E.B. 1938. Studies in the Uredinales, the genus Chaconia.
external.
Bull. Torrey Bot. Club 65:625-629.
Ono, Y.; Hennen, J.F. 1983. Taxonomy of the Chaconiaceous
TYPE: Chaconia alutacea Juel
genera (Uredinales). Trans. Mycol. Soc. Jpn. 24:309-402.
On: Pithecellobium divaricatum (Borg.) Benth.
Ono, Y.; Kakishima, M.; Lohsomboon, P.; Manoch, L.;
(Fabaceae) Visarathnonth, N. 1988. Two new species of Uredinales from
Type locality: Paraguay Thailand. Mycologia 80:261-—263.
Santiago, D.V.R. 1999. Taxonomia de Uredinales em plantas da
REMARKS: familia Leguminosae do Cerrado Braxildiro. Ph.D. thesis,
Seven species of Chaconia are known, four on Univ. Brasilia, Brasilia, Brazil. 262 p.
Fabaceae, one on Euphorbiaceae, one on Oleaceae

FIGURE 54. Chaconia. A. Teliospores and a basidium of C. ingae (Syd.) Cummins. B. An aeciospore of C. ingae
(reprinted with permission, from Santiago 1999). C. A urediniospore and teliospores of C. alutacea Juel.

94
 GOPLANA Racib.
Parasit. Algen Pilze Javas, Pt. 2. p. 1-46 (24-25). 1900.

FAMILY: Chaconiaceae distinguish Goplana from Coleosporium, with which

it probably has no close relationship. This genus,
DESCRIPTION: having inconspicuous teliospores embedded in a
Spermogonia and aecia unknown. Uredinia sub- gelatinous matrix with internal basidia, is often
epidermal, erumpent, Uredo-type; urediniospores considered close to or is placed in other hetero-
borne singly on pedicels, pores obscure, super- basidiomycete groups such as Septobasidiales
equatorial in 1 species. Telia subepidermal, erum- (Gaumann 1964) or Auriculariaceae (Raciborski
pent as gelatinous (when wet) cushions; teliospores 1900), and may represent a primitive group in
in a single layer, sessile, embedded in a gelatinous Uredinales.
matrix; germination occurs without dormancy;
basidia internal, becoming 4-celled by division of REFERENCES:
protoplast, each cell producing a basidiospore. Berndt, R. 1999. Neotropical rust fungi: new species and observa-
tions. Mycologia 91:1045-1059.
TYPE: Goplana micheliae Racib. Cummins, G.B. 1935. Notes on some species of the Uredinales.
Mycologia 27:605-614.
On: Michelia velutina DC. (Magnoliaceae)
Ono, Y. 1982. Rusts of yams in Southeast Asia and South Pacific.
Type locality: Java, Indonesia
Trans. Br. Mycol. Soc. 79:423-428.
Gaumann, E. 1964. Die Pilze. Grundzuge ihrer Entwicklungs-
REMARKS: geschichte und Morphologie. Birkhauser Verlag, Basel. p.541.
About 12 species are known (Berndt 1999; Ono, Y.; Hennen, J.F. 1983. Taxonomy of the Chaconiaceous
Cummins 1935; Ono and Hennen 1983; Sydow and genera (Uredinales). Trans. Mycol. Soc. Jpn. 24:369-402.
Sydow 1915), all of which occur in the warmer re- Raciborski, M. 1900. Parasitische Algen und Pilze Javas. Bot. Inst.
gions of the world. They are on plants of Asteraceae, Buitenzorg, Batavia. Pt. 2. p. 1-46.
Dioscoreaceae, Euphorbiaceae, Grossulariaceae, Sydow, H.; Sydow, P. 1915. Monographia Uredinearum. Vol. 3.
Fratres Borntraeger, Leipzig. p. 674-677.
Lauraceae, Magnoliaceae, Meliosmaceae, Rubiaceae,
and Vitaceae. The pedicellate urediniospores

FIGURE 55. Goplana. A telium of G. dioscoreae Cummins.

95
 HEMILEIA Berk. & Broome
in Berkeley, Gard. Chron. 45:1157. 1869.

FAMILY: Chaconiaceae karyogamy and meiosis located in the so-called

urediniospores...” Hemileia may not be a legitimate
DESCRIPTION: name. In the original description, only uredinio-
Spermogonia and aecia unknown. Uredinia typi- spores are described. The illustration is of clustered
cally suprastomatal, but in some species erumpent urediniospores, as seen from the upper side of the
through the ruptured epidermis, Wardia-type; sorus, and a single urediniospore, and a “...group of
urediniospores borne singly on short pedicels, typi- threads with young immature spores highly magni-
cally strongly asymmetrical with 1 flat or concave fied...” The threads and spores can be interpreted as
side, usually smooth, and a dorsal, convex, germinating teliospores with long basidia. How-
echinulate or aculeate side. Telia as the uredinia; te- ever, Ono et al. (1986) consider Hemileopsis Racib.
liospores borne singly on short pedicels, 1-celled, (Paras. Algen Pilze Javas 1:25, 1900) to be the legiti-
usually broader than high, often angular, wall uni- mate name replacing Hemileia.
formly thin, pale, germ pore not visible, perhaps not Hemileia vastatrix is an important disease of culti-
differentiated; germination occurs without dor- vated coffee.
mancy; basidium external.
REFERENCES:
TYPE: Hemileia vastatrix Berk. & Broome Berkeley, M.J. 1869. (No title.) Gard. Chron. 45:1157.
On: Coffea sp. (Rubiaceae) Gopalkrishnan, K.S. 1951. Notes on the morphology of the genus
Type locality: Sri Lanka (Ceylon) Hemileia. Mycologia 43:271-283.
Harr, J. 1979. Hemileia vastatrix Berk. & Br. Publ. Div. Agron. Dep.
REMARKS: Investig. Sandoz, S.A. 27 p.
More than 50 species have been described, most McCain, J.W.; Hennen, J.F. 1984. Development of the uredinial
thallus and sorus in the orange coffee rust fungus, Hemileia
from Africa and Asia. Species of the genus parasitize
vastatrix. Phytopathology 74:714-721.
both monocot and dicot hosts. The “humpbacked” Ono, Y.; Kakishima, M.; Kudo, A.; Sato, S. 1986. Blastospora
urediniospores are so characteristic that numerous smilacis, a teleomorph of Caeoma makinoi and its sorus devel-
species have been described without telia being opment. Mycologia 78:253-262.
known and therefore may not be legitimately named. Rajendren, R.B. 1967. A new type of nuclear life cycle in Hemileia
No monographical work on this genus is available. vastatrix. Mycologia 59:279-285.
Despite many studies and inoculations, no aecial Rajendren, R.B. 1972. Development and parasitism in the genus
Hemileia (Uredinales). Mycopathol. Mycol. Appl. 47: 81-91.
state is known (Thirumalachar 1947). Because
Rayner, R.R.W. 1972. Micologia, historia y biologia de la roya del
the teliospores germinate without dormancy, the cafeto. Inst. Interam. Cienc. Agric. O.E.A., Publ. Misc. 94. 68 p.
aecial state is expected to be systemic or otherwise Thirumalachar, M.J. 1947. Some noteworthy rusts. II. Mycologia
persistent; but, in H. vastatrix, Rajendren (1967) has 39:231-248.
reported a “...unique nuclear behaviour, with

96
FIGURE 56. Hemileia. H. vastatrix Berk. & Broome. A. A uredinium. B. Urediniospores. C, D. Germinating
urediniospores. (Reprinted with permission, from Harr 1979.)

97
 MARAVALIA Arthur
Bot. Gaz. 73:58-69 (60). 1922.

FAMILY: Chaconiaceae or 7 spermogonia. We include them all in Maravalia.

Botryorhiza was considered a synonym of Maravalia
DESCRIPTION: (Ono 1978), but we retain it here as an independent
Spermogonia subcuticular, Group VI (type 5 or genus, in agreement with Ono (1984), until more
7). Aecia subepidermal or subcuticular, erumpent, information becomes available and some nomencla-
with or without paraphyses, Uredo-type; aecio- tural problems have been solved. Dai and Shen
spores borne singly on pedicels, mostly echinulate, (1993) compared many morphological characteris-
germ pores various, often unizonate. Uredinia sub- tics and concluded that Maravalia should be classi-
epidermal, erumpent, Uredo-type, similar to aecia; fied in Chaconiaceae together with Chaconia, Olivea,
urediniospores as aeciospores. Telia subepidermal, and Chrysocelis, but Ochropsora is not closely related
erumpent; teliospores 1-celled, borne singly on to them.
pedicels or occasionally sessile, with or without
sporogenous basal cells, wall thin, pigmented or REFERENCES:
pale, germ pore differentiated or not; germination Arthur, J.C. 1922. Uredinales collected by R. Thaxter and J.B.
Rorer in Trinidad. Bot. Gaz. 73:58-69.
apical and without dormancy; basidia external.
Cummins, G.B. 1950. The genus Scopella of the Uredinales. Bull.
Torrey Bot. Club 77:204-213.
TYPE: Maravalia pallida Arthur & Thaxt.
Dai, Y.C.; Shen, R.X. 1993. A numerical taxonomic study on the
On: Pithecellobium latifolium (L.) Benth. position of Maravalia and other genera of Uredinales.
(Fabaceae) Mycotaxon 48:193—200.
Type locality: Trinidad Evans, H.C. 1993. Studies on the rust Maravalia cryptostegiae, a
potential biological control agent of rubber-vine weed
(Cryptostegia grandiflora, Asclepiadaceae: Periplocoideae) in
REMARKS: Australia, I: Life-cycle. Mycopathologia 124:163-174.
Thirty-one species were recognized by Ono Laundon, G.F. 1964. Angusia (Uredinales). Trans. Br. Mycol. Soc.
(1984) on nine host families. Fabaceae has the largest 47:327-329.
number of species (13) followed by Sapotaceae (10). Mains, E.B. 1939. Scopella gen. nov. of the Pucciniaceae. Ann.
Since Ono’s monograph, two additional species Mycol. 37:57-60.
have been reported from Thailand (Ono et al. 1988a, Mains, E.B. 1939. Studies in the Uredinales, the genus Maravalia.
Bull. Torrey Bot. Club 66:173-179.
b), one on Fabaceae and another on Zingiberaceae.
Ono, Y. 1978. Taxonomic revisions of the tribe Oliveae and
Maravalia, as often treated, is heterogeneous
morphologically related genera (Uredinales). Ph.D. thesis,
because species with type 4 spermogonia are Purdue Univ., West Lafayette, IN. 254 p.
included. When such species as M. ascotele (Syd.) Ono, Y. 1984. A monograph of Maravalia (Uredinales). Mycologia
Mains, M. elata (H. & P. Syd.) Mains, and M. pura 76:892-911.
(Syd.) Mains are recognized as Uromyces species and Ono, Y.; Kakishima, M.; Lohsomboon, P.; Sato, S.; Manoch, L.;
removed from this genus, as we do here, Maravalia, Visarathnonth, N. 1988a. Two rust fungi with pseudo-
superstomatal sori collected in Thailand. Trans. Br. Mycol.
Scopella, and Angusia have most features in Soc. 91:467-472.
common; all are tropical, all have pedicellate,
Ono, Y.; Kakishima, M.; Lohsomboon, P.; Manoch, L.;
nonresting teliospores, all have uredinoid aecia or Visarathnonth, N. 1988b. Two new species of Uredinales from
probably do, and, insofar as known, all have type 5 Thailand. Mycologia 80:261-263.

98
FIGURE 57. Maravalia. A Teliospores of M. pallida Arthur & Thaxt. B. Teliospores of M. kevorkianti
(Cummins) Cummins & Y. Hirats.

99
 OCHROPSORA Dietel
Ber. Dtsch. Bot. Ges. 13:401—402. 1895.

FAMILY: Chaconiaceae Dietel, and O. nambuana (Henn.) Dietel (Hiratsuka

et al. 1992). Aecial stages are known in O. ariae
DESCRIPTION: (Anemone, Ranunculaceae) and O. kraunhiae
Spermogonia subcuticular, Group VI (type 7). (Corydalis, Fumariaceae) (Hiratsuka and Kaneko
Aecia subepidermal, erumpent, with peridium, 1978). Both aecial states are systemic. Because there are
Aecidium-type; aeciospores catenulate, verrucose. no “resting” spores, systemic infections or some other
Uredinia subepidermal, erumpent, the type species compensatory development would be expected. Dai
with abundant, peripheral paraphyses joined at the and Shen (1993) suggested that Ochropsora may not
bottom to peridial tissue, Malupa-type; uredinio- belong in Chaconiaceae.
spores borne singly on short pedicels, echinulate,
pores obscure. Telia subepidermal, in crusts 1 cell REFERENCES:
deep, waxy in appearance, becoming erumpent; te- Dai, Y.C.; Shen, R.X. 1993. A numerical taxonomic study on the
liospores 1-celled, becoming 4-celled by division of position of Maravalia and other genera of Uredinales.
Mycotaxon 48:193-200.
the protoplast (internal basidia), each cell producing
Dietel, P. 1895. Ochropsora, eine neue Uredineengattung. Ber.
1 basidiospore; basidiospores produced on short Dtsch. Bot. Ges. 13:401—402.
sterigmata; germination occurs without dormancy. Hiratsuka, N.; Kaneko, S. 1978. Heteroecism of the Wistaria rust,
Ochropsora kraunhiae (Diet.) Diet. Proc. Jpn. Acad. Ser. B, Phys.
TYPE: Ochropsora sorbi (G. Winter) Dietel Biol. Sci. 54 (6):300-303.
=Caeoma sorbi G. Winter Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
=Ochropsora ariae (Fuckel) Ramsb. Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
Nakayama, K. 1992. The rust flora of Japan. Tsukuba
On: Sorbus aucuparia L. (Rosaceae) Shuppankai, Ibaraki. 1205 + 159 p.
Type locality: Germany

REMARKS:
Four species are known: O. ariae, O. daisensis T.
Hirats. & S. Uchida ex Hirats. f., O. kraunhiae (Dietel)

FIGURE 58. Ochropsora. O. ariae (Fuckel) Ramsb. A. A uredinial paraphysis. B. A urediniospore.

C. Teliospores.

100
 OLIVEA Arthur
Mycologia 9:55-104 (60-61). 1917.

FAMILY: Chaconiaceae REMARKS:

There are 7 or 8 species distributed in warm re-
DESCRIPTION: gions of the earth on members of the Euphorbiaceae,
Spermogonia subcuticular, Group VI (type 7). Lamiaceae, Sapotaceae, and Verbenaceae (Ono and
Aecia subepidermal, deep-seated, opening by rup- Hennen 1983). Some species have strongly asym-
ture of the epidermis, peridium lacking, Caeoma- metrical, lobed urediniospores and aeciospores
type, or subcuticular, erumpent, with basally united (when of the Malupa-type). The latter is true of
peripheral paraphyses, Malupa-type; aeciospores Olivea fimbriata (Mains) Cummins & Y. Hirats.
catenulate in subepidermal aecia, verrucose with (=Tegillum fimbriatum Mains). We consider Tegillum
rodlike verrucae, or borne singly on pedicels and to represent a life cycle variant rather than a distinc-
similar to urediniospores. Uredinia subepidermal tive genus.
or substomatal, erumpent, with peripheral paraphyses
united basally, Malupa-type; urediniospores borne REFERENCES:
singly on pedicels, echinulate, pores equatorial. Arthur, J.C. 1917. Uredinales of Porto Rico based on collections
Telia similar to uredinia in origin, erumpent; telio- by H.H. Whetzel and E.W. Olive. Mycologia 9:55-104.
spores sessile on sporogenous basal cells, 1-celled, Ono, Y.; Hennen, J.F. 1983. Taxonomy of Chaconiaceous genera
wall thin and pale, germ pore apparently not differ- (Uredinales). Trans. Mycol. Soe. Jpn. 24:369-402.

entiated; germination occurs without dormancy;

basidia external.

TYPE: Olivea capituliformis Arthur

On: Alchornea latifolia Sw. (Euphorbiaceae)
Type locality: Puerto Rico

FIGURE 59. Olivea. O. capituliformis Arthur. A. Urediniospores. B. A paraphysate telium.

101
 DASYSPORA Berk. & M.A. Curtis
J. Acad. NatSaiyPhilas Ii Sere27 2277-294 (281). 1854.

FAMILY: Uropyxidaceae REMARKS:

The type is the only species and it occurs in the
DESCRIPTION: American tropics. Dasyspora differs from Puccinia in
Spermogonia subepidermal, Group VI (type 5): the type of spermogonium. The previously reported
Aecia and uredinia unknown, probably not pro- “hyphoid” aecium was found to be an associated
duced. Telia subepidermal, erumpent,; teliospores 2- alga (Hennen and Figueiredo 1981).
celled by transverse septum, borne singly on pedi-
cels, wall pigmented, beset with rodlike verrucae REFERENCES:
mostly branched apically, germ pore 1 in each cell, Berkeley, M.J.; Curtis, M.A. 1854. Exotic fungi from the
Schweinitzian Herbarium, principally from Surinam. J. Acad.
basidia external. Nat. Sci. Phila. II. Ser.2,p. 277-294.
Hennen, J.E; Figueiredo, M.B. 1981. The hyphoid aecium, a rust-
TYPE: Dasyspora foveolata Berk. & M.A. Curtis alga association (Dasyspora- Stomatochroon), and other correc-
=D. gregaria (Kunze) Henn. tions to neotropical rusts (Uredinales). Mycologia 73:350-355.
=Puccinia gregaria Kunze
On: Xylopia sp. (Annonaceae)
Type locality: Surinam

aLe, “aesWe
sites
I vey

msitn
m

FIGURE 60. Dasyspora. Teliospores of D. gregaria

(Kunze) Henn.

102
 DESMELIA H. & P. Syd.
Ann. Mycol. 16:240—248 (241). 1918 (issued 1919).

FAMILY: Uropyxidaceae REMARKS:

There are three or four species of uncertain dis-
DESCRIPTION: tinction, all on ferns. The relationships of the genus
Spermogonia and aecia unknown. Uredinia will remain obscure at least until the aecial state is
suprastomatal, as the telia, Wardia-type; uredinio- known. Edythea has been treated as synonymous
spores borne singly on pedicels, echinulate. Telia with Desmella (Thirumalachar and Cummins 1948),
substomatal, emerging from stomates by means of a but we recognize them as separate genera (Hennen
few sporogenous cells, each of which may produce and Ono 1978; see discussion under Edythea).
several spores, the mature sorus thus suprasto- Hennen and Ono (1978) discussed the similarities of
matal; teliospores borne singly on pedicels, 2-celled telial development in Desmella, Edythea, Cerradoa,
by transverse, oblique, or vertical septa, wall pale or and Prospodium.
pigmented, germ pore 1 in each cell; germinating
without dormancy; basidia external. REFERENCES:
Hennen, J.F.; Ono, Y. 1978. Cerradoa palmaea: the first rust fungus
TYPE: Desmella anemiae H. & P. Syd. on Palmae. Mycologia 70:569-576.
On: Anemia tomentosa Swartz. var. fulva Swartz. Sydow, H.; Sydow, P. 1919. Mykologische Mitteilungen. Ann.
Mycol.16:240-248.
(Schizaeaceae)
Thirumalachar, M.; Cummins, G.B. 1948. Status of the rust genera
=A. fulva Swartz. Allopuccinia, Leucotelium, Edythea, and Ypsilospora. Mycologia
Type locality: Brazil 40:417-422.

FIGURE 61. Desmella. Urediniospores and teliospores of D. anemiae H. & P. Syd.

103
 DIDYMOPSORELLA Thirum.
Sci. Cult. 16:210-215 (210). 1950. (Sydowia 5:23-29 (28). 1951.)

FAMILY: Uropyxidaceae REMARKS:

Three species have been recognized. All of them
DESCRIPTION: are on Toddalia (Rutaceae) from India, Sri Lanka,
Spermogonia subepidermal, Group VI (type 5). Africa, Taiwan, China, and Japan. The paraphyses
Aecia subepidermal, erumpent, Uredo-type; aecio- usually have been overlooked, but were described
spores borne singly on pedicels, echinulate. and illustrated by Ramakrishnan (1951) when he
Uredinia subepidermal, erumpent, Uredo-type; described the synonymous genus Gymnopuccinia.
urediniospores as the aeciospores. Telia subepi-
dermal, with a peripheral layer of hyphae and host REFERENCES:
cells, erumpent as loosely adherent filaments of Ramakrishnan, T.S. 1951. Two new rusts from South India. Trans.
teliospores and intermixed gelatinizing paraphyses; Br. Mycol. Soc. 34:141-145.

teliospores 2-celled, borne singly on pedicels, germ Thirumalachar, M.J. 1951. Critical notes on some plant rusts. II.

pore | or 2 in each cell, wall pale; germination occurs Sydowia 5:23-29.

without dormancy; basidia external.

TYPE: Didymopsorella toddaliae (Thirum. &

Mundkur) Thirum.
=Didymopsora toddaliae Thirum. & Mundkur
=Didymopsorella lemanensis (Doidge) Hirats. f.
On: Toddalia aculeata Pers. var. gracilis (Rutaceae)
Type locality: India

FIGURE 62. Didymopsorella. D. lemanensis (Doidge) Hirats. f. A. A telium. B. Teliospores. C. A germinating

teliospore. D. A paraphysis. E. Urediniospores.

104
 DIPYXIS Cummins & J.W. Baxter
Mycologia 59:367-370 (368). 1967.

FAMILY: Uropyxidaceae REMARKS:

In addition to the type, there is one other species,
DESCRIPTION: D. viegasti Jerst.) Cummins & J.W. Baxter on Ar-
Spermogonia subcuticular, Group VI (type 7). rabidaea sp. in Brazil (Cummins and Baxter 1967;
Aecia subepidermal, erumpent, Uredo-type; Hennen et al. 1982). Both hosts are Bignoniaceae and
aeciospores borne singly on pedicels, as the both rusts have echinulate teliospores and reniform
urediniospores. Uredinia subepidermal, erumpent, urediniospores, i.e. with a concave side with germ
Uredo-type; urediniospores borne singly on pedicels, pores near the hilum or strongly subequatorial.
echinulate, pores zonate. Telia subepidermal, erum-
pent; teliospores borne singly, 2-celled by transverse REFERENCES:
septum, wall pigmented, germ pores 3, rarely 4, per Cummins, G.B.; Baxter, J.W. 1967. Dipyxis, a new genus of
cell; basidia external. Uredinales. Mycologia 59:367-370.
Hennen, J.F.; Hennen, M.M.; Figueiredo, M.B. 1982. Indice das
ferrugens (Uredinales) do Brasil. [Index of rust fungi
TYPE: Dipyxis mexicana Cummins & J.W. Baxter (Uredinales) of Brazil.] Arq. Inst. Biol., Sao Paulo 49 (Supl.
On: Adenocalymma calderonii (Standl.) Sieb. 1):1-202.
(Bignoniaceae)
Type locality: Sinaloa, Mexico

FIGURE 63. Dipyxis. D. mexicana Cummins & J.W. Baxter. A. Urediniospores.

B. Teliospores.

105
 EDYTHEA H.S. Jacks.
Mycologia 23:96—116 (97-98); 1 plate. 1931.

FAMILY: Uropyxidaceae of the Palmae, differs in that the teliospores are

deeply pigmented. To accept degrees of pigmen-
DESCRIPTION: tation of spore walls as generic characteristics is
Spermogonia and aecia unknown. Uredinia hardly tenable. Consequently, Cerradoa palmaea
suprastomatal like the telia, Wardia-type; uredinio- (Hennen and Ono 1978) has been reduced to syn-
spores borne singly on pedicels, echinulate. Telia onymy as Edythea palmaea (J.F. Hennen & Y. Ono)
subepidermal, but emerging through stomata by Cummins & Y. Hirats. (Cummins and Hiratsuka
means of elongate cells that produce sporogenous 1983). Edythea differs from Desmella, in which the
cells apically, which, in turn, produce teliospores teliospores arise on the apices of elongate sporo-
borne singly on pedicels; teliospores 2-celled, sep- genous cells that emerge through the stomata.
tum vertical or transverse, germ pore 1 in each cell; Whether this is a significant distinction is question-
basidium external. able.

TYPE: Edythea quitensis (Lagerh.) H.S. Jacks. & REFERENCES:

Holw. Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
=Uropyxis quitensis Lagerh. fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
On: Berberis sp. (Berberidaceae) Hennen, J.F.; Ono, Y. 1978. Cerradoa palmaea: the first rust fungus

Type locality: Quito, Ecuador on Palmae. Mycologia 70:569-576.

Jackson, H.S. 1931. The rusts of South America based on the
Holway collections. III. Mycologia 23:96-116; 1 plate.
REMARKS:
Thirumalachar, M.J.; Cummins, G.B. 1948. Status of the rust
The type and two other species occur on Berberis genera Allopuccinia, Leucotelium, Edythea, and Ypsilospora.
in South America. Cerradoa palmaea J.P. Hennen & Y. Mycologia 40:417-422.
Ono, which parasitizes Attalea ceraensis Barb. Rodr.

FIGURE 64. Edythea. A. Teliospores of E. tenella H.S. Jacks. B. Teliospores of E. palmaea (J.F. Hennen &
Y. Ono) Cummins & Y. Hirats.

106
 KIMUROMYCES Dianese, Santos, Medeiros & Furlanetto
Fitopatol. Bras. 20:251-255 (251-252). 1995.

FAMILY: Uropyxidaceae clearly differs from K. cerradensis Dianese, Santos,

Medeiros & Furlanetto in teliospore morphology.
DESCRIPTION: Dianese et al. (1995) consider Kimuromyces to be
Spermogonia and aecia not known. Uredinia more closely related to genera of the Uropyxidaceae
subepidermal, becoming erumpent, dense (e.g. Uropyxis, Prospodium, Sorataea) than to those of
paraphyses mostly peripheral, Calidion-type; the family Pucciniaceae (e.g. Puccinia, Cleptomyces),
urediniospores colored (brown), borne singly on an affiliation suggested by Hawksworth et al. (1995).
sympodially proliferating cells, echinulate, with 2 Very prominent ornamentation on the teliospores
equatorial germ pores. Telia brown, erumpent, somewhat resembles that of Esalque holwayi (HS.
sometimes with uredinia, paraphysate, paraphyses Jacks.) J.F. Hennen, A.A. Carvalho & Figueiredo.
similar to those of uredinia; teliospores pedicellate,
2-celled, with transverse septum, both cells of telio- REFERENCES:
spores with prominent simple or sometimes Dianese, J.C.; Santos, L.T.P.; Medeiros, R.B.; Furlanetto, C. 1995.
branched appendages, germ pore 1 per cell; basidia Kimuromyces cerradoensis gen. et sp. nov., the rust fungus of
external. “Gongalo Alves” (Astronium fraxinifolium-Anacardiaceae)
from the Brazilian cerrado. Fitopatol. Bras. 20:251-255.
Hawksworth, D.H.; Kirk, P.M.; Sutton, B.C.; Pegler, D.N. 1995.
TYPE: Kimuromyces cerradensis Dianese, Santos,
Ainsworth & Bisby’s dictionary of the fungi. 8th ed. Int.
Medeiros & Furlanetto Mycol. Inst., CAB Int., Surrey, UK. 616 p.
On: Astronium fraxinifolium Schott. ex Spreng.
(Anacardiaceae)
Type locality: Brazil

REMARKS:
Another species of rust, Leptinia brasiliensis Juel
(Puccinia brasiliensis), is known on Astronium sp., but

FIGURE 65. Kimuromyces. A, B. Teliospores of K. cerradensis Dianese, Santos, Medeiros & Furlanetto (reprinted with
permission, courtesy of Dr. J.C. Dianese).

107
 MACRUROPYXIS Azbukina
Komarov Read., Vladivost. 19:15-62 (20-21). 1972.

FAMILY: Uropyxidaceae Macruropyxis fraxini (Kom.) Azbukina is microcyclic

and has type 7 (Group VI) spermogonia. The exam-
DESCRIPTION: ined spermogonia of Dipyxis were too old to classify
Spermogonia subcuticular, Group VI (type 7). with certainty, but type 3 (Group I) or perhaps type 7
Aecia and uredinia unknown, probably not (Group VI) were reported originally. A possible
produced. Telia subepidermal, erumpent, without difference of spermogonial types and the sievelike
paraphyses; teliospores 2-celled by transverse sep- appearance of the germ pores are all that separate
tum, wall bilaminate, the inner wall pigmented, the Macruropyxis and Dipyxis. Some favor placing M.
outer wall hyaline, germ pores 3, rarely 4, in each fraxini in Uropyxis (Hiratsuka et al. 1992), but we
cell, sievelike in appearance and apparently with recognize this as a separate genus.
multiple perforations, spores borne singly on pedi-
cels; basidia external. REFERENCES:
Azbukina, Z. 1972. Rust fungi of far east. Lec. Komarovianae
TYPE: Macruropyxis fraxini (Kom.) Azbukina (Komarov Read.), Vladivost. 19:15-62.

=Puccinia fraxini Kom. Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
=Uropyxis fraxini (Kom.) Magnus
Nakayama, K. 1992. The rust flora of Japan. Tsukuba
On: Fraxinus sieboldiana Blume (Oleaceae) Shuppankai, Tsukuba. 1205 + 159 p.
= F. lanaginosa Koidz.
Type locality: Far Eastern Region, Russia

REMARKS:
This monotypic genus is similar to and perhaps
not separable from Dipyxis in significant features.

FIGURE 66. Macruropyxis. Teliospores of M. fraxini (Kom.) Azbukina.

108
 MIMEMA HS. Jacks.
Mycologia 23:332-364 (338-339). 1931.

FAMILY: Uropyxidaceae as a synonym of Sorataea (Cummins and Hiratsuka

1983), but as discussed by Dianese et al. (1994), their
DESCRIPTION: teliospore morphology and the mode of basidial
Spermogonia and aecia not known. Uredinia formation clearly distinguish between the two;
with abundant peripheral paraphyses, Calidion- therefore we recognize these genera as different.
type; urediniospores finely and closely Teliospores of Mimema are very similar to those of
echinulate/verrucose (M. holwayi H.S. Jacks.) or Hamaspora, which occurs only on Rubus (Rosaceae)
echinulate (M. venturae Dianese et al.), germ pores and has Group IV (type 8 or 10) spermogonia, but
equatorial (1 to 6). Telia develop following uredinia they may not be closely related. The spermogonial
in the same sori, emerging from the dense periph- and aecial states of Mimema are not known, but if
eral paraphyses as columns of teliospore tendrils; discovered, their relationship to Hamaspora or other
teliospores cylindrical or fusiform, multicelled, usu- genera could be clarified. Superficially, the columns
ally 4 (3-6), with long tapering hyaline pedicels; ger- of teliospore tendrils emerging from the dense
mination through discrete germ pores; basidia peripheral paraphyses are similar to some species of
external. Crossopsora.

TYPE: Mimema holwayi H.S. Jacks. REFERENCES:

On: Cassia sp. (possibly C. versicolor Mey.) Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
(Fabaceae) fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
Type locality: Bolivia Dianese, J.C.; Santos, L.T.; Medeiros, R.B.; Sanchez, M. 1994.
Mimema venturae sp. nov. on Delbergia miscolobium in Brazil.
REMARKS: Mycol. Res. 98:786-788.
Jackson, H.S. 1931. The rusts of South America based on the
The type species and another one, M. venturae
Holway collections. IV. Mycologia 23:332-364.
Dianese et al. from Brazil (Dianese et al.1994), both
on Fabaceae, are known. Mimema has been treated

FIGURE 67. Mimema. A. Teliospores of M. holwayi H.S. Jacks. B. A uredinium of M. venturae Dianese,
Santos, Medeiros & Sanchez. C. A telium of M. venturae. (A reprinted with permission, from
Jackson 1931, B and C from Dianese et al. 1994.)

109
 NEWINIA Thaung
Mycologia 65:702-704 (702). 1973.

FAMILY: Uropyxidaceae REMARKS:

In addition to the type species, there are two
DESCRIPTION: others, N. kigeliae Eboh on Kigelia africana (L.) Benth.
Spermogonia subcuticular, Group VI (type 7). from Nigeria (Eboh 1983) and N. thaiana Kakish. & Y.
Aecia subepidermal, erumpent, with paraphyses, Ono on Stereospermum cylindricum Pierre ex D. Dop.
Uredo-type; aeciospores borne singly on pedicels, from Thailand (Kakishima et al. 1988). All three host
essentially as the urediniospores. Uredinia sub- genera belong in the family Bignoniaceae. The telio-
epidermal, erumpent, with paraphyses, Uredo-type; spores are similar to those of Kuehneola, Frommeélla,
urediniospores borne singly on pedicels, echinulate, and Xenodochus, but these taxa have Group IV (type
pores equatorial. Telia subepidermal, erumpent;, 10 or 11) spermogonia.
teliospores borne singly on pedicels, 3- to many-
celled by transverse septa, wall pigmented, smooth, REFERENCES:
germ pore 1, apical in terminal cell, 1 or less often 2 Eboh, D.O. 1983. A new species of Newinia from Nigeria.
next to the septum in other cells; basidia external. Mycologia 75:316-318.
Kakishima, M.; Lohsomboon, P.; Ono, Y.; Manoch, L.;
Visarathanonth, N. 1988. Newinia thaiana, a new rust fungus
TYPE: Newinia heterophragmae Thaung from Thailand. Mycologia 80:397-400.
On: Heterophragma sulphureum Kurz. Thaung, M.M. 1973. A new genus of rusts from Burma.
(Bignoniaceae) Mycologia 65:702-704.
Type locality: Burma

FIGURE 68. Newinia. A. Urediniospores and teliospores of N. heterophragmae Thaung. B. Urediniospores of N. thaiana
Kakish. & Y. Ono. C, D. Teliospores of N. thaiana. (B—D reprinted with permission, from Kakishima et al.
1988.)

110
 PHRAGMOPYXIS Dietel
in Engler and Prantl, Nat. Pflanzenfam. 1(1):24-81 (70). 1897.

FAMILY: Uropyxidaceae TYPE: Phragmopyxis deglubens (Berk. & M.A.

Curtis) Dietel
DESCRIPTION: =Triphragmium deglubens Berk. & M.A. Curtis
Spermogonia subcuticular, Group VI (type 7). On: Unidentified plant (Fabaceae)
Aecia subepidermal, erumpent; aeciospores catenu- Type locality: Mexico
late, verrucose, Caeoma-type, or echinulate with
REMARKS:
spores borne singly on pedicels, Uredo-type.
There are four species, all autoecious and all on
Uredinia subepidermal or subcuticular, erumpent,
legumes (Cummins 1978). Phragmopyxis acuminata
Uredo-type; urediniospores borne singly on pedi-
cels, echinulate, germ pores scattered, obscure. Telia
(Long) P. & H. Syd. is microcyclic, and P. leonensis
subepidermal or subcuticular, erumpent; telio-
Cummins, from Sierra Leone, is the only extra-
spores borne singly on usually hygroscopic pedi- American species. It differs from the others in
cels, mostly 3-celled by transverse septa, wall having subcuticular uredinia and telia.
conspicuously bilaminate, the outer layer pale to REFERENCES:
colorless and easily separable, germ pores 3 or 4 in Cummins, G.B. 1978. Rust fungi on legumes and composites in
each cell, pedicel often swollen basally and some- North America. Univ. Arizona Press, Tucson, AZ. 424 p.
times dissolving into the mounting medium; Dietel, P. 1897. Hemibasidii und Uredinales. In A. Engler and K.
basidium external. Prantl. Nat. Pflanzenfam. 1(1):24-81.

FIGURE 69. Phragmopyzxis. A. A teliospore, a paraphysis, and a urediniospore of P. noelti J.W. Baxter. B. Two
teliospores and a urediniospore of P. deglubens (Berk. & M.A. Curtis) Dietel.

NAGI
 POROTENUS Viégas
Bragantia 19:95-99 (98). 1960.

FAMILY: Uropyxidaceae asymmetrical. The species occur on Memora and

Amphilophium of the Bignoniaceae and Lippia of
DESCRIPTION: Verbenaceae. Three species assigned first to Puccinia
Spermogonia Group VI (type 5, subepidermal, or and then to Prospodium (Cummins 1940) were trans-
type 7, subcuticular). Aecia subepidermal, erum- ferred to Porotenus: P. elatipes (Arthur & Holw.)
pent, Uredo-type; aeciospores borne singly on pedi- Cummins & Y. Hirats., P. permagnum (Arthur &
cels, echinulate, pores equatorial. Uredinia sub- Holw.) Cummins & Y. Hirats., and P. depallens
epidermal, erumpent, Uredo-type; urediniospores as (Arthur & Holw.) Cummins & Y. Hirats. (Cummins
the aeciospores. Telia subepidermal, erumpent; and Hiratsuka 1983). Hennen and Sotao (1996)
teliospores 2-celled by transverse septum, borne added two new species on Memora from Brazil (P.
singly on pedicels; each cell germinating by elonga- bibasiporulus J.F. Hennen & Sotao and P. biporus J.F.
tion of the apex, with a differentiated area in the Hennen & Sotao).
wall, or with a discrete germ pore. Porotenus has characters in common with Pro-
spodium, and the teliospores of P. concavus are simi-
TYPE: Porotenus concavus Viegas
lar to those of Sorataea.
On: Memora glaberrima K. Schumann
(Bignoniaceae) REFERENCES:
Type locality: Brazil Cummins, G.B. 1940. The genus Prospodium (Uredinales).
Lloydia (Cincinnati) 3:1-78.
REMARKS: Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
This is a small genus of seven species, all from the fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
Neotropics. They are autoecious and either macro- Hennen, J.F.; Sotao, H.M.P. 1996. New species of Uredinales on
cyclic or microcyclic. The type species may attack Bignoniaceae from Brazil. SIDA Contrib. Bot. 17:173-184.

leaf blades and stems, often causing hypertrophy Viegas, A.P. 1960. Porotenus n.gen. Uredinearum. Bragantia
19:95-99.
and distortion. The urediniospores are radially

FIGURE 70. Porotenus. A. A teliospore and urediniospores of

P. elatipes (Arthur & Holw.) Cummins & Y. Hirats.
B. Urediniospores and teliospores of P. concavus Viégas.

L2.
 PROSPODIUM Arthur
J. Mycol. 13:28-32 (31). 1907.

FAMILY: Uropyxidaceae REMARKS:

About 50 species of Prospodium have been
DESCRIPTION: described; all are autoecious, and some are micro-
Spermogonia subcuticular, Group VI (type 7). cyclic. Most occur on Bignoniaceae and the rest on
Aecia subepidermal, erumpent, Uredo-type; aecio- Verbenaceae (Cummins 1940; Hennen and Sotao
spores borne singly on pedicels, mostly echinulate. 1996). The Uredo-type aecia are never supra-
Uredinia subepidermal, erumpent in some species, stomatal, although the uredinia of the same species
Uredo-type; or substomatal, emerging through may be. Microcyclic species never produce supra-
stomata to sporulate above the leaf surface, usually stomatal telia, although the putative parental
with basally united peripheral paraphyses, species may; they always assume the aecial habit.
basketlike, with paraphyses on the rim, Malupa- Three sections have been recognized (but not for-
type; urediniospores borne singly on pedicels, malized as subgenera): Euprospodium, with erum-
mostly echinulate, the wall simple, or with an outer pent uredinia and telia; Nephlyctis, which consists
partial, hyaline, hygroscopic envelope that forms of microcyclic species only; and Cyathopsora, with
one or two apical thickenings, pores 2 (or rarely 3), suprastomatal uredinia and telia. Despite the diver-
equatorial. Telia as the uredinia of a species; telio- gency in sorus position, the species are obviously
spores 2-celled by transverse septum, borne singly closely related.
on pedicels, which are often appendaged basally,
spore wall pigmented, often obviously bilaminate REFERENCES:
and usually echinulate or verrucose, germ pore | in Arthur, J.C. 1907. New genera of Uredinales. J. Mycol. 13:28-32.
Cummins, G.B. 1940. The genus Prospodium (Uredinales).
each cell; basidia external.
Lloydia (Cincinnati) 3:1—78.
Hennen, J.F.; Sotao, H.M.P. 1996. New species of Uredinales on
TYPE: Prospodium appendiculatum (G. Winter) Arthur Bignoniaceae from Brazil. SIDA Contrib. Bot. 17:173-184.
=Puccinia appendiculata G. Winter
On: An unidentified plant of Bignoniaceae
Type locality: Mexico
FIGURE 71. Prospodium. A. Schematic drawings of three types of urediniospores: a simple echinulate urediniospore
(left), a urediniospore with partial hygroscopic envelope with one apical thickening (center), and a
urediniospore with partial hygroscopic envelope with two apical thickenings (right). B. Schematic
drawing of a uredinium of Malipa-type. C. A teliospore of P. couraliae Syd. D. Urediniospores of
P. couraliae. E. Teliospores of P. appendiculatum (G. Winter) Arthur.

114
 SORATAEA Syd.
Ann. Mycol. 28:432-447 (432). 1930.

FAMILY: Uropyxidaceae Cummins (1948) suggested that Leucotelium did not

differ substantially from Sorataea. The species of
DESCRIPTION: Leucotelium are heteroecious with aecial states on
Spermogonia subcuticular, Group VI (type 7). Ranunculaceae and telial states on Rosaceae. Insofar
Aecia subepidermal, erumpent, Uredo- or Aecidium- as known, the species have type 7 spermogonia.
type, with peridium; aeciospores either borne singly Cummins and Hiratsuka (1983) believed that
or catenulate. Uredinia subepidermal, erumpent, Leucotelium should be treated as a synonym of
with or without paraphyses, Uredo-type; uredinio- Sorataea and so proposed the following transfers:
spores borne singly on pedicels, echinulate, pores Sorataea cerasi (Castagne) Cummins & Y. Hirats.
equatorial. Telia subepidermal, erumpent, with or (Puccinia pruni-persicae Hori, Leucotelium pruni-
without peripheral paraphyses; teliospores 2- to persicae Tranzschel) and S. padi (Tranzschel)
several-celled by transverse septa, borne singly on Cummins & Y. Hirats. (Leucotelium padi Tranzschel).
pedicels grouped on basal sporogenous cells in at Cummins and Hiratsuka (1983) treated Mimema asa
least some species, wall pale but usually pigmented, synonym of Sorataea, but following Dianese et al.
pore obscure if differentiated; germinating without (1994), we recognize them as separate genera.
dormancy, apparently mostly by growth of the apex
of the cells; basidia external. REFERENCES:
Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
TYPE: Sorataea amiciae Syd. fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
On: Amicia lobbiana Benth. (Fabaceae) Dianese, J.C.; Santos, L.T.; Medeiros, R.B.; Sanchez, M. 1994.
Type locality: Sorata, Bolivia Mimema venturae sp. nov. on Delbergia miscolobium in Brazil.
Mycol. Res. 98:786-788.
REMARKS: Eboh, D.O.; Cummins, G.B. 1980. Species of Sorataea (Uredinales).
Mycologia 72:203-204.
About 10 species are recognized; most of them
Savile, D.B.O. 1971. Generic disposition and pycnium type in
were originally described as species of Puccinia or Uredinales. Mycologia 63:1089-1091.
related genera, but were transferred to this genus Sydow, H. 1930. Novae fungorum species. XX. Ann. Mycol.
because of the different type of spermogonia (type 7 28:432-447.
instead of type 4) (Eboh and Cummins 1980; Thirumalachar, M.J.; Cummins, G.B. 1948. Status of Allopuccinia,
Cummins and Hiratsuka 1983; Savile 1971). Most of Leucotelium, Edythea, and Ypsilospora. Mycologia 40:417-422.
the species are on Fabaceae. Thirumalachar and

FIGURE 72. Sorataea. A. Teliospores of S. cerasi (Castagne) Cummins & Y. Hirats. B. Urediniospores and
teliospores of S. amiciae Syd.

MS
 TRANZSCHELIA Arthur
Rés. Sci. Congr. Int. Vienne. p. 331-348 (340). 1906.

FAMILY: Uropyxidaceae and Hennen 1990). Most macrocyclic species are

heteroecious, producing aecia on the Ranuncu-
DESCRIPTION: laceae and telia on the Rosaceae, but there are two
Spermogonia subcuticular, Group VI (type 7). macrocyclic autoecious species on Anemone
Aecia subepidermal, erumpent, with peridium, (Ranunculaceae). The microcyclic species, includ-
Aecidium-type; aeciospores catenulate, verrucose. ing the type species, Tranzschelia cohaesa (Long)
Uredinia subepidermal, erumpent, Uredo-type, Arthur, are on Ranunculaceae, in accordance with
with oblong to capitate paraphyses; urediniospores Tranzschel’s Law. The genus occurs circumglobally
borne singly on pedicels, echinulate, germ pores in the north temperate region and southward on cul-
equatorial. Telia subepidermal, erumpent; telio- tivated crops.
spores usually 2-celled by transverse septum, borne
singly on pedicels that tend to adhere basally in REFERENCES:
groups, the spores thus tending to be in bunches, Arthur, J.C. 1906. Eine auf die Struktur und Entwicklungs-
wall pigmented, echinulate or verrucose, germ pore geschichte begrtindete Klassifikation der Uredineen. Reés. Sci.
Congr. Int. Vienne. p. 331-348.
1 in each cell; basidia external.
Bennell, A.P.; Henderson, D.M. 1978. Urediniospore and telio-
spore development in Tranzschelia (Uredinales). Trans. Br.
TYPE: Tranzschelia cohaesa (Long) Arthur Mycol. Soc. 71:271-278.
=Puccinia cohaesa Long Dunegan, J.C. 1938. The rusts of stone fruits. Phytopathology
On: Anemone caroliniana Walter 28:411-427.
=A. berlandieri Pritzel (Ranunculaceae) Laundon, G.F. 1975. Taxonomy and nomenclature notes on
Type locality: Texas, USA Uredinales. Mycotaxon 3:133-161.
Lépez-Franco, R.M.; Hennen, J.F. 1990. The genus Tranzschelia
(Uredinales) in the Americas. Syst. Bot. 15:560-591.
REMARKS:
Tranzschel, W.; Litvinov, M. 1939. [Rust fungi of the genus
There are about 15 species worldwide and 8 spe-
Tranzschelia on the Prunoideen.]J. Bot. 24:247—253. (In Russian.)
cies are known from the Americas (L6pez-Franco

FIGURE 73. Tranzschelia. A. Urediniospores, a paraphysis, and a teliospore of T. discolor (Fuckel) Tranzschel.
B. Teliospores of T. pruni-spinosae (Pers.) Dietel. C. A teliospore of T. pruni-spinosae (courtesy of Dr. S. Sato
and Dr. M. Kakishima).

116
 UROPYXIS J. Schrot.
Hedwigia 14:161-172 (165). 1875.

FAMILY: Uropyxidaceae REMARKS:

The species are more common in warm regions,
DESCRIPTION: but U. amorphae (Curtis) J. Schrot. is known in
Spermogonia subcuticular, Group VI (type 7). Canada. The 15 or so species are all autoecious;
Aecia subepidermal, similar to uredinia, Calidion- some are macrocyclic and others are microcyclic
type; aeciospores borne singly on pedicels. Uredinia (Baxter 1959). Most occur on legumes. Uropyxis
subepidermal, erumpent, with peripheral differs from Puccinia and Cumminsiella because of
paraphyses, Calidion-type; urediniospores borne the type 7 spermogonia, and from Prospodium be-
singly on pedicels, echinulate, pores scattered, cause of the two pores per teliospore cell.
obscure. Telia subepidermal, erumpent; teliospores
borne singly on pedicels that are often hygroscopic, REFERENCES:
2-celled by transverse septum, wall usually conspic- Baxter, J.W. 1959. A monograph of the genus Uropyxis. Mycologia
uously hygroscopic and bilaminate, the outer wall 51:210-226.
often easily separable, sculptured or smooth, germ Cummins, G.B. 1978. Rust fungi on legumes and composites in
North America. Univ. Arizona Press, Tucson, AZ. 424 p.
pores 2 in each cell; basidia external.
Schréter, J. 1875. Uber einige amerikanische Uredineen.
Hedwigia 14:161-172.
TYPE: Uropyxis amorphae (Curtis) J. Schrot.
=Puccinia amorphae Curtis
On: Amorpha herbacea Walt. (Fabaceae)
Type locality: S. Carolina, USA

FIGURE 74. Uropyxis. A. A teliospore, a paraphysis, and a urediniospore of U. amorphae (Curtis) J. Schrét.
B. A teliospore of U. holwayi (Arthur) Arthur. C. Teliospores of U. heterospora J.F. Hennen & Cummins.

UL
 ATELOCAUDA Arthur & Cummins
Ann. Mycol. 31: 41-45 (41). 1933.

FAMILY: Pileolariaceae It is less easily separable from Pileolaria

(Thirumalachar and Kern 1955), although it is
DESCRIPTION: doubtful that the two genera are closely related.
Spermogonia subepidermal or subcuticular, Atelocauda (at least its Acacia-infecting species) is
Group VI (type 5 or 7). Aecia subepidermal, erum- more likely related to Uromycladium. Two endo-
pent, Uredo-type; aeciospores borne singly on pedi- cyclic species related to Atelocauda are Endoraecium
cels, similar to the urediniospores. Uredinia sub- hawaiiense Hodges & D.E. Gardner and E. acaciae
epidermal, erumpent, Uredo-type; urediniospores Hodges & D.E. Gardner (Hodges and Gardner 1984)
borne singly on pedicels, pigmented, most and per- (see Endoraecium). Chen et al. (1996) reported that
haps all reticulate, germ pores zonate (equatorial or although it is demicyclic, A. koae produces aecio-
bizonate). Telia subepidermal, erumpent; telio- spores together with teliospores; the aeciospores
spores 1-celled, borne singly on pedicels, wall pale cannot initiate infections and thus it behaves as a
or pigmented, germ pore 1, pedicels simple; basidia microcyclic species.
external.
REFERENCES:
TYPE: Atelocauda incrustans Arthur & Cummins Arthur, J.C.; Cummins, G.B. 1933. New species of Uredinales.
On: Lonchocarpus sp. (Fabaceae) Ann. Mycol. 31:41-45.
Type locality: Panama Chen, W.Q.; Gardner, D.E.; Webb, D.T. 1996. Biology and life cycle
of Atelocauda koae, an unusual demicyclic rust. Mycoscience
37:91-98.
REMARKS:
Gardner, D.E. 1981. Nuclear behavior and clarificaton of the
The type species is microcyclic. Three others are spore stages of Uromyces koae. Can. J. Bot. 59:939-946.
demicyclic or macrocyclic and occur on Acacia in Gardner, D.E. 1991. Atelocauda angustiphylloda n. sp.,a microcyclic
Australia, New Zealand, China, Taiwan, Japan, and rust on Acacia koa in Hawaii. Mycologia 83:650-653.
Hawaii. They are A. bicincta (McAlpine) Cummins Gardner, D.E.; Hodges, C.S., Jr. 1985. Spore surface morphology
& Y. Hirats. (Uromyces bicinctus McAlpine), A. koae of Hawaiian Acacia rust fungi. Mycologia 77:575-586.
(Arthur) Cummins & Y. Hirats. (U. koae Arthur), Hodges, C.S., Jr.; Gardner, D.E. 1984. Hawaiian forest fungi. IV.
Rusts on endemic Acacia species. Mycologia 76:332-349.
A. digitata (G. Winter) Cummins & Y. Hirats. (U.
Ono, Y. 1984. A monograph of Maravalia (Uredinales). Mycologia
digitatus G. Winter) and A. hyalospora (Sawada) Y.
76: 892-911.
Ono. Another microcyclic species, A. angustiphylloda Thirumalachar, M.J.; Kern, FD. 1955. The rust genera Allotelium,
D.E. Gardner was described from Hawaii on Acacia Atelocauda, Coniostelium and Monosporidium. Bull. Torrey Bot.
koa A. Gray (Gardner 1991). Atelocauda is readily Club 82:102-107.
separable from Uromyces because of the spermogonia.

118
FIGURE 75. Atelocauda. A. A germinating teliospore of A. koae (Arthur) Cummins & Y. Hirats. (redrawn from Hodges
and Gardner 1984). B. A teliospore of A. digitata (G. Winter) Cummins & Y. Hirats. C. A teliospore of
A. incrustans Arthur & Cummins. D. Teliospores of A. koae. E. Teliospores and a urediniospore of
A. bicincta (McAlpine) Cummins & Y. Hirats. F. An aeciospore of A. digitata (reprinted with permission,
from Gardner and Hodges 1985).

PLS
 PILEOLARIA Castagne
Obs. Plantes Acotylédonées Fam. Urédinées. 1. 35 p. (22). 1842.

FAMILY: Pileolariaceae Teliospores are typically somewhat discoid and

verrucose or reticulately verrucose, and the
DESCRIPTION: urediniospores are ridged or beaded in longitudinal
Spermogonia subcuticular, Group VI (type 1), or spiral patterns. A few species on Fabaceae have
Aecia subepidermal, erumpent, Uredo-type; aecio- been referred to Pileolaria but we treat these sepa-
spores borne singly on pedicels, verrucose or rately as species of Atelocauda. The life cycle of P.
echinulate, often in spiral or longitudinal lines, klugkistiana Dietel was proven by inoculation exper-
pores zonate. Uredinia subepidermal, erumpent, iments (Kakishima et al. 1984), and an excellent
Uredo-type; urediniospores as the aeciospores. Telia morphological work was published on three Japa-
subepidermal, erumpent; teliospores 1-celled, nese species by Katsuya et al. (1980). No comprehen-
borne singly on pedicels, wall thick and pigmented, sive monograph is available for this genus.
variously sculptured, germ pore 1; germination
mostly after dormancy; basidia external. REFERENCES:
Castagne, L. 1842. Observations sur quelques plantes
TYPE: Pileolaria terebinthi Castagne acotylédonées de la famille des Urédinées et dans les sous-
On: Pistacia terebinthus L. (Anacardiaceae) tribus des Nemasporées et des Aecidinées, recueillies dans le
Dep. des Bouches-du-Rhoéne. 1. Marseille. 35 p.
Type locality: France
Kakishima, M.,; Sato, T.; Sato, S. 1984. Notes on two rust fungi,
Pileolaria klugkistiana and Nyssopsora cedrelae. Trans. Mycol.
REMARKS: : Soc. Jpn. 25:355-359.
There are about 20 to 24 species, most of which Katsuya, K.; Kakishima, M.; Sato, S. 1980. Spore surface structure
are macrocyclic and autoecious. They parasitize of three Pileolaria species in Japan. Rep. Tottori Mycol. Inst.
species of Rhus, Pistacia, and other Anacardiaceae. Jpn. 18:163-167.

FIGURE 76. Pileolaria. A. Teliospores and a urediniospore of P. brevipes Berk. & Rabenh. B. Teliospores of P. effiisa
Peck. C, D. Urediniospores of P. shiraiana (Dietel & Syd.) S. Ito. E, F. Teliospores of P. shiraiana.
(C-F reprinted with permission, from Katsuya et al. 1980.)

120
 UROMYCLADIUM McAlpine
Ann. Mycol. 3:303-323 (321). 1905.

FAMILY: Pileolariaceae REMARKS:

Seven species have been recognized, all occur-
DESCRIPTION: ring on Acacia and Albizia of the Fabaceae. Both
Spermogonia subepidermal, Group VI (type 5 or macrocyclic and microcyclic species are known; all
perhaps sometimes type 7). Aecia subepidermal, are autoecious. Some species cause conspicuous
erumpent, Uredo-type; aeciospores borne singly on branch galls and distortions of the host plant. Atelo-
pedicels, verrucose or reticulate, pores equatorial. cauda may be related to Uromycladium. Some species
Uredinia subepidermal, erumpent, Uredo-type; of Atelocauda also produce distortions of host tissues
urediniospores similar to aeciospores. Telia subepi- and have urediniospores with surface markings
dermal, erumpent; teliospores 1-celled with 1 to 3 similar to Uromycladium. These species have type 5
borne on a usually branched and septate pedicel spermogonia, as does Uromycladium, but they have
that may also bear hygroscopic sporelike cysts, nonseptate pedicels and no sterile cells.
germ pore 1, the teliospores usually pigmented;
basidia external. REFERENCES:
Cunningham, G.H. 1931. The rust fungi of New Zealand.
MclIndoe, Dunedin, NZ. 261 p.
TYPE: Uromycladium simplex McAlpine McAlpine, D. 1905. A new genus of Uredineae-Uromycladium.
(Lectotype) Ann. Mycol. 3:303-323.
On: Acacia pycnantha Benth. (Fabaceae) McAlpine, D. 1906. The rusts of Australia. Gov. Print.,
Type locality: Australia Melbourne. 349 p.

FIGURE 77. Uromycladium. U. aritimum McAlpine. A. Urediniospores. B. Teliospores.

 ENDORAECIUM Hodges & D.E. Gardner
Mycologia 76:332-349 (339). 1984.

FAMILY: Pileolariaceae genus closely related to the genus Atelocauda and the
most common of the several obviously related rusts
DESCRIPTION: on endemic Acacia spp. in Hawaii (Hodges and
Spermogonia associated with telia, Gardner 1984). Both species are on Acacia koa.
subepidermal, conical with flat hymenium, Group Together with several related Atelocauda species on
VI (type 5). Telia covering surface of phyllodes, the same host, they exhibit interesting life cycle
paraphysate, paraphyses slender and with the tip modifications and speciation (see Atelocauda).
rounded and inflated; teliospores pedicellate, 1-
celled, resembling aeciospores of Atelocauda; basidia REFERENCES:
external. Gardner, D.E. 1981. Nuclear behavior and clarification of the
spore stages of Uromyces koae. Can. J. Bot. 59:939-946.
TYPE: Endoraeciumacaciae Hodges & D.E. Gardner Gardner, D.E. 1991. Atelocauda angustiphylloda n. sp.,a microcyclic
rust of Acacia koa in Hawaii. Mycologia 83:650-653.
On: Acacia koa Gray (Fabaceae)
Gardner, D.E.; Hodges, C.S., Jr. 1985. Spore surface morphology
Type locality: Kauai, Hawaii of Hawaiian Acacia rust fungi. Mycologia 77:575-586.
Hodges, C.S., Jr.; Gardner, D.E. 1984. Hawaiian forest fungi. IV.
REMARKS: Rusts on endemic Acacia species. Mycologia 76:332-349.
Two species, E. acaciae and E. hawatiense Hodges
& D.E. Gardner, are known. This is an endocyclic

FIGURE 78. Endoraecium. A teliospore of E. hawatiense

Hodges & D.E. Gardner (reprinted with
permission, from Hodges and Gardner 1984).

L222
 ALLOTELIUM Syd.
Ann. Mycol. 37:275-438 (312-313). 1939.

FAMILY: Raveneliaceae Allotelium, although distinct. Allotelium closely re-

sembles Diabole and Dicheirinia and undoubtedly
DESCRIPTION: belongs to the same family, Raveneliaceae, but dif-
Spermogonia, aecia, and uredinia unknown. fers in several important morphological characters;
Telia subcuticular, covered by the cuticle and mem- for example, Allotelium has a prominent peridium,
branous peridium for a long time; teliospores 4- whereas the other two genera do not. Allotelium
celled, cells arranged rectangularly, pedicel with is considered synonymous with Diorchidium by
apical cells; basidia undoubtedly external. Cummins and Hiratsuka (1983) and with Dicheirinia
by Hennen et al. (1998).
TYPE: Allotelium mirabile Syd.
On: Calliandra trinervia Benth. (Fabaceae) REFERENCES:
Type locality: Ecuador Berndt, R. 1995. Diabolidium, a new genus of rust fungi
(Uredinales). Mycotaxon 51:263-271.
REMARKS: Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
The type species, A. mirabile, and another species fungi. Rev. ed. APS Press, St. Paul, MN. 152 p-
(A. calliandrae (R. Berndt) Y. Hirats. comb. nov.) are Hennen, J.F.; Sotao, H.M.P.; Hennen, M.M.W. 1998. The genus
known. The second species was described as Dia- Diorchidium in the Neotropics. Mycologia 90:1079-1086.
Sydow, H. 1939. Fungi aequatoriensis. Ann. Mycol. 37:275-438.
bolidium calliandrae R. Berndt (Berndt 1995), but we
consider it to be closely related to the type species of

FIGURE 79. Allotelium. Teliospores of A. calliandrae (R. Berndt) Y. Hirats. (reprinted with permission, from
Berndt 1995).

123
 ANTHOMYCES Dicetel
Hedwigia 38:248-259 (253-254). 1899.

FAMILY: Raveneliaceae REMARKS:

The single species occurs in Brazil on an unidenti-
DESCRIPTION: fied legume. The genus differs from Anthomycetella
Spermogonia and aecia unknown. Uredinia sub- because the latter has multihyphal pedicels. The
epidermal, erumpent, with basally united periph- teliospores with simple pedicels with apical cells are
eral paraphyses, Malupa-type; urediniospores borne like Dicheirinia, but Dicheirinia has ornamented
singly on pedicels, echinulate, pores obscure. Telia teliospores and Anthomyces has smooth teliospores.
similar to the uredinia; teliospores 1-celled, firmly If Anthomyces proves to have type 7 spermogonia
united laterally to forma radial cluster of 3 to several and uredinoid aecia there would be little to separate
spores, wall pigmented, germ pore | in each spore, the genera.
apical, pedicels with apical cells of the same number
as the spores (or at least as the peripheral ones), REFERENCE:
basally simple; germination occurs without dor- Dietel, P. 1899. Uredineae brasilienses a cl. E. Ule lectae. Il.
mancy; basidia external. Hedwigia 38:248-259,

TYPE: Anthomyces brasiliensis Dietel

On: Caesalpinia sp. or Piptadenis sp. (Fabaceae)
Type locality: Brazil

FIGURE 80. Anthomyces. Paraphyses and a group of teliospores of

A. brasiliensis Dietel.
 ANTHOMYCETELLA H. & P. Syd.
Ann. Mycol. 14:353-375 (353-354). 1916.

FAMILY: Raveneliaceae REMARKS:

The type is the only known species. Thirumalachar
DESCRIPTION: (1947) demonstrated that the basal layer does not
Spermogonia and aecia unknown. Uredinia sub- consist of teliospores, but of sporogenous cells, each
epidermal, erumpent, with peripheral paraphyses, capable of producing one or more teliospores.
Calidion-type; urediniospores borne singly on pedi-
cels, echinulate, pores obscure. Telia as the uredinia; REFERENCES:
teliospores 1-celled, firmly united to form a radial Sydow, H.; Sydow, P. 1916. Weitere Diagnosen neuer philip-
cluster 2 to several spores wide, pigmented, over- pinischer Pilze. Ann. Mycol. 14:353-375.
laid by a pale layer and subtended by short spore- Thirumalachar, M.J. 1947. Brief notes on the genera Stereostratum
Magn. and Anthomycetella Syd. Mycologia 39:334-340.
like basal cells, germ pore 1 in each spore, apical,
pedicel compound, corresponding to the number of
basal cells; germination occurs without dormancy;
basidia external.

TYPE: Anthomycetella canarii H. & P. Syd.

On: Canarium villosum Craib. (Burseraceae)
Type locality: Philippines

FIGURE 81. Anthomycetella. Teliospores and a paraphysis of

A. canarit H. & P. Syd.
 APRA J.F. Hennen & F.O. Freire
Mycologia 71:1053-1057 (1053). 1979.

FAMILY: Raveneliaceae REMARKS:

Only the type species is known. Apra differs from
DESCRIPTION: Dicheirinia and Diorchidiella in having free teliospores;
Spermogonia splitting the epidermis, Group VI in the latter two genera, the teliospores develop on
(type 7). Aecia subepidermal, deep-seated, becom- separate apical stalk cells but are adherent to others
ing erumpent, Aecidium-type, with peridium; aecio- on the same pedicel. Diabole, also on Mimosa, has
spores catenulate. Uredinia morphologically two free teliospores on a single apical cell and with
identical with aecia. Telia subepidermal, with a one or more pairs on a common pedicel.
marginal layer of compact hyphae and collapsed
palisade cells, becoming erumpent; teliospores REFERENCE:
1-celled, borne in laterally free pairs on a common Hennen, J.F; Freire, FO. 1979. Apra,a new genus of Uredinales on

pedicel that has 2 apical cells, 1 for each spore, germ Mimosa from Brazil. Mycologia 71:1053-1057.

pore not seen; basidia doubtless external.

TYPE: Apra bispora J.F. Hennen & F.O. Freire

On: Mimosa micracantha Benth. (Fabaceae)
=M. rufescens Benth. var. amnis-nigri
Barneby
Type Locality: Brazil

FIGURE 82. Apra. A. bispora J.P. Hennen & F.O. Freire. A. A telium. B. Teliospores.

126
 CUMMINSINA Petr.
Sydowia 9:474-477 (474). 1955.

FAMILY: Raveneliaceae REMARKS:

The type is the only known species. The genus
DESCRIPTION: has the features of Anthomycetella, but produces
Spermogonia and aecia unknown. Uredinia sub- more complex (often more so than illustrated) spore
epidermal, erumpent, with peripheral paraphyses, heads that appear like a bundle of Phragmidium
Calidion-type; urediniospores borne singly on pedi- teliospores or a giant Alternaria spore. The relation-
cels, echinulate. Telia subepidermal, erumpent; ship of Cumminsina to other genera is obscure.
teliospores consisting of chains of laterally adherent
cells forming a club-shaped, pigmented head, REFERENCE:
pedicel simple basally but with apical cells equal in Petrak, F. 1955. Cumminsina n.gen., eine neue Gattung der
number to the chains of cells, germ pores not seen; Uredineen. Sydowia 9:474—447,
germination unknown, but basidia undoubtedly
external.

TYPE: Cumminsina clavispora Petr.

On: Grewia nenensis (author’s name not found)
(Tiliaceae)
Type locality: Angola

FIGURE 83. Cumminsina. C. clavispora Petr. A. A teliospore. B. Uredinial paraphyses. C. Urediniospores.

 CYSTOMYCES Syd.
Ann. Mycol. 24:283-426 (290). 1926.

FAMILY: Raveneliaceae REMARKS:

The type is the only known species and it is
DESCRIPTION: microcyclic. Cystomyces is similar to Ravenelia, but
Spermogonia subepidermal, Group VI (type 5). has only 3-celled teliospores and a simple pedicel.
Aecia and uredinia not produced in the type. Telia The pedicel attaches to the cysts, which are therefore
subepidermal, erumpent;, teliospores mostly 3- similar to the apical cells of Dicheirinia. Lopez-
celled by vertical septa, thus radially arranged, Franco and Hennen (1989) reported a germ slit near
deeply pigmented, opaque, subtended by 3 (2 the inner walls but no evidence of germ tube exit
according to Hennen) colorless, hygroscopic cysts through the slit was observed.
surmounting a simple pedicel, 1 germ pore (or germ
slit) in each cell; germination unknown but basidia REFERENCES:
undoubtedly external. Cummins, G.B. 1978. Rust fungi on legumes and composites in
North America. Univ. Arizona Press, Tucson, AZ. 424 p.

TYPE: Cystomyces costaricensis Syd. Lopez-Franco, R.M.; Hennen, J.F. 1989. The slit and other sorts of
On: Unidentified Fabaceae, probably metabasidia exits in rust fungus probasidia. Am. Mycol. Soc.
Newsl. 40:37.
Lonchocarpus sp. (L. guatemalensis
Bentam. according to Cummins (1978)) Sydow, H. 1926. Fungi in itinere costaricensi collecti. Ann. Mycol.
24:283-426.
Type locality: Costa Rica

FIGURE 84. Cystomyces. A teliospore of

C. costaricensis Syd.
 DIABOLE Arthur
Bull. Torrey Bot. Club 49:189-196 (194). 1922.

FAMILY: Raveneliaceae REMARKS:

Only the type species is known. It occurs in Cuba,
DESCRIPTION: Central America, Mexico, and Brazil on Mimosa.
Spermogonia subcuticular, Group VI (type 7). Diabole is one of a few genera that have subcuticular
Aecia and uredinia unknown. Telia subcuticular, telia. The genus presumably is related to Ravenelia or
erumpent; teliospores paired on each cell at the Dicheirinia, or perhaps Uromycladium, where Arthur
pedicel apex and 1-3 such cells borne on each pedi- first placed it (Cummins 1935).
cel, thus 2 to 6 1-celled spores on a pedicel, wall
pigmented, verrucose, germ pores uncertain, per- REFERENCES:
haps in a basal pale area of each spore; germination Arthur, J.C. 1922. New species of Uredinales. XIV. Bull. Torrey
unknown, but the basidia undoubtedly external. Bot. Club 49:189-196.
Cummins, G.B. 1935. The genus Dicheirinia. Mycologia
PAE AS IANS),
TYPE: Diabole cubensis (Arthur & J.R.Johnst.) Arthur
=Uromycladium cubense Arthur & J.R. Johnst.
On: Mimosa sp. (Fabaceae)
Type locality: Cuba

FIGURE 85. Diabole. A telium and teliospores of D. cubensis (Arthur & J.R. Johnst.) Arthur.
 DICHEIRINIA Arthur
N. Am. Flora 7:83-160 (147). 1907.

FAMILY: Raveneliaceae REMARKS:

There are 12 species in the genus, all on Fabaceae.
DESCRIPTION: Dicheirinia canariensis Urries (Canary Islands), D.
Spermogonia subcuticular, Group VI (type 7). trispora Cummins (Mauritius), D. maderensis Gjaerum
Aecia subepidermal, erumpent, Uredo-type; aecio- (Madeira Islands), and D. viennotii B. Huguenin
spores borne singly on pedicels. Uredinia subepi- (New Caledonia) are the extra-American species.
dermal, erumpent, Calidion-type, similar to the aecia Hennen et al. (1998) consider Allotelium to be a syn-
but without spermogonia, mostly with peripheral onym of Dicheirinia, not of Diorchidium, as indicated
paraphyses that are often branched and elaborate, by Cummins and Hiratsuka (1983), but we treat
Calidion-type; urediniospores borne singly on pedi- them as separate genera here.
cels, as the aeciospores, pores equatorial or basal.
Telia subepidermal, erumpent, often surrounded by REFERENCES:
peripheral paraphyses similar to those of uredinia; Arthur, J.C. 1907. Uredinales. N. Am. Flora 7:83-160.
teliospores 2- to 4-celled by vertical septa, wall pig- Cummins, G.B. 1935. The genus Dicheirinia. Mycologia 27:151—
mented, mostly ornamented with blocklike warts, 159.
the pedicel with 1 apical cell for each cell of the Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
spore, germ pore 1 in each cell; basidia external.
Gjaerum, H.B. 1982. Rust fungi from Madeira. Bol. Mus. Munic.
Funchal 34 (143):5-22.
TYPE: Dicheirinia binata (Berk. & M.A. Curtis) Arthur Hennen, J.F.; Sotao, H.M.P.; Hennen, M.M.W. 1998. The genus
=Triphragmium binatum Berk. & M.A. Curtis Diorchidium in the Neotropics. Mycologia 90:1079-1086.
On: Unknown host plant, now known to be
Erythrina sp. (Fabaceae)
Type locality: Nicaragua

FIGURE 86. Dicheirinia. A. Paraphyses and teliospores of D. binata (Berk. & M.A. Curtis) Arthur. B. A teliospore of
D. manaosensis (Henn.) Cummins.

130
 DIORCHIDIELLA J.C. Linda.
Darwiniana (B. Aires) 11:415-417 (416-417). 1957.

FAMILY: Raveneliaceae (Ferreira and Carvalho 1995). Both occur on Mimosa

species. The genus is separable from Dicheirinia
DESCRIPTION: because of the two germ pores per cell instead of one
Spermogonia subepidermal, Group VI (type 5). as in Dicheirinia; Diorchidiella also lacks prominent
Aecia and uredinia unknown. Telia subepidermal, peripheral paraphyses, which surround the telia of
erumpent; teliospores 2-celled by vertical septum, Dicheirinia.
borne singly on pedicels, the pedicel having 2 apical
cells, 1 for each teliospore cell, wall pigmented, REFERENCES:
germ pores 2 in each cell; basidium external. Ferreira, F.A.; Carvalho, A.O. 1995. Anew species of Diorchidiella
with a spermogonial state. Mycol. Res. 99:885-886.
TYPE: Diorchidiella australis (Speg.) J.C. Lind. Lindquist, J.C. 1957. Diorchidiella, un nuevo género de Uredinales
=Diorchidium australe Speg. (Pucciniaceae-Ravenelieae). Darwiniana (B. Aires) 11:415—-417.

On: Mimosa rocae Lor. & Niederl. (Fabaceae)

Type locality: Argentina

REMARKS:
Two species, the type and D. verlandii F.A.
Ferreira & A.O. Carvalho (from Brazil) are known

FIGURE 87. Diorchidiella. Teliospores of D. australis (Speg.) J.C. Linda.

13)
 DIORCHIDIUM Kalchbr.
in Kalchbr. and Cooke, Grevillea 11:18-27 (26). 1882.

FAMILY: Raveneliaceae described D. quadrifidum Cummins and D. tetra-

sporum Cummins, both with 4-celled teliospores,
DESCRIPTION: thus broadening the concept of the genus. We be-
Spermogonia subcuticular, Group VI (type sas lieve that D. copaifera (Syd.) Cummins & Y. Hirats.
Aecia subepidermal, erumpent, Uredo-type; aecio- (=Sphenospora copaiferae Syd.) belongs here. It is simi-
spores borne singly on pedicels, echinulate, pores lar to D. quadrifidum and D. tetrasporum, but with
equatorial. Uredinia subepidermal, erumpent, Uredo- two-celled teliospores. Diphragmium Boedijn, based
type; urediniospores as the aeciospores. Telia on Diorchidium koordersii Wurth, differs because the
subepidermal, erumpent; teliospores 2- or less com- teliospore pore is apical. We do not consider this to
monly 4-celled by vertical septa, borne singly on be important. Hennen et al. (1998) described four
pedicels, wall pigmented, germ pore 1 in each cell, species from the Neotropics, including a new
apical; basidium external. species, D. amapaensis J.F. Hennen & Sotao, with
excellent illustrations.
TYPE: Diorchidium woodii Kalchbr. & Cooke
On: Milletia sp. (Fabaceae) REFERENCES:
Type locality: South Africa Bernat, R. 1996. Diorchidium taiwanensis sp. nov. (Uredinales), a
new Diorchidium from Taiwan. Mycotaxon 59:253-257.

REMARKS: Boedijn, K.B. 1959. The Uredinales of Indonesia. Nova Hedwigia

Twelve species (five from Africa, four from South 1:463-496.
Cummins, G.B. 1960. Descriptions of tropical rusts. IX. Bull.
America, one each from Sri Lanka, Taiwan, and
Torrey Bot. Club 87:31—45.
Indonesia) are reported. The number is uncertain M.M.W. 1998. The genus
Hennen, J.E.; Sotao, H.M.P.; Hennen,
because species of Puccinia with more or less vertical Diorchidium in the Neotropics. Mycologia 90:1079-1086.
septa have been described as Diorchidium spp. The Kalchbrenner, K.; Cooke, M.C. 1882. Fungi Macowanianl.
spermogonia of Puccinia are type 4 and the telio- Grevillea 11:18-27.
spores rarely have vertical septa. Cummins (1960)

FIGURE 88. Diorchidium. A. Teliospores of D. woodii Kalchbr. & Cooke. B. A paraphysis, a urediniospore, and
teliospores of D. tetrasporum Cummins. C. Paraphyses, teliospores, and a urediniospore of D. copaifera
(Syd.) Cummins & Y. Hirats. D. Teliospores of D. puiggarii Speg. E. Urediniospores of D. puiggarit.
(D, E reprinted with permission, from Hennen et al. 1998).
 ESALQUE J.F. Hennen, Figueiredo & A.A. Carvalho
Mycologia 92:312-316 (315). 2000.

FAMILY: Raveneliaceae confirmed as a species of Hapalophragmium, not

Triphragmium, as stated in Cummins and Hiratsuka
DESCRIPTION:
(1983). However, a species described as Triactella
Spermogonia and aecia unknown. Uredinia sub- holwayi H.S. Jacks. cannot be placed in Hapalo-
epidermal, Uredo-type; urediniospores pedicellate,
phragmium because of the arrangement of the cells in
1-celled, walls echinulate with 2 or more germ
the teliospores. Therefore Hennen et al. (2000)
pores. Telia subepidermal; teliospores often mixed
created a new genus, Esalque, to accommodate this
in uredinia, each with 3 cells, 1 basal (proximal),
species. This genus is similar to another recently
which is attached to the pedicel, and 2 apical (distal)
established genus, Hennenia (Buritica 1995).
cells, germ pores not seen; basidia undoubtedly
external. REFERENCES:
Buritica, P. 1995. Nuevos registros de Uredinales para la flora
TYPE: Esalque holwayi (HLS. Jacks.) J.F. Hennen, Colombiana y paises adyacentes. Rev. Acad. Colomb. Cienc.
Figueiredo & A.A. Carvalho Exactas Fis. Nat. 19:463-467.
=Triactella holwayi H.S. Jacks. Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
fungi. Rev. ed. APS Press, St. Paul, MN. 152 p-
=Hapalophragmium holwayi (H.S. Jacks.) Syd.
Hennen, J.F.; Figueiredo, M.B.; Carvalho, A.A. de. 2000. Esalque
On: Caesalpinia sp. (originally reported
holwayi gen. et comb. nov., a rust of Brazilian ironwood
erroneously as Cassia sp.) (Fabaceae) (Caesalpinia species). Mycologia 92:312-316.
Type locality: Rio de Janeiro, Brazil

REMARKS:
Hennen et al. (2000) pointed out that the type
species of the genus Triactella, T. pulchrum Racib., is

FIGURE 89. Esalque. E. holwayi (H.S. Jacks.) J.F. Hennen, Figueiredo & A.A. Carvalho. A. A teliospore, urediniospores,
and a paraphysis. B. Teliospores (courtesy Dr. J.F. Hennen). C. Teliospores and urediniospores (courtesy
Dr. D.R. Santiago).
 HAPALOPHRAGMIUM H. & P. Syd.
Hedwigia 40:62-65 (64). 1901.

FAMILY: Raveneliaceae All are autoecious, and most are macrocyclic. We

treat Hapalophragmium and Hapalophragmiopsis
DESCRIPTION: Thirum. as synonymous, as does Monoson (1977) and
Spermogonia Group VI (type 7, subcuticular, or Lohsomboon et al. (1992), but Laundon (1975) cites
less commonly type 5, subepidermal). Aecia subepi- possible reasons for keeping them separate, as did
dermal, erumpent, mostly with peripheral pa- Thirumalachar (1950). Lohsomboon et al. (1992) pro-
raphyses, Calidion- or Uredo-type; aeciospores borne vides extensive keys to species as well as detailed
singly on pedicels, echinulate. Uredinia similar to descriptions and excellent figures.
aecia, Calidion- or Uredo-type; urediniospores borne
singly on pedicels, echinulate, pores equatorial. REFERENCES:
Telia subepidermal, erumpent; teliospores borne Laundon, G.F. 1975. Taxonomy and nomenclature notes on

singly on simple pedicels, triquetrous, 3-celled, with Uredinales. Mycotaxon 3:133-161.

Lohsomboon, P.; Kakishima, M.; Ono, Y. 1992. A monograph of
2 cells basal and 1 above, wall pigmented, smooth or
Hapalophragmium. Mycol. Res. 96:461-472.
sculptured, germ pore 1 in each cell; basidia external. Monoson, H.L. 1977. A synopsis of the genus Hapalophragmium
(Uredinales). Mycologia 69:21-33.
TYPE: Hapalophragmium derridis H. & P. Syd. Sydow, H.; Sydow, P. 1901. Hapalophragmium, ein neues Genus
On: Derris uliginosa Benth. (Fabaceae) der Uredineen. Hedwigia 40:62-65.
Type locality: Tanzania and Madagascar Thirumalachar, M.J. 1950. Some noteworthy rusts. III. Mycologia
(syntypes) 42:224-232.

REMARKS:
This is a genus of 15 species, all on legumes in
tropical Asia and Africa (Lohsomboon et al. 1992).

FIGURE 90. Hapalophragmium. A. A sorus with teliospores and a urediniospore of H. derridis H. & P. Syd.
B. A paraphysis, a teliospore, and urediniospores of H. ornatum Cummins.
 HENNENIA Buritica
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 19:463-468 (465-466). 1995.

FAMILY: Raveneliaceae tropical areas on Annonaceae and Fabaceae. The

type species of this genus is unique in exhibiting two
DESCRIPTION: kinds of teliospore formation: one in which the telia
Spermogonia subepidermal, flat, Group VI (type occur in the same area as the aecia, and another type
5). Aecia cup-shaped, with peridium, Aecidium- in which teliospores occur on large areas of the
type; aeciospores catenulate, verrucose, with some leaves (undersides) and single teliospores emerge
refractive granules. Uredinia unknown. Telia through the stomata. The second type of teliospore
subepidermal, erumpent, or telia not formed and formation gives a coffee-brown color to the
individual teliospores emerge through stomata sporulating areas. This genus is considered to be
(suprastomatal); teliospores pedicellate, tricellular closely related to the newly described genus Esalque
with 2 apical cells mounted on 1 basal cell, or 3 cells J.E Hennen et al. (Hennen et al. 2000), which has
in a row, one below the other, germ pore obscure. similar teliospore morphology.

TYPE: Hennenia ditelia Buritica REFERENCES:

On: Annona sp. (Annonaceae) Buritica, P. 1995. Nuevos registros de Uredinales para la flora
Type locality: Colombia Colombiana y paises adyacentes. Rev. Acad. Colomb. Cienc.
Exactas Fis. Nat. 19:463-468.
REMARKS: Hennen, J.F; Figueiredo, M.B.; Carvalho, A.A. de. 2000. Esalque
holwayi gen. et comb. nov., a rust of Brazilian ironwood
Only the type species is known. Buritica (1995)
(Caesalpinia species). Mycologia 92:312-316.
considers this genus related to Hapalophragmium
and Sphaerophragmium, which commonly occur in

FIGURE 91. Hennenia. Teliospores of H. ditelia Buritica

(courtesy of Dr. J.F. Hennen).
 KERNKAMPELLA Rajendren
Mycologia 62:837-843 (839). 1970.

FAMILY: Raveneliaceae monograph of Ravenelia and its possible segregates.

Ono (1987) examined the specimens identified as K.
DESCRIPTION: breyniae-patentis (Mundkur & Thirum.) Rajendren
Spermogonia subcuticular, Group VI (type 7). and K. breyniae (H. & P. Syd.) Rajendren and con-
Aecia subepidermal, erumpent, with peridium, cluded that they are conspecific; he chose K. breyniae
Aecidium-type, or without, Caeoma-type; aeciospores as the type species of the genus. Rajendren (1970b)
catenulate, verrucose. Uredinia subepidermal, described the repeated production of uredinio-
erumpent, with or without paraphyses, Uredo-type; spores by basal cells, with new spores arising within
urediniospores borne singly on pedicels, echinulate, the pedicel of the preceding spore, a method
pores mostly equatorial. Telia subepidermal, erum- reported by Hennen and Figueiredo (1979) for the
pent; teliospores 1-celled, each cell with a central anamorphic genus Intrapes. It is not known whether
appendage, strongly adherent laterally, forming this method applies to all species of Kernkampella.
discoid heads subtended by a patelliform layer of All species occur on Euphorbiaceae.
cells, below which are hygroscopic cysts, germ pore
1 in each cell, pedicel compound; basidia external. REFERENCES:
Bagyanarayana, G.; Ramachar, P. 1985. Spermogonial and aecial
stages of Kernkampella kirganelliae. Trans. Br. Mycol. Soc.
TYPE: Kernkampella breyniae (H. & P. Syd.) 84:171-173.
Rajendren (Lectotype, Ono 1987) Hennen, J.F; Figueiredo, M.B. 1979. Intrapes,a new genus of fungi
=Ravenelia breyniae H. & P. Syd. imperfecti (Uredinales) from Brazilian Cerrado. Mycologia
=Kernkampella breyniae-patentis (Mundkur & 71:836-840.
Thirum.) Rajendren Laundon, G.F. 1975. Taxonomy and nomenclature notes on
Uredinales. Mycotaxon 3:133-161.
On: Breynia rhamnoides Blume (Euphorbiaceae)
Ono, Y. 1987. Identity of Kernkampella breyniae-patentis with K.
Type locality: Pusa, India breyniae and its occurrence in Japan. Trans. Mycol. Soc. Jpn.
28:237-243.
REMARKS: Rajendren, R.B. 1970a. Kernkampella: a new genus in the
This genus of eight species is distinguished from Uredinales. Mycologia 62:837-843.
Ravenelia because of the patelliform layer of cells Rajendren, R.B. 1970b. Cytology and developmental morphol-
between the spores and the cysts. Whether this ogy of Kernkampella breyniae-patentis and Ravenelia hobsoni.
Mycologia 62:1112-1121.
structure warrants segregation of a genus or of a sec-
Tyagi, R.N.S. 1974. A critical account of the Kernkampella. Indian J.
tion of Ravenelia is a matter of opinion. Laundon Mycol. Plant Pathol. 3:63-66.
(1975) and Ono (1987) accepted the genus; Tyagi and Tyagi, R.N.S.; Prasad, N. 1972. The monographic studies on ge-
Prasad (1972) and Tyagi (1974) did not. We favor nus Ravenelia occurring in Rajasthan. Indian. J. Mycol. Plant
retaining Kernkampella in the absence of a modern Pathol. 2:108-135.

FIGURE 92. Kernkampella. K. appendiculata (Lagerh. & Dietel) Laundon.

A. A teliospore. B. A paraphysis.
 LIPOCYSTIS Cummins
Bull. Torrey Bot. Club 64:39-44 (39). 1937.

FAMILY: Raveneliaceae REMARKS:

Only the type species is known. It is autoecious
DESCRIPTION: and occurs in the West Indies on Mimosa. It is one of
Spermogonia subcuticular, Group VI (type 7). the few genera that have all spore forms sub-
Aecia subcuticular, erumpent, Uredo-type; aecio- cuticular. There is little to indicate relationship ex-
spores borne singly on pedicels, echinulate, pores cept the host and the spermogonia, both of which
equatorial. Uredinia subcuticular, erumpent, Uredo- suggest Ravenelia. Lipocystis differs from Ravenelia
type; urediniospores as the aeciospores. Telia sub- by the absence of cysts.
cuticular, erumpent; teliospores 1-celled, borne
singly on short, broad pedicels, laterally free or REFERENCE:
tending to adhere in small groups, wall pigmented, Cummins, G.B. 1937. Descriptions of tropical rusts. Bull. Torrey
germ pore 1; basidium external. Bot. Club 64:39-44.

TYPE: Lipocystis caesalpiniae (Arthur) Cummins

=Ravenelia caesalpiniae Arthur
On: Mimosa ceratonia L. (Fabaceae)
Type locality: West Indies

FIGURE 93. Lipocystis. L. caesalpiniae (Arthur) Cummins. A. Urediniospores. B. A paraphysis. C. Teliospores.

37
 NYSSOPSORA Arthur
Rés. Sci. Congr. Int. Vienne. p. 331-348 (342). 1906.

FAMILY: Raveneliaceae al. 1990). Information about life cycles is scarce. Six
species are known only in the telial state, two
DESCRIPTION: species have uredinia and telia, and only one species
Spermogonia unknown. Aecia, when present (in (N. cedrelae (Hori) Tranzschel) is known to have
N. cedrelae (Hori) Tranzschel), similar to uredinia. aecia (Uredo-type), confirmed by inoculation experi-
Uredinia subepidermal, erumpent, Uredo-type; ments (Kakishima et al. 1984). The strongly spinose
urediniospores borne singly on pedicels, echinulate. teliospores are the conspicuous feature and separate
Telia subepidermal, erumpent; teliospores borne the genus from Triphragmium (with one germ pore
singly on pedicels, triquetrous, 3-celled with a basal, per cell) and Triphragmiopsis (with two pores per
pedicellate cell surmounted by 2 cells, wall pig- cell).
mented and bearing conspicuous spines, which are
often apically branched, germ pores 2 in each cell; REFERENCES:
basidia external. Arthur, J.C. 1906. Eine auf die Structur und Entwicklungs-
geschichte begriindete Klassifikation der Uredineen. Reés. Sci.
TYPE: Nyssopsora echinata (Lév.) Arthur Congr. Int. Vienne. p. 331-348.
Henderson, D.M. 1973. The rust genus Nyssopsora and its host
=Triphragmium echinatum Lév.
relations. Notes R. Bot. Gard. Edinb. 32:217-221.
On: Meum athamanticum Jacq. (Apiaceae) Kakishima, M.; Sato, T.; Sato, S. 1984. Notes on two rust fungi
Type locality: France Pileolaria klugkistiana and Nyssopsora cedrelae. Trans. Mycol.
SOCy |PN. 20000-0008
REMARKS: Lohsomboon, P.; Kakishima, M.; Ono, Y. 1990. A revision of the
There are nine species, four on Araliaceae, two genus Nyssopsora (Uredinales). Mycol. Res. 94:907-922.
on Sapindaceae, and one each on Meliaceae, Pitto- Monoson, H.L. 1974. The species of Triphragmium, Nyssopsora,

sporaceae, and Apiaceae, distributed in Asia, Aus- and Triphragmiopsis. Mycopathol. Mycol. Appl. 52:115-131.
tralia, Europe, and North America (Lohsomboon et

FIGURE 94. Nyssopsora. A. Teliospores of N. echinata (Lév.) Arthur. B. Teliospores of N. clavellosa (Berk.) Arthur.

138
 RAVENELIA Berk.
Gard. Chron. 13:132. 1853.

FAMILY: Raveneliaceae worldwide in warm climates, but absent from

Europe and Australia, and exclusively on Fabaceae.
DESCRIPTION: According to J.F. Hennen (pers. comm.), of 116
Spermogonia mostly subcuticular, Group VI described species of Ravenelia in the Americas, 55
(type 7), in a few species subepidermal (type 5). species are in Mexico, 40 species in Brazil, and 22
Aecia subepidermal or sometimes subcuticular, mostly along the southern border areas of the USA.
erumpent, mostly with pedicellate spores, Uredo- All species reported on Euphorbiaceae belong in
type, or in a few species spores catenulate, with or Kernkampella. J.F. Hennen (pers. comm.) thinks that
without peridium, Aecidium- or Caeoma-type. the host of R. sarmentoi J.C. Lindgq., reported as
Uredinia mostly subepidermal, but sometimes Bulnesia (Zygophyllaceae), is misidentified. Most of
subcuticular, erumpent, Uredo-type; urediniospores the species are autoecious and likely macrocyclic,
borne singly on pedicels, mostly echinulate, often but a few microcyclic species are known. There is no
with paraphyses. Telia subepidermal or sometimes recent comprehensive monograph.
subcuticular, dark brown to nearly black; telio-
spores complex, consisting of discoid heads, typi- REFERENCES:
cally 1 cell thick, but in a few species 2 cells thick in Berkeley, M.J. 1853. (No title.) Gard. Chron. 13:132.
the center part of the spore head due to transverse or Cummins, G.B. 1978. Rust fungi on legumes and composites in
oblique septa, cells strongly adherent laterally, up to North America. Univ. Arizona Press, Tucson, AZ. 424 p.

80 or more cells in a spore head, pedicellate; surface Doidge, E.M. 1927. South African rust fungi. Bothalia 2(1a):1—228.

of teliospore heads smooth or sculptured; teliospore Kakishima, M.; Lohsomboon, P.; Sato, S. 1987. Life cycle and
morphology of Ravenelia japonica. Trans. Mycol. Soc. Jpn.
heads subtended by colorless, hygroscopic cysts, 28:189-196.
pedicels fasciculate; germ pore 1 in each cell; basidia Lindquist, J.C. 1954. Las especies Argentinas de Ravenelia. Rev.
external. Fac. Agron. 30:103-128.
Rezende, D.V. 1999. Taxonomia de Uredinales em plantas da
TYPE: Ravenelia glandulosa Berk. & M.A. Curtis familia Leguminosae do Cerrado Brasiliero. Ph.D. thesis,
(Lectotype) Univ. Brasilia, Brasilia. 262 p. [Originally published as
Santiago, D.V.R. 1999. ]
=R. epiphylla (Schwein.) Dietel
Savile, D.B.O. 1989. Raveneliaceae revisited. Can. J. Bot. 67:2983-
On: Tephrosia virginiana (L.) Pers. (Fabaceae) 2994,
=Cracca virginiana L. Sydow, P.; Sydow, H. 1915. Monographia Uredinearum. 3:224—
Type locality: South Carolina, USA 310.
Tyagi, R.N.S.; Prasad, N. 1972. The monographic studies on ge-
REMARKS: nus Ravenelia occurring in Rajasthan. Indian J. Mycol. Plant
Ravenelia is the third largest genus after Puccinia Pathol. 2:108-127.

and Uromyces, with about 200 species distributed

FIGURE 95. Ravenelia. A. Dorsal view of a teliospore of R. fragrans Long var.

fragrans. B. Side view of a teliospore head of R. echinata Lagerh.
FIGURE 96. Ravenelia. Teliospores and urediniospores of Ravenelia species. A, top surface of teliospore heads; B, side
view of teliospore ornamentation; C, urediniospores: 1, R. hermosa Cummins & J.W. Baxter; 2, R. mexicana
Tranzschel; 3, R. holwayi Dietel (A and B) and R. cumminsii J.W. Baxter; 4, R. stevensii Arthur (A and B)
and R. spegazziniana J.C. Lindq.; 5, R. bella Cummins & J.W. Baxter; 6, R. corbula J.W. Baxter.

140
FIGURE 97. Ravenelia. A. A urediniospore of R. lonchocarpi Lagerh. & Dietel. B. Urediniospores of R. pileolarioides
H. & P. Syd. C. Teliospores of R. bezerrae Dianese, Medeiros & Furlanetto. D. Bottom view of a teliospore
of R. bezerrae. E. Teliospores of R. dieteliana Henn. F. Side view of a teliospore of R. dieteliana. (Reprinted
with permission, from Rezende 1999.)

141
 SPHAEROPHRAGMIUM Magnus
Ber. Dtsch. Bot. Ges. 9:118-124 (121). 1891.

FAMILY: Raveneliaceae REMARKS:

There are 16 known species; 13 are on Fabaceae
DESCRIPTION: and 3 on Annonaceae. They occur circumglobally in
Spermogonia subepidermal, Group VI (type 5): warm to tropical areas (Lohsomboon et al. 1994;
Aecia subepidermal, erumpent, with peridium, Alcorn and Walker 1996; Monoson 1974).
Aecidium-type; aeciospores catenulate, verrucose.
Uredinia subepidermal, erumpent, Calidion-type; REFERENCES:
urediniospores borne singly on pedicels, echinulate, Alcorn, J.L.; Walker, J. 1996. Sphaerophragmium quadricellulae sp.
nov. (Mycota, Uredinales) on Acacia in Australia, with brief
pores equatorial, the spores often asymmetrical and notes on some related rust genera. Mycotaxon 58:325-330.
reniform, mostly with peripheral paraphyses. Telia Lohsomboon, P.; Kakishima, M.; Ono, Y. 1994. A monograph of
subepidermal, erumpent; teliospores borne singly Sphaerophragmium (Uredinales). Mycol. Res. 98:107-119.
on pedicels, 4- to several-celled by vertical and Magnus, P. 1891. Einige Beobachtungen zur naheren Kenntniss
transverse septa, usually more or less globoid, wall der Arten von Diorchidium und Triphragmium. Ber. Dtsch. Bot.
pigmented, with simple or often apically furcate Ges. 9:118-124.
spines or projections, germ pores obscure, uncer- Monoson, H.L. 1974. The genus Sphaerophragmium. Mycologia
66:791-802.
tain; basidia external.

TYPE: Sphaerophragmium acaciae (Cooke) Magnus

=Triphragmium acaciae Cooke
On: Albizia lebbeck (L.) Benth. (Fabaceae)
=Acacia lebbeck (L.) Willd.
Type locality: India

FIGURE 98. Sphaerophragmium. A. Paraphyses, urediniospores, and teliospores of S. acaciae

(Cooke) Magnus. B. A teliospore of S. artabotrydis Doidge.
 SPHENOSPORA Dietel
Ber. Dtsch. Bot. Ges. 10:57—63 (63). 1892.

FAMILY: Raveneliaceae Sphenospora copaiferae Syd. was described on Copaifera

of the Fabaceae, but we consider this species to be-
DESCRIPTION: long to Diorchidium. Sphenospora dalbergiellae Vienn.-
Spermogonia and aecia unknown. Uredinia sub- Bourg. on Dalbergiella sp. (Fabaceae) from Guinea,
epidermal, becoming erumpent, but often slowly, Africa, resembles S. copaiferae, but without the oily
Uredo-type; urediniospores borne singly on pedi- gelatinous matrix, and should also belong to
cels, echinulate, germ pores equatorial or obscure. Diorchidium (Hennen et al. 1998). An abundance of
Telia subepidermal, erumpent as waxy-oily cush- orange-colored oil and an apparently gelatinous
ions when moist, hard when dry; teliospores borne matrix is characteristic of Sphenospora.
singly on pedicels, 2-celled by vertical septum, Three species occur on orchids and are often
mostly narrowly ellipsoid or conical, wall pale to intercepted by the United States Plant Quarantine
colorless, smooth, germ pore 1 in each cell if differ- Service in air cargo to the US.
entiated, apical; germination occurs without dor-
mancy; basidia external. REFERENCES:
Berndt, R. 1996. Comparative studies on the teliospore mor-
TYPE: Sphenospora pallida (G. Winter) Dietel phology and ontogeny of Spumula serispora, spec. nova, and
=Diorchidium pallidum G. Winter Ravenelia texensis (Uredinales, Raveneliaceae). Plant Syst.
On: Possibly Dioscorea sp. (Dioscoreaceae) Evol. 200:79-88.
Dietel, P. 1892. Zu Beurtheilung der Gattung Diorchidium. Ber.
Type locality: Bolivia
Dtsch. Bot. Ges. 10:57-63.
Hennen, J.F.; Sotao, H.M.P.; Hennen, M.M.W. 1998. The genus
REMARKS:
Diorchidium in the Neotropics. Mycologia 90:1079-1086.
Six species are known and five occur in tropical Linder, D.H. 1944. Anew rust of orchids. Mycologia 36:464468.
America on monocots. Sphenospora xylopiae J.M. Yen, J.-M.; Sulmont, P. 1969. Les Urédinées du Gabon. II. Un nou-
Yen & Sulmont, on Annonaceae in Gabon, is the veau Sphenospora parasite du Xylopia: Sphenospora xylopiae
extra-American species (Yen and Sulmont 1969). (nov. sp.). Bull. Soc. Mycol. Fr. 85:351-353.

FIGURE 99, Sphenospora. A. A telium of S. smilacina Syd. B. Teliospores and urediniospores of S. saphena Cummins.

143
 SPUMULA Mains
Mycologia 27:638-641. 1935.

FAMILY: Raveneliaceae complicated heads, the former with mostly 2-10

teliospores and the latter much more variable with
DESCRIPTION: up to 20. The teliospore heads have simple pedicels,
Spermogonia subcuticular, Group VI (type 7). but the cysts do not seem to be hygroscopic and are
Aecia subepidermal, erumpent, with peridium, not always of the same number as the spores in a
Aecidium-type; aeciospores catenulate, verrucose. head. Both species resemble Dicheirinia as much as
Uredinia subepidermal, erumpent, Uredo-type; they do Spumula. Berndt (1996) compared the telio-
urediniospores borne singly on pedicels, echinulate, spore heads of two species of Spumula, including the
germ pores obscure. Telia subepidermal, erumpent, type species, S. quadrifida Mains. and Ravenelia
teliospores 1-celled, laterally united in 3- to several- texensis Ellis & Galloway. He concluded that the
celled discoid, pigmented heads subtended by spore head ontogeny of the three species was simi-
colorless, hygroscopic cysts, germ pore 1 in each lar, but that the origin of the hygroscopic cysts sus-
cell, pedicel simple; basidia external. taining the spore heads was different.

TYPE: Spumula quadrifida Mains REFERENCES:

On: Calliandra bijuga Rose (Fabaceae) Baxter, J.W. 1966. New species of rust fungi from Mexico.
Type locality: Jalisco, Mexico Mycologia 58:336-338.
Berndt, R. 1996. Comparative studies on the teliospore mor-
phology and ontogeny of Spumuila serispora, spec. nova, and
REMARKS:
Ravenelia texensis (Uredinales, Raveneliaceae). Plant Syst.
About 8 species occur in tropical America. All Evol. 200:79-88.
species are autoecious and on Fabaceae. Spumula Leén Gallegos, H.M.;Cummins, G.B. 1981. Uredinales (Royas) de
differs from Ravenelia because of the simple pedicel. Mexico. Vol. II. Secretaria Agricultura Recuresos Hidraulicos,
The type and two other species are similar, with four Culiacan, Sinaloa, Mexico. 492 p.
spores per head. Spumula bottomleyae (Doidge) Mains, E.B. 1935. Spumula, a new genus of rusts. Mycologia
Thirum. and S. heteromorpha J.W. Baxter have more 27:638-641.

FIGURE 100. Spumula. A. Teliospores of S. quadrifida Mains. B. A teliospore head of S. heteromorpha J.W. Baxter.

144
 TRIPHRAGMIOPSIS Naumov
Bull. Soc. Mycol. Fr. 30:64-83 (78). 1914 (published 1915).

FAMILY: Raveneliaceae teliospore cell, and from Nyssopsora in the less con-
spicuous spore wall sculpturing (Lohsomboon et al.
DESCRIPTION: 1990). The appearance of the teliospores is more like
Spermogonia and aecia unknown. Uredinia sub- that of Triphragmium than of Nyssopsora. Triphrag-
epidermal, erumpent, with well-developed miopsis and Triphragmium were both accepted and
peridium with catenulate, verrucose uredinio- submerged in Nyssopsora by Majewski (1977). How-
spores, Aecidium-type, or with paraphyses with ever, Monoson (1974) considered Triphragmiopsis
echinulate urediniospores borne singly on pedicels, distinct from the two others. We recognize all three
Uredo-type. Telia subepidermal, erumpent; telio- genera separately in this manual. Lohsomboon et al.
spores borne singly on pedicels, triquetrous, (1990) provided a key to the three known species,
3-celled with a basal pedicellate cell surmounted by descriptions, and illustrations.
2 cells, wall pigmented and verrucose, germ pores
2 or 3 in each cell; basidia external. REFERENCES:
Henderson, D.M. 1973. The rust fungus Nyssopsora and its
TYPE: Triphragmiopsis jeffersoniae Naumov relations. Notes R. Bot. Gard. Edinb. 32:217-221.
On: Jeffersonia dubia (Maxim.) Benth. & Hook. Lohsomboon, P.; Kakishima, M.; Ono, Y. 1990. The genus

(Berberidaceae) Triphragmiopsis (Uredinales). Trans. Mycol. Soc. Jpn. 31:335—

343.
Type locality: Vladivostok, Russia
Majewski, T. 1977. Grzyby (Mycota) Tom 9, Uredinales I.
Panstwowe Wydawnictwo Naukowe, Warsaw. 394 p.; 3 pl.
REMARKS: Monoson, H.L. 1974. The species of Triphragmium, Nyssopsora,
There are three species, the type on Berberidaceae and Triphragmiopsis. Mycopathol. Mycol. Appl. 52:115-131.
in far eastern Russia and Korea, T. isopyri (Moug. & Naumov, N. 1914 (published 1915). Matériaux pour la flore
Nestl.) Tranzschel on Ranunculaceae in Europe, and mycologique de la Russie. Bull. Soc. Mycol. Fr. 30:64-83.
T. laricinum (Chou) EL. Tai on Larix spp. (Pinaceae) Ono, Y.; Azbukina, Z.M.; Kakishima, M.; Kaneko, S. 1997. Distri-
in northern China and far eastern Russia (Ono et al. bution of the larch brown rust caused by Triphragmiopsis
laricinum (Uredinales) in Russian Far East. J. For. Res. 2:227—
1997). The genus differs from Triphragmium in Pele
having two or three germ pores rather than one per

FIGURE 101. Triphragmiopsis. Teliospores of T. jeffersoniae Naumov.

 TRIPHRAGMIUM Link
in Linnaeus, Species Plantarum. 4'" ed. Vol. 6. Pt. 2. p. 84. 1825.

FAMILY: Raveneliaceae The type species of Triactella (Triphragmium pul-

chrum Racib.) on Derris, despite the host being in the
DESCRIPTION: Fabaceae rather than the Rosaceae, is not separable
Spermogonia subcuticular, Group VI (type 7). by substantial characters from Hapalophragmium,
Aecia subepidermal, erumpent, Uredo-type; aecio- thus making the genus Triactella invalid. A species
spores borne singly on pedicels, echinulate. knownas Triactella holwayi H.S. Jacks. differs in mor-
Uredinia subepidermal, erumpent, with pa- phology and cannot be placed in Hapalophragmium,;
raphyses, Uredo-type; urediniospores borne singly thus Hennen et al. (2000) created the new genus
on pedicels, echinulate, pores obscure. Telia Esalque to accommodate this fungus (see Esalque).
subepidermal, erumpent, teliospores borne singly Figures labeled as Triphragmium pulchrum in
on pedicels, triquetrous, 3-celled with 1 basal cell Cummins and Hiratsuka (1983) are actually of
surmounted by 2 cells, wall pigmented, sculptured, Esalque holwayi (H.S. Jacks.) J.-F. Hennen, Figueiredo
especially around the pores, germ pore 1 in each cell; & A.A. Carvalho (=Triactella holwayi HS. Jacks.) from
basidia external. Brazil (Hennen et al. 2000).

TYPE: Triphragmium ulmariae (DC.) Link REFERENCES:

=Puccinia ulmariae DC. Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
On: Spiraea ulmaria L. (Rosaceae) fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
=Filipendula ulmaria (L.) Maxim. Hennen, J.F.; Figueiredo, M.B.; de Carvalho, A.A. 2000. Esalque
holwayt gen et comb. nov., a rust of Brazilian ironwood
Type locality: France
(Caesalpinia species). Mycologia 92:312-316.

REMARKS: Link, H.F. 1925. Species Hyphomycetum et Gymnomycetum. [11

Linnaeus Species Plantarum. 4" ed. Vol. 6. Pt. 2. Gymno-
Four species parasitize the Rosaceae in the mycetes. G.C. Nauk, Berolini. 128 p.
Northern Hemisphere and two are on Fabaceae Lohsomboon, P.; Kakishima, M.; Ono, Y. 1990. A revision of the
(Lohsomboon et al. 1990). The genus differs from genus Triphragmium (Uredinales). Trans. Mycol. Soc. Jpn.
Nyssopsora and Triphragmiopsis in having one pore 31:215-226.
Monoson, H.L. 1974. The species of Triphragmium, Nyssopsora,
per cell and from Hapalophragmium because the
and Triphragmiopsis. Mycopathol. Mycol. Appl. 52:115-131.
pedicel attaches to one of the three teliospore cells
rather than two. Triphragmiopsis and Triphragmium
have teliospores with similar surface sculpture,
which is unlike that of Nyssopsora.

FIGURE 102. Triphragmium. Teliospores and urediniospores of T. ulmariae (DC.) Link.

146
 YPSILOSPORA Cummins
Bull. Torrey Bot. Club. 68:43-48 (47). 1941.

FAMILY: Raveneliaceae was described from Argentina but is also reported

from Costa Rica, Guatemala, Honduras, and Puerto
DESCRIPTION: Rico in anamorph states (Uredo ingae Henn.).
Spermogonia subcuticular, Group VI (type 7). Thirumalachar and Cummins (1948) reduced
Aecia subepidermal, erumpent, with or without Ypsilospora to synonymy with Chaconia because they
peripheral paraphyses, Uredo-type; aeciospores interpreted the pedicels as elongated sporogenous
borne singly on pedicels, echinulate, pores equa- basal cells. Thirumalachar and Mundkur (1949) and
torial. Uredinia Uredo-type, similar to aecia. Telia Cummins (1959) did likewise, but Ono and Hennen
subepidermal, erumpent, with or without (1979) reestablished Ypsilospora, stating that the
paraphyses; teliospores borne in Y-shaped pairs on pedicels of the teliospores develop successively on
a common pedicel, 1-celled, wall pale to colorless, sporogenous cells. We accept their interpretation.
germ pore 1 if differentiated, obscure, basidia exter-
nal; germination occurs without dormancy. REFERENCES:
Cummins, G.B. 1941. New rusts from America and Africa. Bull.
TYPE: Ypsilospora baphiae Cummins Torrey Bot. Club. 68:43-48.
On: Baphia nitida Lodd. (Fabaceae) Cummins, G.B. 1959. Illustrated genera of rust fungi. Burgess
Type locality: Sierra Leone Publ. Co., Minneapolis, MN. 131 p.
Ono, Y.; Hennen, J.F. 1979. Teliospore ontogeny in Ypsilospora
baphiae and Y. africana sp. nov. (Uredinales). Trans. Br. Mycol.
REMARKS:
Soc. 73:229-233.
The type species is microcyclic; the second
Thirumalachar, M.J.; Cummins, G.B. 1948. Status of rust genera
described species, Y. africana Y.Ono & J.F. Hennen is Allotelium, Leucotelium, Edythea, and Ypsilospora. Mycologia
demicyclic. Both parasitize species of Baphia 40:417-422.
(Fabaceae) in western Africa. The third species, Thirumalachar, M.J.; Mundkur, B.B. 1949. Genera of rusts. I.
Y. tucumanensis Hernandez & J.F. Hennen (to be Indian Phytopathol. 2:65-101.
published), is on Inga edulis Martius (Fabaceae) and

FIGURE 103. Ypsilospora. Teliospores of Y. baphiae Cummins.

 FROMMEELLA Cummins & Y. Hirats.
Illus. Genera Rust Fungi. Rev. ed. p. 1-152 (120). 1983.

FAMILY: Phragmidiaceae Hennen 1990). The species of Frommeélla are

autoecious and restricted to Potentilla or close rela-
DESCRIPTION: tives of the Rosaceae. The genus differs from Phrag-
Spermogonia intraepidermal, Group IV (type midium because of the single germ pore per
10). Aecia subepidermal, erumpent, Uredo-type; teliospore cell, but doubtless is closely related.
aeciospores borne singly on pedicels, echinulate, When Arthur described Frommea he designated
pores equatorial, obscure. Uredinia subepidermal, Uredo obtusa F. Strauss as the type. Laundon (1975)
erumpent, Uredo-type; urediniospores similar to has pointed out that U. obtusa is a synonym of
aeciospores. Telia subepidermal, erumpent; telio- Phragmidium potentillae (Pers.) P. Karst., and by
spores borne singly on short pedicels, 3- to several- designated typification, Frommea is a synonym of
celled by transverse septa, wall pigmented, smooth, Phragmidium. However, the fungus described by
germ pore 1 per cell; germinating without dor- Arthur is not a Phragmidium; hence a new generic
mancy; basidia external. name, Frommeélla, was established (Cummins and
Hiratsuka 1983).
TYPE: Frommeélla tormentillae (Fuckel) Cummins &
Y. Hirats. REFERENCES:
=Phragmidium tormentillae Fuckel Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
On: Potentilla erecta (L.) Rauschel (Rosaceae) fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.

=Tormentilla erecta L. Laundon, G.F. 1975. Taxonomy and nomenclature notes on

Uredinales. Mycotaxon 3:133-161.
Type locality: Europe
McCain, J.W.; Hennen, J.H. 1990. Taxonomic notes on Frommeélla
(Uredinales). 1. Mycotaxon 39:249-256.
REMARKS:
Parmelee, J.A. 1986. Frommeélla tormentillae. Fungi Can. 306. Natl.
The type species and two varieties of F. mexicana Mycol. Herb., Biosyst. Res. Cent., Res. Branch, Agric. Can.,
(Mains) J.W. McCain & J.F. Hennen on Duchesnea, Ottawa, ON. 2 p.
from USA and Mexico, are recognized (McCain and

FIGURE 104. Frommeélla. Teliospores

and a urediniospore of
F. tormentillae (Fuckel)
Cummins & Y. Hirats.

148
 GERWASIA Racib.
Bull. Acad. Sci. Crac. Cl. Sci. Math. Nat. 1909:266-279 (270-271). 1909.

FAMILY: Phragmidiaceae Jacks., and takes precedence over it. It is probable

that Mainsia rubi (Dietel & Holw.) H.S. Jacks. is the
DESCRIPTION: same fungus as G. rubi Racib. because both have
Spermogonia within the hypertrophied epider- suprastomatal telia. Most of the 16 or so species of
mis, Group IV (type 6). Aecia intraepidermal, Uredo- Mainsia have been transferred to this genus (Buritica
type; aeciospores borne singly on pedicels, 1994). The species occur on Rosaceae and more espe-
echinulate. Uredinia intraepidermal or subepi- cially on Rubus. All species are autoecious and most
dermal, becoming erumpent, Uredo-type; occur in the Asian and American tropics.
urediniospores borne singly on pedicels, echinulate,
pores obscure, perhaps equatorial. Telia as the REFERENCES:
uredinia or strictly suprastomatal; teliospores borne Buritica, P. 1994. Cambios taxonomicos y nuevos registros de
singly on pedicels, 1-celled, wall pale, germ pore 1; Uredinales de la flora Andina. Inst. Cienc. Nat. Ecol. Univ.
germination occurs without dormancy; basidia Nac. Colomb., Medellin 5:184-188.

external. Jackson, H.S. 1931. The rusts of South America based on the
Holway Collections. II]. Mycologia 23:96-116.
Raciborski, M. 1909. Uber einige javanische Uredineae. Bull.
TYPE: Gerwasia rubi Racib.
Acad. Sci. Crac. Cl. Sci. Math. Nat. 1909:266-279.
On: Rubus sp. (Rosaceae)
Type locality: Java, Indonesia

REMARKS:
Raciborski’s type has not been seen, but there is
no doubt that the genus is the same as Mainsia H.S.

et
J

Pay
fA

[GHEE
A)

= —DPey ine
ee a a ey

FIGURE 105. Gerwasia. A. A uredinium of G. rubi Racib. B. Urediniospores (left) and

teliospores (right) of G. standleyi (Cummins) Cummins & Y. Hirats.
C. A telium of G. standleyi.

149
 GYMNOCONIA Lagerh.
Tromso Mus. Aarsheft. 16:105-152 (142). 1894.

FAMILY: Phragmidiaceae Laundon (1975) suggested applying Gymmoconia to

the endocyclic form, and Cummins and Hiratsuka
DESCRIPTION: (1983) followed Laundon’s (1975) argument that the
Spermogonia intraepidermal, Group IV (type 6). name Gymmnoconia should be based on Aecidium
Aecia without peridium or paraphyses, indetermi- nitens Schwein. and applied to the endocyclic forms.
nate, Caeoma-type; aeciospores catenulate, ver- They created the genus Arthuriomyces for these
rucose. Uredinia not produced. Telia subepidermal, forms. However, Laundon’s argument was flawed
becoming erumpent; teliospores borne singly on (R.S. Peterson, personal communication 2001). As
pedicel, 2-celled, wall pigmented, germ pore 1 in Dodge (1923) showed, the fungus on the type host in
each cell; basidia external. the type area of Aecidium nitens was (at least in
1923) the long-cycled rust, not the endocyclic form.
TYPE: Gymnoconia peckiana (Howe in Peck) Trotter Therefore, Gymnoconia should be the name for the
=Puccinia peckiana Howe in Peck long-cycled rust, with Arthuriomyces as a synonym.
=Arthuriomyces peckianus (Howe in Peck)
Cummins & Y. Hirats. REFERENCES:
=Gymnoconia interstitiale Lagerh. Cummins, G.B.; Hiratsuka,Y. 1983. Illustrated genera of rust
On: Rubus sp. (Rosaceae) fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.

Type locality: New York, USA Dodge, B.O. 1923. The distribution of the orange-rusts of Rubus.
Phytopathology 13:61-74.
REMARKS: Dodge, B.O. 1924. Uninucleated aecidiospores in Caeoma nitens,
and associated phenomena. J. Agric. Res. 28:1045-1058.
Only the type species is known; it is widespread
Kunkel, L.O. 1913. The production ofa promycelium by aecio-
in North America, Europe, and Asia. It varies spores of Cacoma nitens Burrill. Bull. Torrey Bot. Club 40:361—
morphologically; for instance, a new variety was 366.
recently described from China (Zhang et al. 1997). It Lagerheim, G. von. 1894. Uber Uredineen mit variablem Pleo-
also varies in life cycle: two endocyclic forms are morphismus. Ein Beitrag zur Biologie der Rostpilze. Tromse
in North America (Dodge 1924; Kunkel Mus. Aarsheft. 16:105-152.
known
Laundon, G.F. 1975. Taxonomy and nomenclature notes on
1913), one with spermogonia that produce four
Uredinales. Mycotaxon 3:133-161.
basidiospores per basidium and one that gener- A.F. 1969. Gymnoconia_ nitens. Com-
Laundon, G.F,; Rainbow,
ally lacks spermogonia and produces two monw. Mycol. Inst. Description of pathogenic fungi and bac-
basidiospores per basidium. We do not have a teria 20. 2 p.
proper generic name and a legitimate species epi- Zhang, N.; Zhuang, J.Y.; Wei, S.X. 1997. Fungal flora of the Daba
thet for the endocyclic forms at the present time. Mountains: Uredinales. Mycotaxon 61:49-79.

FIGURE 106. Gymmnoconia. G. peckiana (Howe in Peck) Trotter. A. Aeciospores. B. Teliospores.

(A and B reprinted with permission, from Laundon and Rainbow 1969.)
C. Germinating teliospores of two endocyclic forms.
 HAMASPORA Korn.
Hedwigia 16:17-31 (22-23). 1877.

FAMILY: Phragmidiaceae REMARKS:

About 15 species are reported. They are pre-
DESCRIPTION: sumed to be autoecious, all occur on Rosaceae, and
Spermogonia subepidermal, Group IV (type 8 or all occur in warm areas of Africa, Australia, and
10). Aecia subepidermal, erumpent, Uredo-type; widespread locations in Asia. The species have long
aeciospores borne singly on pedicels, echinulate. nematode-like teliospores. The genus presumably
Uredinia subepidermal, erumpent, Uredo-type, has a single germ pore per cell in the teliospores.
paraphysate; urediniospores borne singly on pedi- This genus is distinguished from Phragmidium and
cels, echinulate, pores obscure. Telia subepidermal, Frommeélla by its long, narrow, colorless teliospores
erumpent in pale, feltlike or yarnlike masses; telio- without distinct germ pores.
spores 2- to several-celled by transverse septa, borne
singly on stout pedicels, spore wall pale and more or REFERENCES:
less uniformly thin except the apex, germ pore Kornicke, F. 1877. Mykologische Beitrage. Hedwigia 16:17-31.
obscure, but probably 1 per cell; germination occurs Monoson, H.L.1969. The species of Hamaspora. Mycopathol.
without dormancy; basidia external. Mycol. Appl. 37:263-272.

TYPE: Hamaspora longissima (Thum.) Korn.

=Phragmidium longissimum Thum.
On: Rubus rigidus Smith (non Merc.) (Rosaceae)
Type locality: South Africa

FIGURE 107. Hamaspora. A. A germinating teliospore,

a part of a paraphysis, and uredinio-
spores of H. longissima (Thum.) Korn.
B. A teliospore and a urediniospore of
H. hashiokae Hirats. f.
 JOERSTADIA Gjaerum & Cummins
Mycotaxon 15:420-424 (420). 1982.

FAMILY: Phragmidiaceae REMARKS:

There are four species, two demicyclic and two
DESCRIPTION: microcyclic; all occur on species of Alchemilla in
Spermogonia subepidermal, Group IV (type 8). Africa. The genus differs from other genera that
Aecia subepidermal, erumpent, Caeoma-type; aecio- have pedicellate, 2-celled teliospores because of the
spores catenulate, verrucose. Uredinia unknown, type 8 spermogonia.
probably not produced. Telia subepidermal, erum-
pent; teliospores borne singly on pedicels, 2-celled REFERENCE:
by transverse septum, germ pore not seen. Gjaerum, H.B.; Cummins, G.B. 1982. Rust fungi (Uredinales) on
East African Alchemilla. Mycotaxon 15:420-424.

TYPE: Joerstadia alchemillae (Bacc.) Gjaerum &

Cummins
=Gymnoconia alchemillae Bacc.
On: Alchemilla pedata Hochst. ex A. Rich.
(Rosaceae)
Type locality: Scioa ad Entotto, Ethiopia

FIGURE 108. Joerstadia. A. Teliospores of J. alchemillae (Bacc.) Gjaerum &

Cummins. B. Teliospores of J. keniensis Gjaerum & Cummins.
(Reprinted with permission, from Gjaerum and Cummins 1982.)
 KUEHNEOLA Magnus
Bot. Centralbl. 74:165-170 (169). 1898.

FAMILY: Phragmidiaceae autoecious and macrocyclic. Kuehneola uredinis may

cause defoliation of commercial raspberry plants
DESCRIPTION: and also can infect stems, causing “cane blight”. The
Spermogonia subcuticular, Group IV (type 11). genus Spirechina, described on Rubus from Bolivia
Aecia subepidermal, erumpent, Uredo-type; aecio- (Arthur 1907), is considered to be synonymous with
spores borne singly on pedicels. Uredinia Kuehneola (Jackson 1931). Thirumalachar (1960) con-
subepidermal, erumpent, Uredo-type; uredinio- sidered Catenulospora Mundkur as a synonym of
spores borne singly on pedicels, echinulate or other- Kuehneola rather than of Cerotelium (as in Cummins
wise marked, pores equatorial or obscure. Telia and Hiratsuka 1983) and transferred four species of
subepidermal, erumpent; teliospores borne singly Catenulospora to Kuehneola.
on short pedicels, 2- to several-celled by transverse
septa, wall pale or colorless, smooth, germ pores 1 in REFERENCES:
each cell; germination occurs without dormancy; Arthur, J.C. 1907. New genera of Uredinales. J. Mycol. 13:28-32.
basidia external. Arthur, J.C. 1912. Uredinales. N. Am. Flora 7:182-187.
Bagyanarayana, G.; Rao, K.N. 1985. A new species of Kuelneola
TYPE: Kuehneola albida J.G. Kihn) Magnus montana from India. Can. J. Bot. 63:762-764.
=Chrysomyxa albida J.G. Kuhn Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
fungi. Rev. Ed. APS Press, St. Paul, MN. 152 p.
=Kuehneola uredinis (Link) Arthur
Jackson, H.S. 1931. The rusts of South America based on the
On: Rubus fruticosus L. (Rosaceae) Holway collections. III. Mycologia 23:96-116.
Type locality: Germany Magnus, P. 1898. Einige Bemerkungen zu P. Dietel’s Bearbeitung
der Hemibasidii u. Uredinales in Engler-Prantl, Nattirlichen
REMARKS: Planzenfamilien. Bd. I. Bot. Centralbl. 74:165-170.
The genus has only a few species; most are on Parmelee, J.A.; de Carteret, P.M. 1986. Kuehneola uredinis. Fungi
Rosaceae but others are on Anacardiaceae, Can. 307.
Malvaceae, Verbenaceae, and Celastraceae Thirumalachar, M.J. 1960. Critical notes on some plant rusts. III.
Mycologia 52:688-693.
(Bagyanarayana and Rao 1985). All species are

FIGURE 109. Kuehneola. A. A telium with one germinating teliospore of K. uredinis (Link) Arthur. B. Urediniospores
of K. uredinis. C. A teliospore and a urediniospore of K. loesneriana (Arthur) HS. Jacks. & Holw.

Faye)
 PHRAGMIDIUM Link
Mag. Ges. Naturfr. Freunde Berlin 7:25—45 (30). 1816.

FAMILY: Phragmidiaceae sections are recognized by Arthur (1934), Earlea,

with smooth or almost smooth teliospores with firm
DESCRIPTION: non-hygroscopic pedicels, and Euphragmidium, with
Spermogonia Group IV (type 11, subcuticular, or verrucose teliospores with hygroscopic pedicels.
type 10, intraepidermal). Aecia subepidermal, There are about 10 species that have uredinoid
erumpent, Caeoma-type, with catenulate spores, or aecia and terete, basally septate pedicels. These
less often, Uredo-type, with spores borne singly; are sometimes placed in Phragmotelium, as by
aeciospores verrucose or echinulate, pores scattered. Thirumalachar and Mundkur (1949).
Uredinia subepidermal, erumpent, with peripheral Aregma Fr. is an earlier name for this genus and
paraphyses, Uredo-type or Calidion-type; uredinio- the name Phragmidium needs conservation
spores borne singly on pedicels, mostly echinulate, (Laundon 1965).
pores scattered, obscure. Telia subepidermal, erum-
pent; teliospores borne singly on often hygroscopic REFERENCES:
pedicels, 1- to several-celled by transverse septa, Arthur, J.C. 1934. Manual of the rusts in United States and
wall pigmented, smooth or more often verrucose, Canada. Purdue Research Foundation, Lafayette, IN.
often obviously bilaminate, germ pores 2 or usually Reprinted with a supplement by G.B. Cummins (1962),
Hafner Publ. Co., New York. 438 + 24 p.
3 in each cell; basidium external.
Cummins, G.B. 1931. Phragmidium species of North America:
differential teliospores and aecial characters. Mycologia
TYPE: Phragmidium mucronatum (Pers.) Schltdl. 23:433-445.
=Puccinia mucronata Pers. Hiratsuka, N.; Kaneko, S.; Nishigaki, H. 1980. A taxonomic
On: Rosa centifolia L. (Rosaceae) revision of the species of Phragmidium in the Japanese Archi-
Type locality: Europe pelago. Rep. Tottori Mycol. Inst. Jpn. 18:53-88.
Laundon, G.F. 1965. The generic names of Uredinales. Com-
REMARKS: monw. Mycol. Inst. Mycol. Pap. 99. Kew, Surrey. 24 p.
About 60 to 65 species have been recognized, but Link, H.F. 1816. Observationes in ordines plantarum naturales.
Mag. Ges. Naturfr. Freunde Berlin 7:25-45.
not all are distinctive. Sydow and Sydow (1915) list
Sydow, P.; Sydow, H. 1915. Phragmidium Link. Monogr.
63 species. There is no recent world monograph. All Uredinearum 3:89-159.
species are autoecious and most are macrocyclic, but Thirumalachar, M.J.; Mundkur, B.B. 1949. Genera of rusts. II.
there are demicyclic and microcyclic species. All Indian Phytopathol. 2:193-244.
species occur on the Rosaceae, with Rosa, Rubus, and
Potentilla the common hosts. The genus predomi-
nantly inhabits the Northern Hemisphere. Two

154
FIGURE 110. Phragmidium. A. A teliospore of P. biloculare Dietel & Holw. B. A teliospore of P. boreale Tranzschel.
C. A teliospore, a paraphysis, and a urediniospore of P. peckianum Arthur. D. A teliospore and a
urediniospore of P. fusiforme J. Schrot.
 TRACHYSPORA Fuckel
Bot. Z. 19:249-252 (250). 1861.

FAMILY: Phragmidiaceae REMARKS:

Trachyspora is a genus of four species on
DESCRIPTION: Alchemilla of the Rosaceae. Henderson (1973) re-
Spermogonia Group IV (type 10), but usually not ported that the surface sculpture is similar to that of
produced or not collected. Aecia subepidermal, some species of Phragmidium, thus fortifying the
erumpent, without peridium, Petersonia-type; aecio- opinion that the spores are aeciospores and that the
spores catenulate but without intercalary cells, relationship is with that genus. Gjerum and
appearing irregularly verrucose under moderate Cummins (1982) reported that spermogonia of T.
magnification, but actually spinose (Henderson intrusa were intraepidermal and indeterminate,
1973), with spines isolated or in groups on basal with a flat hymenium, corresponding to type 10 of
plaques. Uredinia not produced, but some sup- Cummins and Hiratsuka (1983), the phragmidioid
posed urediniospores occasionally occur in the telia, type. A previously recognized species, T. vestita
echinulate, pores obscure. Telia subepidermal, (Dietel) J.C. Lindgq., was found to bea hyphomycete,
erumpent, produced both on systemic mycelium in Chlamydomyces palmarum (Cooke) E.W. Mason.
the old aecia or localized; teliospores 1-celled, borne
singly on 1-septate pedicels, the upper cell short, REFERENCES:
spore wall pigmented, germ pore obscure; basidium Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
external. fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
Fuckel, L. 1861. Mykologisches. Bot. Z. 19:249-252.
TYPE: Trachyspora alchemillae Fuckel Gjeerum, H.B.; Cummins, G.B. 1982. Rust fungi (Uredinales) on
East African Alchemilla. Mycotaxon 15:420-424.
=T. intrusa (Grev.) Arthur
Henderson, D.M. 1973. Studies in the morphology of fungal
On: Alchemilla vulgaris L. (Rosaceae)
spores, Trachyspora intrusa. Rep. Tottori Mycol. Inst. 10:163-
=A. xanthochlora Rothm. 168.
Type locality: Java

FIGURE 111. Trachyspora. A. Teliospores and urediniospores of T. keniensis Gjaerum

& Cummins. B. Teliospores of T. intrusa (Grev.) Arthur. (Reprinted
with permission, from Gjarum and Cummins 1982.)
 XENODOCHUS Schltdl.
Linnaea 1:235-249 (237). 1826.

FAMILY: Phragmidiaceae REMARKS:

The type and X. minor Arthur are known. Both
DESCRIPTION: are circumboreal on species of Sanguisorba of the
Spermogonia intraepidermal, type 10. Aecia Rosaceae. The type is autoecious and macrocyclic;
subepidermal, erumpent, Caeoma-type; aeciospores X. minor is microcyclic. The genus differs from
catenulate, with papillate verrucae. Uredinia and Frommeella because of two germ pores in all but the
urediniospores similar to aecia and aeciospores apical teliospore cell. Sato and Sato (1980) have
except not accompanied by spermogonia. Telia determined the complete life cycle of X. carbonarius
subepidermal, erumpent; teliospores borne singly by inoculations.
on short pedicels, 2- to many-celled by transverse
septa, wall smooth, pigmented, germ pore 1 (rarely REFERENCES:
2) in apical cell, others with 2 pores; basidia external. Sato, T.; Sato, S. 1980. The caeomoid uredinium of Xenodochus
carbonarius (Uredinales). Trans. Mycol. Soc. Jpn. 21:411-416.
TYPE: Xenodochus carbonarius Schltdl. Schlechtendal, D.F.L. de. 1826. Fungorum novorum et descrip-
On: Sanguisorba officinalis L. (Rosaceae) torum illustrationes. Linnaea 1:235-249.
Type locality: Germany

FIGURE 112. Xenodochus. A. Teliospores of X. minor Arthur. B. Urediniospores and

a teliospore of X. carbonarius Schltdl.

7
 CHRYSELLA Syd.
Ann. Mycol. 24:283—426 (292). 1926.

FAMILY: Pucciniaceae REMARKS:

The type is the only species in the genus. Buritica
DESCRIPTION: (1974) treats this species as belonging in Chryso-
Spermogonia subepidermal, Group V (type 4). cyclus.
Aecia and uredinia unknown, probably not pro-
duced. Telia subepidermal, erumpent, waxy when REFERENCES:
wet, hard when dry; teliospores borne singly on Buritica, P. 1974. A revision of rust genera (Uredinales) with re-
pedicels; germination occurs without dormancy; duced life cycles. Ph.D. thesis, Purdue Univ., West Lafayette,
IN. 114 p.
basidia produced internally by division of telio-
Sydow, H. 1926. Fungi in itinere costaricensi collecti. Ann. Mycol.
spore contents into 4 cells by transverse septa, each 24:283-426.
cell producing a long sterigma and a large basidio-
spore.

TYPE: Chrysella mikaniae Syd.

On: Mikania hirsutissima DC. (Asteraceae)
Type locality: Costa Rica

FIGURE 113. Chrysella. Teliospores, internal basidia, and

basidiospores of C. mikaniae Syd.
 CHRYSOCYCLUS Syd.
Ann. Mycol. 23:308-429 (322). 1925.

FAMILY: Pucciniaceae that the genus is little more than an exaggerated

micro-Puccinia. The basidium of the lower cell is not
DESCRIPTION: in the teliospore cell, but that of the upper cell comes
Spermogonia subepidermal, Group V (type 4). close to being an internal basidium. Perhaps the
Aecia and uredinia unknown, probably not pro- genus is midway between Puccinia and Chrysopsora
duced. Telia subepidermal, erumpent; teliospores and both are derivatives of Puccinia. Buritica (1974)
2-celled, borne singly on pedicels, wall pale, each treats Chrysella as a synonym of Chrysocyclus. This
cell becoming 4-celled by apical growth and by the genus may be closer to Mikronegeriaceae because of
subsequent division of the protoplast into 4 parts, the deep-seated spermogonia (almost type 12) and
each of which produces a sterigma and basidio- teliospore germination by apical elongation of cells
spore, germination occurs without dormancy; without germ pores (semi-internal basidia).
basidia semi-internal.
REFERENCES:
TYPE: Chrysocyclus cestri (Dietel & Henn.) Syd. Buritica. P. 1974, A revision of rust genera (Uredinales) with
(Lectotype) reduced life cycles. Ph.D. thesis, Purdue Univ., Lafayette, IN.
=Puccinia cestri Dietel & Henn. 114 p.
On: Cestrum megalophyllum Dun. (Solanaceae) Jackson, H.S. 1932. The rusts of South America based on the
Holway collections. VI. Mycologia 24:62-186.
Type locality: Costa Rica
Sydow, H. 1925. Fungi initinere costaricensi collecti. Ann. Mycol.
23:308-429.
REMARKS:
Four species are known in South and Central
America (Buritica 1974). Jackson (1932) suggested

FIGURE 114. Chrysocyclus. Germinating teliospores

of C. cestri (Dietel & Henn.) Syd.

R59.
 CHRYSOPSORA Lagerh.
Ber. Dtsch. Bot. Ges. 9:344-348 (345-346). 1891 (issued 1892).

FAMILY: Pucciniaceae point of interest is that it proves that an y)“internal

basidium” is not characteristic of only the Coleo-
DESCRIPTION: sporiaceae. Two other species, C. cestri (Henn.)
Spermogonia subepidermal, Group V (type 4). Arthur and C. mikaniae Arthur, which were consid-
Aecia and uredinia not produced. Telia subepider- ered to be distinct and were transferred to Hol-
mal, erumpent, waxy when fresh or moist; teliospores wayella by Jackson (1926), are now recognized as
2-celled with pedicels, wall pale; germination species of Chrysocyclus (Jackson 1932).
occurs without dormancy; 4-celled basidia pro-
duced internally by separation of cell contents by REFERENCES:
transverse septa. Jackson, H.S. 1926. Preliminary note on the genus Chrysopsora.
Mycologia 18:48—-49.
TYPE: Chrysopsora gynoxidis Lagerh. Jackson, H.S. 1932. The rusts of South America based on the
On: Gynoxys pulchella DC. (Asteraceae) Holway collections. VI. Mycologia 24:62-186.
Type locality: South America Lagerheim, G. von. 1891 (issued 1892). Pucciniosira, Chrysopsora,
Alveolaria und Trichospora, vier neue Uredineen-Gattungen
mit tremelloider Entwicklung. Ber. Dtsch. Bot. Ges. 9:344-348.
REMARKS:
The type is the only known species. There are no
good clues to the relationship of this genus, but a

FIGURE 115. Chrysopsora. Teliospores

and internal basidia of C.
gynoxidis Lagerh.

160
 CLEPTOMYCES Arthur
Bot. Gaz. 65:460-474 (464). 1918.

FAMILY: Pucciniaceae differs from Puccinia and Prospodium because of

multiple germ pores and from Prospodium in the
DESCRIPTION: type of spermogonium. Stereostratum, with only telia
Spermogonia subepidermal, Group V (type 4). known and parasitizing Poaceae (Bambusoideae),
Aecia and uredinia unknown. Telia subepidermal, has three pores per cell. Thirumalachar (1960) con-
erumpent; teliospores 2-celled by transverse sidered the genus to be synonymous with Stereo-
septum, borne singly on pedicels, wall pigmented stratum and transferred C. lagerheimianus (Dietel)
and bilaminate, germ pores 4 or more in each cell; Arthur to that genus, but we recognize them sepa-
germination not recorded, but basidia undoubtedly rately. When the aecial states become known, the
external. question of relationship may be decided.
TYPE: Cleptomyces lagerheimianus (Dietel) Arthur REFERENCES:
=Puccinia lagerheimiana Dietel Arthur, J.C. 1918. Uredinales of the Andes, based on collections
On: Aegiphila sp. (Verbenaceae) by Dr. and Mrs. Rose. Bot. Gaz. 65:460-474.
Type locality: Equador Thirumalachar, M.J. 1960. Critical notes on some plant rusts. III.
Mycologia 52:688-693.
REMARKS:
The type and one other species on Fabaceae are
known. Both occur in South America. The genus

FIGURE 116. Cleptomyces. Teliospores of C. lagerheimianus

(Dietel) Arthur.

161
 CORBULOPSORA Cummins
Mycologia 32:359-375 (364-365). 1940.

FAMILY: Pucciniaceae REMARKS:

Three species have been recognized, all on
DESCRIPTION: Asteraceae, the type and C. gravida Cummins on
Spermogonia deep-seated, Group V (type 4). Olearia in New Guinea, and C. cumminsii Thirum. on
Aecia deep-seated, becoming erumpent, with Lactuca in India. Corbulopsora differs from Uromyces
peridium, Aecidium-type; aeciospores catenulate. because of the palisadelike peridium, just as Miyagia
Uredinia subepidermal, becoming erumpent, witha differs from Puccinia. Cummins (1959) treated Cor-
tubular peridium composed of long, laterally bulopsora as a synonym of Puccinia, but we consider
united, palisadelike cells, Uredostilbe-type; uredinio- this a separate genus as in Cummins and Hiratsuka
spores borne singly on pedicels, echinulate, pores (1983).
scattered or obscure. Telia subepidermal, erumpent,
with a peridium, as the uredinia; teliospores 1- REFERENCES:
celled, borne singly on pedicels, several of which Cummins, G.B. 1940. Uredinales of New Guinea. Mycologia
arise from single basal sporogenous cells, germ pore 32:359-375.
1; basidia external. Cummins, G.B. 1959. Illustrated genera of rust fungi. Burgess
Publ. Co., Minneapolis, MN. 131 p.
Cummins, G.B.; Hiratsuka, Y. 1983. Illustrated genera of rust
TYPE: Corbulopsora clemensiae Cummins fungi. Rev. ed. APS Press, St. Paul, MN. 152 p.
On: Olearia sp. (Asteraceae) Hiratsuka, N. 1969. Notes on the genus Miyagia Miyabe ex
Type locality: New Guinea Sydow. Trans. Mycol. Soc. Jpn. 10:89-90.

FIGURE 117. Corbulopsora. C. clemensiae Cummins. A. Peridial cells.

B. A urediniospore. C. Teliospores.

162
 CUMMINSIELLA Arthur
Bull. Torrey Bot. Club 60:465-476 (475). 1933.

FAMILY: Pucciniaceae REMARKS:

There are eight species, all on Mahonia or Berberis
DESCRIPTION:
(Berberidaceae), all autoecious and macrocyclic,
Spermogonia subepidermal, Group V (type 4).
and all native to North and South America (Baxter
Aecia subepidermal, erumpent, with peridium,
1957; McCain and Hennen 1982). Cumminsiella mira-
Aecidium-type; aeciospores catenulate, verrucose.
bilissima (Peck) Nannf. has been introduced with its
Uredinia subepidermal, erumpent, with or without
host into other regions. The aecial state was first
paraphyses, Uredo-type; urediniospores borne
recognized in Europe and has been collected only
singly on pedicels, echinulate or verrucose, pores
rarely in its original area. Cumminsiella differs from
mostly zonate. Telia subepidermal, erumpent; telio-
Puccinia because of the two pores per teliospore cell
spores 2-celled, borne singly on pedicels, wall
and from Uropyxis because of Group V (type 4) sper-
pigmented, often bilaminate, the surface variously
mogonia and Aecidium-type aecia.
sculptured, germ pores 2 in each cell; basidia exter-
nal.
REFERENCES:
Arthur, J.C. 1933. New genera and species of Uredinales. Bull.
TYPE: Cumminsiella sanguinea (Arthur) Arthur Torrey Bot. Club 60:465-476.
(Designated type) Baxter, J.W. 1957. The genus Cumminsiella. Mycologia 49:864-873.
=Uropyxis sanguinea Arthur McCain, J.W.; Hennen, J.F. 1982. Is the taxonomy of Berberis and
=Cumminsiella mirabilissima (Peck) Nannf. Mahonia (Berberidaceae) supported by their rust pathogens,
On: Mahonia sp. (Berberidaceae) Cumminsiella santa sp. nov. and other Cumminsiella species
(Uredinales)? Syst. Bot. 7:48-59.
Type locality: USA

FIGURE 118. Cumminsiella. A. A teliospore and urediniospores of C. mirabilissim

a (Peck) Nannf. B. A teliospore and a
urediniospore of C. wootoniana (Arthur) Arthur. C. A teliospore of C. standleyana Cummins.
 GYMNOSPORANGIUM R. Hedwig
in Lamarck and de Candolle, Fl. Fr. 2. 600 p. (216). 1805.

FAMILY: Pucciniaceae Coleopucciniella as a separate genus, but we consider

it and Coleopuccinia to be synonyms of Gymno-
DESCRIPTION:
sporangium.
Spermogonia subepidermal, Group V (type 4).
Aecia subepidermal, erumpent, with peridia, typi- The variation among species makes a precise
generic description nearly impossible. Type 4 sper-
cally Roestelia-type or in a few species Aecidium-
type; aeciospores catenulate, verrucose, most with
mogonia occur in all species where the life cycle is
pigmented wall. Uredinia subepidermal, Uredo-type, known, and gelatinizing teliospore pedicels charac-
erumpent, lacking in most species; urediniospores terize nearly all of the species.
borne singly on pedicels, verrucose, pores scattered. Leppik (1956) discussed the evolution of biologi-
Telia subepidermal, erumpent as cushions, crests, or cal specialization in the genus.
horns, gelatinizing when moist, mostly perennial in Economically important species include G.
the host, often inducing swelling or fasciation; telio- Juniperi-virginianae Schwein. in North America on
spores I- to several-celled but mostly 2-celled by apples and pears as well as on cultivated Juniperus
transverse septum, borne singly on gelatinizing species, and G. asiaticum Miyabe ex Yamada and G.
pedicels (with perhaps rare exceptions), germ pores yamadae Miyabe ex Yamada on pears in Asia.
1 to several but most often 2 per cell, variously
placed, wall mostly pigmented and smooth; germi- REFERENCES:
nation occurs without dormancy; basidia external. Bernaux, P. 1956. Contribution a l’étude de la biologie des
Gymnosporangium. Ann. Epiphyt. 7:1-210.
Harada, Y. 1984. Pear and apple rusts in Japan, with special
TYPE: Gymnosporangium fuscum R. Hedwig refer-
ence to their life cycles and host ranges. Rep. Tottori Mycol.
(Lectotype) Inst. 22:108-119.
On: Juniperus sabina L. (Pinaceae) Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka,
Y.;
Type locality: France Kaneko, S.; Ono, Y.; Sato, ie aradaye: Hiratsuka, T.;
Nakayama, K. 1992. The rust flora of Japan. Tsukuba
REMARKS: Shuppankai, Ibaraki. 1205 + 159 pp.
According to Kern (1973), 57 species of Gym- Hiratsuka, Y. 1973. Sorus development, spore morphology, and
nosporangium and 14 species of Roestelia that are pos- nuclear condition of Gymnosporangium gaeumannii ssp.
sibly anamorphic states of Gymnosporangium have albertensis. Mycologia 65:137-144.
Kern, ED. 1911. A biologic and taxonomic study of the genus
been reported. This is a genus of predominantly
Gymnosporangium. Bull. N.Y. Bot. Gard. 7:391—483.
north temperate climates. Most species are
Kern, ED. 1973. A revised account of Gymmosporangium. Pennsyl-
heteroecious and demicyclic. For an example of vania State Univ. Press, University Park, PA. 134 p-
unusual urediniospore and teliospore morphology, Lamarck, J.; de Candolle, A.P. 1805. Flore francaise. 2. 600 p.
see Hiratsuka (1973) for G. gaeumannii Zogg ssp. Laundon, G.F. 1975. Taxonomy and nomenclature notes on
albertensis Parmelee. Peterson (1967a, 1982) de- Uredinales. Mycotaxon 3:133-160.
scribed Uredo cupressicola R.S. Peterson, which is Lee, S.K.; Kakishima, M. 1999a. Aeciospore surface structures
of
presumably related to Gymnosporangium, with Gymnosporangium and Roestelia (Uredinales). Mycoscience
occasional two- or even three-celled pedicellate 40:109-120.

urediniospores. Gymnosporangium is unusual be- Lee, S.K.; Kakishima, M. 1999b. Surface structures of peridial
cells
of Gymnosporangium and Roestelia (Uredinales). Mycoscience
cause the telial state is on gymnosperms and the
40:121-131.
aecial state on dicotyledonous plants, especially the
Leppik, E.E. 1956. Some viewpoints on the phylogeny of rust
Pomoideae of the Rosaceae. There are four species, fungi. II. Gymnosporangium. Mycologia 48:637-654.
all microcyclic, all Asian, and all on Rosaceae, Parmelee, J.A. 1965.
that The genus Gymnosporangium in Eastern
have been described as species of Coleopuccinia Canada. Can. J. Bot. 43:239-267.
(including species of Coleopucciniella). It is logical to Parmelee, J.A. 1971. The genus Gymnosporangium in Western
assume that they are derivatives of demicyclic Canada. Can. J. Bot. 49:903-926.
species of Gymnosporangium, in accordance with Parmelee, J.A.; Corlett, M. 1978. Development of the
aecium and
Tranzschel’s Law. Laundon (1975) clearly reported the nuclear condition of Roestelia brucensis. Rep. Tottori
Mycol. Inst. 10:189-201.
the pedicellate nature of teliospores of Coleo-
Peterson, R.S. 1967a. Cypress rusts in California
pucciniella. Hiratsuka et al. (1992) recognized and Baja Cali-
fornia. Madrono 19:47—54,

164
Peterson, R.S. 1967b. Studies of juniper rusts in the West. Thirumalachar, M.J.; Whitehead, M.D. 1954. On the validity of
Madrono 19:79-91, the genera Coleopuccinia and Coleopucciniella (Uredinales).
Peterson, R.S. 1982. Rust fungi (Uredinales) on Cupressaceae. Am. J. Bot. 41:120-122.
Mycologia 74:903-910.

FIGURE 119. Gymnosporangium. Teliospores. A. G. libocedri (Henn.) F. Kern. B. G. clavariiforme (Pers.) DC.
C. G. clavipes (Cooke & Peck) Cooke & Peck. D. G. multiporum F. Kern. E. G. ellisii (Berk.) Ellis.
F. G. juniperi-virginianae Schwein.
 FIGURE 120. Gymnosporangium. Aeciospores. A, B. G. speciosu
m Peck. C, D. Roeste lia sp.
E, F. Roestelia distorta (Arthur) F. Kern. G, H. G. exterum
Arthur & F. Kern. (Reprinted
with permission, from Lee and Kakishima 1999a.)

166
 oe épnnanrs
eee ps > ie

FIGURE 121. Gymnosporangium. Peridial cells of aecia (OW = outer wall; IW = inner wall; SW = side
wall). A. G. biseptatum Ellis. B. G. hyalinum F. Kern ex Cummins. C, D. G. exiguum F. Kern.
E. G. cunninghamianum Barclay. F. G. fuscum R. Hedwig. G. G. fraternum F. Kern. H. G.
yamadae Miyabe ex Yamada. (Reprinted with permission, from Lee and Kakishima 1999b.)

167
 KERNELLA Thirum.
Mycologia 41:97. 1949.

FAMILY: Pucciniaceae REMARKS:

Only the type species is known. This fungus
DESCRIPTION:
was originally named and described as Kernia
Spermogonia, aecia, and uredinia unknown. (Thirumalachar 1946), a preempted name.
Telia subepidermal, deep-seated, without peridium
Thirumalachar (1949) created a new genus, Kerrella,
but with a peripheral “dermal” layer, becoming
and transferred the type. Ragunathan and
erumpent as long, hairlike columns; teliospores Ramakrishnan (1973) treated Kernella as a synonym
2-celled by transverse septum, borne singly on pedi-
of Puccinia, and this is not illogical, but we are retain-
cels, new spores push older spores outward, and ing the genus because of the columnar telia.
because the spores or pedicels adhere to each other,
a column results, wall pale, germ pore 1 in each cell;
REFERENCES:
germination occurs without dormancy; basidia
Ragunathan, A.N.; Ramakrishnan, K. 1973. Rust fungi of
Madras
external. State. VI. Puccinia. Mysore J. Agric. Sci. 7:62-72.
Thirumalachar, M.J. 1946. Kernia, anew genus of the Uredinales.
TYPE: Kernella lauricola (Thirum.) Thirum. Mycologia 38:679-686.
=Kernia lauricola Thirum. Thirumalachar, M.J. 1949. Corrections. Mycologia 41:97.

On: Litsea sp. (Lauraceae)

Type locality: India

LJ
=<)

=
ApYaL
~~
Qe.
AD,

FIGURE 122. Kernella. A telium and teliospores of K. lauricol

a (Thirum.) Thirum.

168
 MIYAGIA Miyabe ex H. & P. Syd.
Ann. Mycol. 11:93-118 (107). 1913.

FAMILY: Pucciniaceae only in the peridiate uredinia and telia. In the type
species, 1-celled teliospores are sometimes more
DESCRIPTION: common than 2-celled spores. Wang and Zhuang
Spermogonia subepidermal, Group V (type 4). (1986) pointed out that the peridial structure of M.
Aecia subepidermal, erumpent, with thick-walled pseudosphaeria is different from the other two
peridium, Aecidium-type; aeciospores verrucose and species, and it should be classified under Puccinia.
catenulate, pores obscure. Uredinia subepidermal, Also on Asteraceae is Corbulopsora, with similarly
erumpent, with a peridium of laterally adherent, peridiate uredinia and telia, but with only 1-celled
palisadelike paraphyses, Uredostilbe-type; teliospores and with Uredo-type aecia. The relation-
urediniospores borne singly on pedicels, echinulate. ship of Miyagia doubtless is with Puccinia and that of
Telia subepidermal, tardily exposed, with a Corbulopsora with Uromyces.
peridium as in the uredinia; teliospores 2-celled by
transverse septum, borne singly on pedicels, wall REFERENCES:
pigmented, germ pore 1 in each cell; basidium Hiratsuka, N. 1969. Notes on the genus Miyagia Miyabe ex Syd.
external. Trans. Mycol. Soe. Jpn. 10:89-90.
Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
TYPE: Miyagia anaphalidis Miyabe ex H. & P. Syd. Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
Nakayama, K. 1992. The rust flora of Japan. Tsukuba
On: Anaphalis margaritacea Benth. & Hook. f. var.
Shuppankai, Ibaraki 1205 + 159 p.
angustior Nakai (Asteraceae) Sydow, H.; Sydow, P. 1913. Ein Beitrag zur Kenntnis der
Type locality: Sapporo, Hokkaido, Japan parasitischen Pilzflora des nordlichen Japans. Ann. Mycol.
11:93-118.
REMARKS: Wang, Y.C.; Zhuang, J.Y. 1986. Notes on the originations of some
The type species and two other species, M. rust fungi from China, Acta Mycol. Sin. Suppl. 1:5-11.
pseudosphaeria (Mont.) Jorst. and M. macrospora Wilson, M.; Henderson, D.M. 1966. British rust fungi. Cambridge
Univ. Press. 384 p.
Hirats. f. are known (Hiratsuka et al. 1992; Wilson
and Henderson 1966). Miyagia differs from Puccinia

FIGURE 123. Miyagia. Miyagia anaphalidis Miyabe ex H. & P. Syd. A. A telium with a
peridium of laterally adherent paraphyses. B. A urediniospore.

169
 POLIOMA Arthur
J. Mycol. 13:28-32 (29). 1907.

FAMILY: Pucciniaceae (Lamiaceae) (Baxter and Cummins 1951); one

macrocyclic species (P. reniformis Leén-Gall. &
DESCRIPTION:
Spermogonia subepidermal, Group V (type 4). Cummins) is on Geranium (Geraniaceae) (Le6n
Aecia subepidermal, erumpent, without peridium,
Gallegos and Cummins 1981). They occur in Mexico
Caeoma-type;
and Ecuador. Polioma differs from Puccinia because
aeciospores catenulate, verrucose.
Uredinia subepidermal, erumpent, without of its sessile teliospores. A rust on Thermopsis
peridium, Uredo-type; urediniospores borne singly, (Fabaceae) with sessile, two-celled, verrucose telio-
echinulate, germ pore 1, basal where known. Telia spores was described from Colorado (Ramaley
subepidermal, erumpent; teliospores sessile and 1987). It might belong to Polioma. However, because
usually grouped on basal cells, 2-celled by trans- the type specimen is missing, we exclude this
verse septum, wall pale, germ pore 1 per cell; germi- species, called Poliomopsis thermopsidis A.W.
nation occurs without dormancy; basidia external. Ramaley.

TYPE: REFERENCES:
Polioma nivea (Holw.) Arthur
Arthur, J.C. 1907. New genera of Uredinales. J. Mycol.
=Puccinia nivea Holw. 13:28-32.
Baxter, J.W.; Cummins, G.B. 1951. Polioma Arth., a valid
On: Salvia purpurea Cav. (Lamiaceae) the Uredinales. Bull. Torrey Bot. Club 78:51—55,
genus of

Type locality: Oaxaca, Mexico Le6én Gallegos, H.M.; Cummins, G.B. 1981. Uredinale
s (Royas) de
Mexico. Vol. II. Secretaria Agricultura Recursos Hidraulic
os,
REMARKS: Culiacan, Sinaloa, Mexico. 492 D:
Four species have been assigned to this genus, Ramaley, A.W. 1987. A new rust on Leguminosae. Mycotaxo
n
Three are microcyclic on species of Salvia 28:361-363.

FIGURE 124. Polioma. Teliospores and urediniospores

of P. nivea (Holw.) Arthur.

170
 PUCCINIA Pers. ex Pers.
Synop. Methodica Fungorum. 708 p. (225-226). 1801.

FAMILY: Pucciniaceae rust (P. helianthi Schwein.), safflower rust (P.

calcitrapae DC. var. centaureae (DC.) Cummins),
DESCRIPTION: cotton rust (P. cacabata Arthur & Holw.), asparagus
Spermogonia subepidermal, Group V (type 4). rust (P. asparagi DC.), mint rust (P. menthae Pers.),
Aecia subepidermal, erumpent, either Aecidium- snapdragon rust (P. antirrhini Dietel & Holw.), and
type with peridium and catenulate and verrucose hollyhock rust (P. malvacearum Bert.).
spores or Uredo-type with mostly echinulate spores Theoretically, the name Puccinia Pers. ex Pers.
borne singly on pedicels. Uredinia subepidermal, (lectotype P. graminis) may need to be conserved
erumpent, without peridium, but may have against Puccinia Adams. ex Pers. (lectotype P. juni-
paraphyses, Uredo-type or sometimes Aecidium-
peri Pers. =Gymnosporangium fuscum DC.) (Laundon
type; urediniospores borne singly on pedicels, 1965).
mostly echinulate, germ pores various, or with
peridium and catenulate and verrucose. Telia REFERENCES:
subepidermal, erumpent in most species, but In addition to the following, see other descriptive
remaining covered by the epidermis and divided manuals listed under “Descriptive manuals and re-
into locules by stromatic paraphyses in some gional monographs of rust fungi”.
species; teliospores typically 2-celled by transverse
septum (but may have 1-celled and sometimes 3- or Arthur, J.C. 1907-1927. Uredinales. N. Am. Flora 7:83-848 (with
4-celled spores in some species), borne singly on Puccinia broken into several “life cycle” genera).

pedicels, basal sporogenous cells none to highly Azbukina, Z.M. 1974. [Rust fungi of the Soviet Far East.]
Akademiya Nauk, Moscow. 527 p. (In Russian.)
developed, spore wall mostly pigmented, but
Azbukina, Z.M. 1984. [Identification of rust fungi of the Soviet
degree of pigmentation varies widely, smooth or Far East.] Akademiya Nauk, Moscow. 288 p. (In Russian.)
variously sculptured, germ pore 1 in each cell, but Cummins, G.B. 1953. The species of Puccinia parasitic on the
not differentiated in a few species; basidia external. Andropogoneae. Uredineana 4:5-90.
Cummins, G.B. 1971. The rust fungi of cereals, grasses and bam-
TYPE: Puccinia graminis Pers. ex Pers. (Lectotype, boos. Springer-Verlag, New York. 570 p.
Cunningham 1931) Cummins, G.B. 1978. Rust fungi on legumes and composites in
On: Triticum vulgare Vill. (Poaceae) North America. Univ. Arizona Press, Tucson, AZ. 424 p-

=T. aestivum L. Cunningham, C.H. 1931. The rust fungi of New Zealand.
MclIndoe, Dunedin. 261 p-
Type locality: Europe
Gaeumann, E. 1959. Die Rostpilze Mitteleuropas. Buechler & Co.,
Bern. 1407 p.
REMARKS:
Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
Puccinia is the largest genus of Uredinales, with Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
about 3000 to 4000 species. Heteroecious and Nakayama, K. 1992. The rust flora of Japan. Tsukuba
autoecious species occur and also life cycle varia- Shuppankai, Tsukuba. 1205 + 159 p.
tions down to the simplest of teliospores only. The Laundon, G.F. 1965. The generic names of Uredinales. Mycol.
Asteraceae, Cyperaceae, Poaceae, and Liliaceae Pap. 99:1-24.

serve as hosts for large numbers of species, but most Leon Gallegos, H.M.; Cummins, G.B. 1981. Uredinales (Royas) de
México. Vol. I. Secretaria Agricultura Recursos Hidraulicos,
groups of vascular plants are parasitized. There are Culiacan, Sinaloa, México. 440 p.
species in all land areas except the polar regions. Lindquist, J.C. 1982. Royas de la Republica Argentina y zonas
Many serious diseases are caused by species of limitrofes. Colecc. Cient. Vol. 20. 574 p.
Puccinia, e.g. black stem rust (P. graminis Pers. ex Majewski, T. 1979. Grzyby (Mycota). Tom. XI. Uredinales II.
Pers.) of small grains and other grasses, leaf or Polska Akademia Nauk, Warszawa-Krakow. 426 p-
brown rust (P. recondita Roberge ex Desm.) of small Persoon, C.H. 1801. Synopsis methodica fungorum. Vol. I, II.
grains and other grasses, stripe or yellow rust (P. G6ttingen. 708 p.
Ul’yanischey, V.I. 1978. [Key to the rust fungi of the USSR.] Vol II.
striiformis Westend.) of wheat and other grasses,
Akademiya Nauk, Leningrad. 383 p. (In Russian.)
crown rust (P. coronata Corda) of oats and other
Zhuang, J.Y.; Wei, S.X.; Wang, Y.C. 1998. Flora fungorum
grasses, sorghum rust (P. purpurea Cooke), common sinicorum. Vol. 10. Uredinales (I). Science Press, Beijing. xi +
corn rust (P. sorghi Schwein.), sugarcane rusts (P. 335 p.
kuehnii E.J. Butler, P. melanocephala Syd.), sunflower

7
FIGURE 125. Puccinia. Teliospores. A. P. spegazziniana De
Toni. B. P. corylopsidis Cummins. C. P. scirpi-ternatani
Hirats. f. D. P. prostti Duby. E. P. scleirae (Pazschke)
Arthur. F. P. heliconiae Arthur. G. P. polysora Underw.
H. P. sorghi Schwein. I. P. hosoniana F. Kern. J. P.
cyani Pass. K. P. cacabata Arthur & Holw.
L. P. melanocephala Syd. M. P. recondita Roberge. N. P. conoclini
i Seym. O. P. solidipes H.S. Jacks. & Holw.
P. P. anisacanthi Dietel & Holw. Q. P. batatae Syd. R. P.
coronata Corda. S. P. longipedicellata Barthol.
T. P. tanaceti DC. U. P. arachidis Speg. V. P. wolgensis Nawashi
n.

i.)
 RAMAKRISHNANIA Ramachar & Bhagyan.
Can. J. Bot. 57:783-786 (783-784). 1979.

FAMILY: Pucciniaceae ontogeny of teliospores, which are produced on

elongating, septate, and branching sporogenous
DESCRIPTION: hyphae, are significantly different from teliospores
Spermogonia, aecia, and uredinia not known. of Puccinia and other genera that are produced
Telia subepidermal; teliospores 2-celled by trans- directly on the sporogenous basal cells. We tenta-
verse septum, produced on the tips of elongating tively put this genus in Pucciniaceae, but it may
and branching sporogenous hyphal cells of indeter- belong to a different family. Information on other
minate growth; germ pore 1 in each cell; basidia spore states, especially the spermogonia, is needed
external. to determine the relationship of this genus to other
groups of rusts.
TYPE: Ramakrishnania ixorae Ramachar & Bhagyan.
On: [xora sp. (Rubiaceae) REFERENCE:
Type locality: India Ramachar, P.; Bhagyanarayana, P. 1979. Ramakrishnania, a new
genus of Uredinales from southern India. Can. J. Bot. 57:783-
REMARKS: 786.
Only the type species is known. Ramachar
and Bhagyanarayana (1979) considered that the

FIGURE 126. Ramakrishnania. R. ixorae Ramachar & Bhagyan. A. Teliospores with septate pedicels. B. Teliospores
produced on elongating basal cells. (Reprinted with permission, from Ramachar and Bhagyanarayana
1979.)

7S
 STEREOSTRATUM Magnus
Ber. Dtsch. Bot. Ges. 17:176-184 (181). 1899.

FAMILY: Pucciniaceae completely replace them, producing urediniospores

between May and July. Production of large
DESCRIPTION: Corticium-like crusts sometimes measuring up to
Spermogonia and aecia unknown. Uredinia 10 cm in diameter and the three germ pores in each
subepidermal, develop under telia, erumpent, teliospore cell distinguish this genus from others.
Uredo-type; urediniospores borne singly on pedi- Thirumalachar (1960) considered Cleptomyces as a
cels, echinulate, germ pores equatorial. Telia synonym of Stereostratum, but we retain them as
subepidermal, becoming erumpent in large separate genera until more information becomes
Stereum-like crusts; teliospores 2-celled by trans- available.
verse septum, borne singly on pedicels, germ pores
3 per cell, obscure; basidia external. REFERENCES:
Berkeley, M.J. 1877. Contribution to the botany of H.M.S. Chal-
TYPE: Stereostratum corticioides (Berk. & Broome) lenger. 38. Enumeration of the fungi collected during the
Magnus expedition of H.M.S. Challenger Ill. J. Linn. Soc. Lond. Bot.
16:38-54.
=Puccinia corticioides Berk. & Broome
He, F; Kakishima, M.; Sato, S. 1990. Annual sporulation cycle of
On: Pleioblastus simonii (Carriere) Nakai the bamboo culm rust, Stereostratum corticioides, on Pleio-
(Poaceae) blastus simonti in Tsukuba, Japan. Rep. Tottori Mycol. Inst.
(Originally as Arundinaria sp.?) 28:243-249.
Type locality: Kobe, Japan He, F; Kakishima, M.; Sato, S. 1991. Inoculation experiments with
urediniospores of Stereostratum corticioides and its ontogenic
observations of sori and spores. Trans. Mycol. Soc. Jpn.
REMARKS:
32:407-415.
Only the type species is known. It parasitizes var-
Hiratsuka, N.; Sato, S.; Katsuya, K.; Kakishima, M.; Hiratsuka, Y.;
ious genera of the Bambusoideae in China, Taiwan, Kaneko, S.; Ono, Y.; Sato, T.; Harada, Y.; Hiratsuka, T.;
Japan, and probably elsewhere. He et al. (1990, 1991) Nakayama, K. 1992. The rust flora of Japan. Tsukuba
investigated the life cycle and development of sori. Shuppankai, Tsukuba. 1205 + 159 p:
According to them, telia are first produced in the fall Magnus, P. 1899. Uber die bei verwandten Arten auftretenden
Modificationen der Charaktere von Uredineen-Gattungen.
as a well-developed crust of hymenia; teliospores
Ber. Dtsch. Bot. Ges. 17:176-184.
with long pedicels develop on the crust during the
Thirumalachar, M.J. 1960. Critical notes on some plant rusts. III.
winter. Uredinia are produced under the telia and Mycologia 52:688-693.

FIGURE 127. Stereostratum. S. corticioides (Berk. & Broome) Magnus.

A. A urediniospore and teliospores. B. Urediniospores (US) among
teliospores (TS) (reprinted with permission, from He et al. 1991).

174
 UROMYCES (Link) Unger
Einfluss Bodens. p. 1-367 (216). 1836.

FAMILY: Pucciniaceae Puccinia only in having 1-celled teliospores. It may

not warrant separate status, but is maintained for
DESCRIPTION: convenience and historical reasons. To treat
Spermogonia subepidermal, Group V (type 4). Uromyces as a synonym of Puccinia would entail
Aecia subepidermal, erumpent, either Aecidium- many nomenclatural changes with no real benefit.
type, with peridium and catenulate, mostly There are several economically important
verrucose aeciospores, or Uredo-type, with mostly species: bean rust (U. appendiculatus (Pers.) Unger,
echinulate aeciospores borne singly on pedicels. two varieties), pea rust (U. pisi (DC.) G.H. Otth),
Uredinia subepidermal, erumpent, Uredo-type; alfalfa or lucerne rust (U. striatus J. Schrot.), clover
urediniospores borne singly on pedicels, usually rust (U. trifolii-repentis Liro), beet rust (U. betae
echinulate, pores various, mostly obvious. Telia (Pers.) Tul.), chickpea rust (U. ciceris-arietinus Jacz.),
subepidermal, erumpent or remaining covered by and carnation rust (U. dianthi (Pers.) Niessl).
the epidermis; teliospores borne singly on pedicels,
which may or may not be grouped on basal REFERENCES
sporogenous cells, 1-celled, wall mostly pigmented, Azbukina, Z.M. 1974. [Rust fungi of the Soviet Far East.]
pore 1; basidia external. Akademiya Nauk, Moscow. 527 p. (In Russian.)
Cummins, G.B. 1971. The rust fungi of cereals, grasses and bam-
TYPE: Uromyces appendiculatus (Pers.) Unger boos. Springer-Verlag, New York, Heidelberg, Berlin. 570 p.
(Conserved type) Cummins, G.B. 1978. Rust fungi on legumes and composites in
North America. Univ. Arizona Press, Tucson, AZ. 424 p.
=Uredo appendiculatus Pers.
Gaeumann, E. 1959. Die Rostpilze Mitteleuropas. Buchdruckerei
On: Phaseolus vulgaris L. (Fabaceae)
Buechler & Co., Bern. 1407 p.
Type locality: Europe Guyot, A.L. 1938, 1951, 1957. Les Urédinées. Genre Uromyces.
Encycl. Mycol. Vol. 8, 438 p.; Vol. 15,331 p.; Vol. 29, 649 p. Paul
REMARKS: Lechevalier, Paris.
Uromyces is the second-largest genus of rust fungi Hiratsuka, N. 1973. Revision of taxonomy of the genus Uromyces
next to Puccinia, and contains more than 600 in the Japanese Archipelago. Rep. Tottori Mycol. Inst. 10:1-98.
reported species. Its species parasitize monocots Lindquist, J.C. 1982. Royas de la Republica Argentina y zonas
and dicots throughout the world. The Asteraceae, limitrofes. Coleccion Cientifica. Vol. 20. Inst. Nac. Tecnol.
Agropecu., Argentina. 574 p.
Poaceae, Liliaceae, Euphorbiaceae, and Fabaceae
Unger, F. 1836. Uber den Einfluss des Bodens auf die Vertheilung
support more species than other families. All varia- der Gewachse, nachgewiesen in der Vegetation des
tions of life cycles occur. For whatever reason, there nordostlichen Tirol’s. Rohrmann und Schweigerd, Wien. Vol.
are many species of Uromyces, but very few species 24. 367 p.
of Puccinia, on legumes. Uromyces differs from

Fie)
FIGURE 128. Uromyces. Teliospores. A. U. viciae-fabaeJ.Schrét. B. U.
appendiculatus (Pers.) Unger. C. U. vignae Barclay.
D. U. amurensis Kom. E. U, glycyrrhizae Magnus. F. U. striatus
J. Schrot. G. U. eragrostidis Tracy.
H. U. aleuropodis-repentis Nattras. I. U. dactylidis G.H. Otth. J. U. halstedii
De Toni. K. U. linearis Berk.
L. U. seditiosus F. Kern. M. U. gemmatus Berk. & M.A. Curtis. N. U. corollocarpi
W.T. Dale. O. U. cucullatus
Syd. P. U. commelinae Cooke. Q. U. sonorensis J.F. Hennen & Cummins.
R. U. socius Arthur & Cummins.
S. U. bidenticola Arthur. T. U. erythronii Pass.

176
 ZAGHOUANIA Pat.
Bull. Soc. Mycol. Fr. 17:182-188 (187). 1901.

FAMILY: Pucciniaceae REMARKS:

Two species are recognized. The type has erum-
DESCRIPTION: pent telia; Z. oleae (E.J. Butler) Cummins, reported
Spermogonia subepidermal, Group V (type 4). from India, has suprastomatal telia. Zaghouania oleae
Aecia subepidermal, erumpent, with peridium, often is treated as Cystopsora oleae E.J. Butler (Butler
Aecidium-ty pe, or without peridium, Caeoma-type in 1910), but maintenance of two genera does not seem
cuplike cavities; aeciospores catenulate in both justified. Zaghouania phillyreae Pat. has four-celled
types. Uredinia subepidermal, erumpent, Uredo- basidia, but Z. oleae has only two-celled basidia.
type; urediniospores borne singly, echinulate. Telia
subepidermal, either rupturing the epidermis or REFERENCES:
developing suprastomatally; teliospores 1-celled, Butler, E.J. 1910. A new genus of the Uredinaceae. Ann. Mycol.
borne singly on pedicels, wall thick, pale, and 8:445-448.
verrucose; germination occurs without dormancy; Patouillard, N. 1901. Champignons Algéro-Tunisiens nouveaux
basidia external, short, broad, and rather thick- ou peu connus. Bull. Soc. Mycol. Fr. 17:182-188.
walled; basidiospores sessile, not ejected.

TYPE: Zaghouania phillyreae Pat.

On: Phillyrea media L. (Oleaceae)
Type locality: Tunisia

FIGURE 129. Zaghouania. Z. oleae (EJ. Butler) Cummins. A. A telium. B. Germinating teliospores.
 ENDOPHYLLUM Lévy.
Mém. Soc. Linn. Paris 4:202—212 (208). 1826.

FAMILY: Pucciniaceae therefore they are pseudogenera or genera of con-

venience. Many times the parental genera cannot be
DESCRIPTION:
predicted, and calling them by anamorphic generic
Spermogonia variable or wanting. Aecia and names also creates confusion. In this book endo-
uredinia not produced. Telia with a peridium, cyclic genera are recognized, but are grouped
morphologically Aecidium; teliospores 1-celled,
closely with the probable parental genera rather
catenulate, like Aecidium spores, except producing a
than in a single group. Buriticd and Hennen (1980)
basidium when germinating. treated Endophyllum within the Pucciniosireae,
together with such genera as Alveolaria, Cionothrix,
TYPE: Endophyllum sempervivi (Alb. & Schwein.)
Didymopsora, and Dietelia, but Buritica (1991) later
de Bary
put all endocyclic genera in the family
=Uredo sempervivi Alb. & Schwein.
Endophyllaceae Dietel. Both of these treatments
On: Sempervivum globiferum L. (Crassulaceae)
create unnatural groupings of the genera involved.
Type locality: Germany
In several species of Puccinia (P. pampeana Speg. and
REMARKS: P. japonica Dietel), two types of spores are known to
Endophyllum is composed of microcyclic species germinate as teliospores (see “Abnormal life cycles”
that simulate the anamorph genus Aecidium. In in the Introduction).
theory, the species may be derived from any macro-
REFERENCES:
cyclic or demicyclic rust fungus that has Aecidium-
Buritica, P. 1991. Familias del orden Uredinales con ciclo de vida
type anamorphs and usually with type 4 spermogo- completamente reducido. Rey. Acad. Colomb. Cienc. Exactas
nia. Previously, Monosporidium (Kulkarniella) was Fis. Nat.18:131-148.
recognized separately, having type 7 or 5 spermogo- Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
nia. In view of the absence of spermogonia in many Pucciniaceae). Flora Neotrop., Monogr. 24:1-50.
endo-species, we recognize Monosporidium as a Léveillé, J.H. 1826. Observations sur deux champignons de la
famille des Urédinées. Mém. Soc. Linn. Paris 4:202-212.
synonym of Endophyllum. The endo forms are really
life cycle variants of macrocyclic species, and

FIGURE 130. Endophyllum. Two types of germination (schemati

c).

178
 ALVEOLARIA Lagerh.
Ber. Dtsch. Bot. Ges. 9:344—348 (346-347). 1891 (issued 1892).

FAMILY: Pucciniosiraceae arrangement of single layers of teliospores in plates

is unique and gives no clue to the relationship of
DESCRIPTION: the genus. In A. cordiae the telia are surrounded by
Spermogonia subcuticular, flat, Group VI (type very long hypertrophied cells of the mesophyll. A
7). Aecia and uredinia not produced. Telia subepi- third species, A. dugetiae Viégas, described on
dermal, deep-seated, becoming erumpent as short Annonaceae, is now known to belong to Dietelia
columns of strongly adherent radial plates, the (Buritica and Hennen 1980).
plates rather easily separable; teliospores 1-celled, Buritica and Hennen (1980) have described both
germ pore 1, apical; germination occurs without subcuticular (epiphyllous with spermatia) and sub-
dormancy; basidia external.
epidermal spermogonia in A. cordiae, but we ques-
tion the nature of the subepidermal structures.
TYPE: Alveolaria cordiae Lagerh. (Lectotype)
On: Cordia sp. (Boraginaceae) REFERENCES:
Type locality: Ecuador Buritica, P.; Hennen, J.-F. 1980. Pucciniosireae (Uredinales-
Pucciniaceae). Flora Neotrop., Monogr. 24:1-54.
REMARKS: Lagerheim, L.G. 1891 (issued 1892). Pucciniosira, Chrysopsora,
Only the type and A. andina Lagerh. are Alveolaria und Trichopsora, vier neue Uredineen-Gattungen
recognized; both parasitize species of Cordia. The mit tremelloider Entwicklung. Ber. Dtsch. Bot. Ges. 9:344-348.

nih
GQ) \

FIGURE 131. Alveolaria. A. cordiae Lagerh. A. Telium. B. Surface view of teliospores from above. C. Germinating
teliospore.

AD
 BAEODROMUS Arthur
Ann. Mycol. 3:18-20 (19). 1905.

FAMILY: Pucciniosiraceae J.F. Hennen) (Buritica 1998). The relationship of the

genus is obscure. Azbukina (1974) established a
DESCRIPTION: tribe, Baeodromeae, for this genus, but Buritica and
Spermogonia subepidermal, Group V (type 4). Hennen (1980) do not think that the Russian and
Aecia and uredinia not produced. Telia sub- Chinese species, B. tranzschelii Azbukina on Urtica
epidermal in origin, erumpent, compact; teliospores Inetevirens Max. (Azbukina 1974; Zhuang and Wei
1-celled, catenulate in adherent chains of a few
1992), is related to other species in the genus.
spores each, or irregularly arranged with or without
intercalary cells, wall mostly lightly pigmented, REFERENCES:
germ pore obscure, probably apical, perhaps not Arthur, J.C. 1905. Baeodromus
holwayi Arth., a new uredineous
differentiated; basidia external. fungus from Mexico. Ann. Mycol. 3:18-20.
Azbukina, Z.M. 1974. Rzhavchinnye griby Dalnego Vostoka.
TYPE: Baeodromus holwayi Arthur (Lectotype) [Rust fungi of the Soviet Far East.] Akad. Nauk, Moscow.
526 p. (p. 149).
On: Senecio cinerarioides Kunth (Asteraceae)
Buritica, P. 1998. La familia Phakopsoraceae en el neotrépico. II.
Type locality: Mexico
Rev. Acad. Colomb. Cienc. Exactas Fis. Nat. 22(84):325-334.
Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
REMARKS:
Pucciniaceae). Flora Neotrop. Monogr. 24:1-50.
Seven species have been described and all are Hernandez, J.R. 2000.Baeodromus ranunculi, a new rust on
microcyclic, occurring on Senecio (Asteraceae) (four Ranunculus from Argentina, and a synopsis of Bacodromus.
species), Eupatorium (Asteraceae) (one species), Mycotaxon 76:329-336.
Ranunculus (Ranunculaceae) (one species), and Zhuang, J-Y.; Wei, S-X. 1992. Materials for study on rust fungi of
Urtica (Urticaceae) (one species) (Hernandez 2000). Eastern Rimland of Qinghai-Xizang (Tibet) Plateau. II.
Baeodromus, an unrecorded genus in China. Mycosystema
Baeodromus dominicana F. Kern has been transferred 5:131-133.
to Uredopeltis (U. dominicana (F. Kern) Buritica &

FIGURE 132. Baeodromus. Cross section of telia of B. senecionis Syd. (courtesy

of Dr.
R. Berndt).

180
 CERATOCOMA Buritica & J.F. Hennen
Rev. Acad. Colomb. Cienc. 18:131-148 (146). 1991.

FAMILY: Pucciniosiraceae REMARKS:

One other species, C. guineensis (Vienn.-Bourg.)
DESCRIPTION: Buritica on Xylopia aethiopica A. Rich. (Annonaceae)
Spermogonia subepidermal, Group V or VI (type from Africa, is known. The genus differs from
4 or 5). Aecia and uredinia lacking. Telia subepi- Cionothrix and Baeodromus in having intercalary
dermal, deep-seated, becoming erumpent as brown, cells and from Dietelia in lacking a peridium.
compact columns of adherent spores; teliospores
catenulate, with intercalary cells and an apparently REFERENCE:
common matrix, 1-celled, germ pore not seen, ger- Buritica, P. 1991. Familias del orden Uredinales con ciclo de vida
minating without dormancy; basidia external. completamente reducido. Rev. Acad. Colomb. Cienc. Exactas
Fis. Nat. 18 (69):131-148.
TYPE: Ceratocoma jacksoniae (Henn. ex McAlpine)
Buritica & J.F. Hennen
=Cronartium jacksoniae Henn.
On: Gompholobium latifolium Sm. (Fabaceae)
Type locality: New South Wales, Australia

FIGURE 133. Ceratocoma. C. jacksoniae (Henn. ex McAlpine) Buritica &

J.F. Hennen. A. Teliospores. B. Habit of C. jacksoniae on the
host plant, showing hairlike telial columns.

181
 CHARDONIELLA F. Kern
Mycologia 31:373-375 (375). 1939.

FAMILY: Pucciniosiraceae REMARKS:

The telia are like those of Cronartium but the
DESCRIPTION: spores are Uromyces-like. The columns are hard
Spermogonia subepidermal, Group V (type 4). when dry. Buriticaé and Hennen (1980) recognize
Aecia and uredinia not produced. Telia subepi- four species, all on Asteraceae and all in South
dermal, becoming erumpent as hairlike columns of America. Further, they consider that the spores orig-
strongly adherent teliospores, the spores in no par- inate in chains with intercalary cells that elongate to
ticular order in the column, 1-celled on long pedicel- simulate pedicels. Trichopsora is similar, but has an
like intercalary cells, germ pore 1 near apex, wall internal basidium.
pale; germination occurs without dormancy;
basidia external. REFERENCES:
Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
TYPE: Chardoniella gynoxidis F. Kern Pucciniaceae). Flora Neotrop., Monogr. 24:1-50.
On: Gynoxys sp. (Asteraceae) Kern, ED. 1939. Chardoniella—a new genus of the Uredinales.
Type locality: Near Bogota, Colombia Mycologia 31:373-375.

FIGURE 134. Chardoniella. A. Cross section of a telium of C. andina (Lagerh.) Buritica

& J.P. Hennen (courtesy of
Dr. R. Berndt). B. Germinating teliospores of C. gynoxidis. C. Teliospores of C. gynoxidis
F. Kern
(reprinted with permission, from Kern 1939)

182
 CIONOTHRIX Arthur
N. Am. Flora 7(2):83-160 (124). 1907.

FAMILY: Pucciniosiraceae REMARKS:

Three species, all neotropical and all microcyclic,
DESCRIPTION: are recognized by Buriticd and Hennen (1980).
Spermogonia subepidermal, Group V (type 4). Cionothrix is generally similar to Skierka, but differs
Aecia and uredinia unknown, probably not pro- in the form of the teliospores and the type of sper-
duced. Telia subepidermal, with peripheral mogonium. Some species, including the type, were
paraphyses; teliospores originally catenulate, soon originally described in Cronartium, but it is unlikely
irregularly arranged and extruded in long flexuous, that they are microcyclic species of that genus. The
hairlike columns of adherent spores, spores 1- relationship of the genus to other genera is obscure,
celled, germ pore perhaps not differentiated; germi- but all three recognized species parasitize the
nation occurs without dormancy by elongation of Asteraceae.
the spore apex to form the basidium; basidium
external. REFERENCES:
Arthur, J.C. 1907. N. Am. Flora 7(2):83-160.
TYPE: Cionothrix praelonga (G. Winter) Arthur Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
=Cronartium praelongum G. Winter Pucciniaceae). Flora Neotrop., Monogr. 24:1—50.
On: Eupatorium sp. (Asteraceae)
Type locality: Brazil

FIGURE 135. Cionothrix. Teliospores of C. praelonga

(G. Winter) Arthur.

183
 DIDYMOPSORA Dietel
Hedwigia 38:248-259 (254-255). 1899.

FAMILY: Pucciniosiraceae Americas on Asteraceae, Melastomataceae,

Solanaceae, and Tiliaceae (Buriticé and Hennen
DESCRIPTION: 1980). Cunningham (1968) transferred Cronartium
Spermogonia subepidermal, Group V (type 4). paraguayensis Speg. to Didymopsora (D. paraguayensis
Aecia and uredinia not produced. Telia subepi-
(Speg.) J.L. Cunn.). He clearly indicated that the
dermal, erumpent, domelike or short columnar,
two-celled teliospores are connected by intercalary
somewhat like Aecidium in appearance but without
cells as in other Didymopsora species (D. chu-
peridium; teliospores 2-celled by transverse septa,
quiraguae Dietel and D. triumfettae H.S. Jacks. &
catenulate in loosely adherent chains, intercalary
Holw.), unlike the one-celled teliospores of
cells present but often seen only basally, wall pale,
Cronartium. Two macrocyclic species reported from
germ pores obscure but probably 1 per cell; germi-
India, D. toddaliae Thirum. & Mundkur and D.
nation occurs without dormancy; basidia external.
macrospora Mundkur & Thirum., are now placed in
TYPE: Didymopsorella because of the pedicellate teliospores
Didymopsora solani-argentei (Henn.) Dietel
and Group VI spermogonia.
(Lectotype, Arthur 1906)
=Aecidium solani-argentei Henn. REFERENCES:
On: Solanum argenteum Dunal ex Poiret Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
(Solanaceae) Pucciniaceae). Flora Neotrop., Monogr. 24:1-50.
Type locality: Brazil Cunningham, J.L. 1968. Ontogeny of teliospores of Cronartium
paraguayense and relationship to Didymopsora. Mycologia
REMARKS: 60:769-755.
Didymopsora and Pucciniosira differ only in the Dietel, P. 1899. Uredineae brasiliensis a.cl. E. Ule lectae. Hedwigia
presence of a peridium in the latter. There are six 38:248-259,
species occurring in warm regions of Africa and the

FIGURE 136. Didymopsora. A telium of D. africana

Cummins.

184
 DIETELIA Henn.
Hedwigia 36:190—248 (215-216). 1897.

FAMILY: Pucciniosiraceae peridium tends to adhere to the spore column.

Endophyllum holwayi H.S. Jacks. has been treated by
DESCRIPTION: Lindquist (1970) as the monotypic genus Jack-
Spermogonia subepidermal, Group V (type 4). soniella, but it has the characteristics of Dietelia, ac-
Aecia and uredinia not produced. Telia subepi- cording to Buritica and Hennen (1980). An African
dermal, erumpent, with peridium, Aecidium-like species, D. codiaei Boerema et al. on Codiaeum spp.
but compact; teliospores 1-celled, catenulate, with (Euphorbiaceae), has been discovered recently in
intercalary cells basally but often not seen, wall pale several European countries on introduced orna-
in most species, smooth or finely verrucose, germ mental plants from Africa (Boerema et al. 1994).
pore 1 where seen, mostly obscure; basidia external.
REFERENCES:
TYPE: Dietelia verruciformis (Henn.) Henn. Boerema, G.H.; Cook, R.T.A.; Dickens, J.S.W.; Pisi, A.; Bellardi,
=Cronartium verruciforme Henn. M.G.; Mordue, J.E.M. 1994. Codiaeum rust. Plant Pathol.
On: Sida flavescens Cav. (Malvaceae) 43:1072-1079.
Type locality: Argentina Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
Pucciniaceae). Flora Neotrop., Monogr. 24:1-50.
REMARKS: Gjeerum, H.B. 1985. East African rusts (Uredinales), mainly from
Eight species are known, five from the Americas, Uganda on families belonging to Apetalae and Polypetalae.
Mycotaxon 24:237-273.
two from Africa, and one from the Philippines on
Hennings, P. 1897. Beitrage zur Pilzflora Sidamerikas. II.
dicotyledonous plants (Boerema et al. 1994; Buritica Hedwigia 36:190-248.
and Hennen 1980; Gjeerum 1985). The telia of this Lindquist, J.C. 1970. Notas uredinologicas. Rev. Fac. Agron. Univ.
genus appear like Aecidium but are compact and the Nac. La Plata 46:199-205.

FIGURE 137. Dietelia. A telium of D. verruciformis

(Henn.) Henn.

185
 PUCCINIOSIRA Lagerh.
Ber. Dtsch. Bot. Ges. 9:344-348 (344-345). 1891 (issued 1892).

FAMILY: Pucciniosiraceae REMARKS:

The genus is composed of 17 species, 7 of which
DESCRIPTION: are neotropical. Buritica and Hennen (1980) treat
Spermogonia subepidermal, Group V (type 4) or Gambleola as a synonym. This seems radical at first
lacking. Aecia and uredinia not produced. Telia thought, because of the strongly adherent teliospores
subepidermal, erumpent, mostly Aecidium-like with
and long dark telial columns of Gambleola, but the
peridium, but compact and hornlike in some
teliospore morphology is similar. Species of
species, teliospores 2-celled by transverse septum,
Pucciniosira, including species formerly recognized
catenulate, with intercalary cells, loosely or firmly
in Gambleola, occur on Berberis (Berberidiaceae) and
united laterally and longitudinally, wall pale or
other dicotyledonous hosts, including Asteraceae,
obviously pigmented, smooth or verrucose, the cells
Solanaceae, and Tiliaceae.
separating easily, germ pores obscure; basidia
external. REFERENCES:
Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
TYPE: Pucciniosira triumfettae Lagerh. (Lectotype) Pucciniaceae). Flora Neotrop., Monogr. 24:1-50.
=P. pallidula (Speg.) Lagerh. (according to Lagerheim, G. von. 1891 (issued 1892). Pucciniosira, Chrysopsora,
Buritica and Hennen 1980) Alveolaria und Trichopsora, vier neue Uredineen-Gattungen
mit tremelloider Entwicklung. Ber. Dtsch. Bot. Ges. 9:344-348.
On: Triumfetta sp. (Tiliaceae)
Type locality: Ecuador

4
=\\ es
\
<—s =
ee
dscSae

=ane
~>

lGneee leeel
reer
S

|eA‘co
=7

Lo“—
RS ZI

ae
[|
ow)

Jos

FIGURE 138. Pucciniosira. A. Telial horns of P. cornuta (Massee) Buritica &

J.F. Hennen (=Gambleola cornuta Massee).
B. Teliospores of P. cornuta. C. A telium of P. pallidula (Speg.) Lagerh.

186
 TRICHOPSORA Lagerh.
Ber. Dtsch. Bot. Ges. 9:344-348 (347). 1891 (issued 1892).

FAMILY: Pucciniosiraceae REMARKS:

Only the type is known and it occurs in South
DESCRIPTION: America. The telial column superficially resembles
Spermogonia subepidermal, Group V (type 4). that of Cronartium, but the germination is different.
Aecia and uredinia not produced. Telia subepider- Coleosporium crowellii Cummins lacks the intercalary
mal, becoming erumpent as more or less gelatinous, cells, but otherwise is similar in appearance. The
hairlike columns; teliospores 1-celled, catenulate, relationship of the genus is obscure, but Buritica and
with elongating and gelatinizing intercalary cells, Hennen (1980) indicate a relation to Chardoniella.
becoming basidia by division of the protoplast into 4
cells, each of which produces 1 basidiospore; germi- REFERENCES:
nation occurs without dormancy. Buritica, P.; Hennen, J.F. 1980. Pucciniosireae (Uredinales,
Pucciniaceae). Flora Neotrop., Monogr. 24:1-50.
TYPE: Trichopsora tournefortiae Lagerh. Lagerheim, C. von. 1891 (issued 1892). Pucciniosira, Chrysopsora,
Alveolaria und Trichopsora, vier neue Uredineen-Gattungen
On: Tournefortia sp. (Boraginaceae)
mit tremelloider Entwicklung. Ber. Dtsch. Bot. Ges. 9:344-348.
Type locality: Ecuador

sila
Aisi
Pease
fa
4-
ees
Siete
=

OEP
/-
SV
oflEPGi
TE —
Vmkey
Oy
HG
SER
eaYA
pp OTe?
y(EE
=sueBALL
yLs
t /

ha
vf.
qh

FIGURE 139. Trichopsora. T. tournefortiae Lagerh. A. A telium (courtesy of Dr. R. Berndt.). B. Germinating teliospores.

187
 HIRATSUKAMYCES Thirum., F. Kern & B.V. Patil
Sydowia Ann. Mycol. (Ser. II) 27:78-81 (80-81). 1973 (published 1975).

FAMILY: Uncertain REMARKS:

Two species (H. salacicola and H. divinum Thirum.,
DESCRIPTION: FE. Kern & B.V. Patil) have been recognized; Caeoma
Spermogonia subepidermal, flask shaped,
callianthum Syd. probably also belongs to the genus.
Group III (type 12). Aecia subepidermal, 3 or 4 cells The aecial states may be localized or cover the entire
deep in the mesophyll, without a peridium but with surface of leaves, causing first an elevation of the
dermal layers consisting of hyphae fused with host host tissue in somewhat cerebroid patterns, then
cells, becoming exposed by extensive rupture of the rupturing it to expose the spores. Under appropriate
epidermis and extruding spores interspersed with conditions the spores and elaters hang from the
elater hyphae (Elateraecium-type); aeciospores sorus. Because of these unique features, Thirumala-
catenulate but separating easily, verrucose. char et al. (1966) described Elateraecium as a new
Uredinia subepidermal, erumpent, with a periph-
aecial type before finding uredinial and telial states.
eral dermal layer, Milesia-type; urediniospores pro- The species are autoecious and parasitize species of
duced in succession on long pedicels, verrucose. Salacia of the Celastraceae.
Telia intraepidermal, not erumpent; teliospores
sessile, l-celled,2 to5inan epidermal cell; germina- REFERENCES:
tion unknown; basidia probably external. Thirumalachar, M.J.; Kern, E.D.; Patil, B.V. 1966. Elateraecium, a
new form genus of Uredinales. Mycologia 58:391-396.
TYPE: Hiratsukamyces salacicola Thirum., F. Kern & Thirumalachar, M.J.; Kern, ED.; Patil, B.V. 1973. Hiratsukamyces, a
Bvebatl: new genus of the Pucciniastreae. Sydowia Ann. Mycol. (Ser.
II) 27:78-81.
(spelled salicicola in the original description)
On: Salacia prinoides DC. (Celastraceae)
Type locality: India

FIGURE 140. Hiratsukamyces. H. salacicola Thirum., F. Kern & B.V. Patil. A. Aeciospores and an elater
hypha.
B. Intraepidermal teliospores.

188
 MASSEEELLA Dietel
Ber. Dtsch. Bot. Ges. 13:332-—335 (332-333). 1895.

FAMILY: Uncertain Euphorbiaceae. Insofar as known, all species are

macrocyclic and autoecious. Masseeélla narasimhanii
DESCRIPTION: Thirum. is the only reported species with caeoma-
Spermogonia subcuticular, Group VI (type 7). toid aecia. This difference was used by Sathe (1966)
Aecia subepidermal, becoming erumpent, with as the basis for the genus Kamatomyces, which we
peridium (Aecidium-type) or without peridium treat as a synonym here. When Thirumalachar
(Caeoma-type); aeciospores catenulate, verrucose. (1943) described M. narasimhanii, he commented
Uredinia subepidermal, small, slightly erumpent, that the structure and development of the telio-
Uredo-type; urediniospores borne singly, echinulate, spores of all known species of Masseeélla show
pores obscure. Telia subepidermal, deep-seated, remarkable resemblance.
lined with upwardly directed hyphae, which may
secrete the gelatinous matrix in which the spores are REFERENCES:
embedded and extruded in hairlike columns; telio- Dietel, P. 1895. Drei neue Uredineen Gattungen, Masseeélla,
spores 1-celled, sessile, without order in the matrix, Phakopsora u. Schizospora. Ber. Dtsch. Bot. Ges. 13:332-335.
not adherent, wall pigmented, germ pore 1; Sathe, A.V. 1966. Revision of Masseeélla narasimhanii Thirum.
basidium external. (Uredinales). Sydowia 19:187-189.
Singh, H.; Singh, B.V. 1967. On some Indian species of Massceella.
Mycopathol. Mycol. Appl. 33:193-199.
TYPE: Masseeélla capparidis (Hobson) Dietel
Thirumalachar, M.J. 1943. Masseeélla narasimhanii, a new species
=Cronartium capparidis Hobson of rust on Flueggia leucopyrus Willd. Proc. Indian Acad. Sci.
On: Flueggea virosa Willd.) Voigt (Euphorbiaceae) Sect. B, 18:36-40.
Type locality: India

REMARKS:
All four species occur in warm regions from
India to the Philippines and all on plants of the

peer
sale.
ello
eee)
7

eer
FIGURE 141. Masseeélla. A. A telium of M. capparidis (Hobson) Dietel (courtesy of Dr. R. Berndt). B. Teliospores of
M. flueggiae Syd.

189
 SKIERKA Racib.
Parasit. Algen Pilze Javas. Pt. 2. 46 p. (30-31). 1900.

FAMILY: Uncertain REMARKS:

There are about 10 circumtropical species (Mains
DESCRIPTION: 1939). All are autoecious and presumably macro-
Spermogonia subepidermal, Group VI (type 5). cyclic, but only telia have been described for S.
Aecia subepidermal, opening by a porelike rupture robusta Doidge. The species occur about equally on
of the epidermis, without peridium, Uredo-type; Geraniales and Sapindales, with one species on
aeciospores borne singly. Uredinia subepidermal, Rhamnales. Skierka differs from Cionothrix in the
similar to the aecia except not associated with sper- type of spermogonium.
mogonia, Uredo-type; urediniospores similar to the
aeciospores, the wall thickened in two opposite, lon- REFERENCES:
gitudinal, somewhat hygroscopic bands or ridges, Mains, E.B. 1939. The genera Skierka and Ctenoderma. Mycologia
the crests of which may be echinulate or crenulate. 31:175-190.
Telia subepidermal, as the uredinia; teliospores Purkayastha, R.P.; Pal, A.K. 1998. SEM studies on a mangrove
1-celled, sessile, produced in irregular succession, rust of Sundarbans, Eastern India. Mycol. Res. 102:692-694.
Raciborski, M. 1900. Parasitische Algen und Pilze Javas. Pt. 2.
strongly adherent and extruded in long, hairlike
Hrsg. Bot. Inst. Buitenzorg, Batavia. 46 p.
columns, germ pore not obvious but doubtless 1 per
cell; germination occurs without dormancy; basidia
external.

TYPE: Skierka canarii Racib.

On: Canarium commune L. (Burseraceae)
Type locality: Java, Indonesia

FIGURE 142. Skierka. A. A telium of S. cristata (Speg.) Mains (courtesy of Dr. R. Berndt). B. Urediniospor
es of
S. holwayi Arthur. C. A telium of S. holwayji.

190
 APPENDIX 1
REFERENCE MATERIALS
(with some annotations)

oroS

GENERAL REFERENCES Petersen, R.H. 1974. The rust fungus life cycle. Bot. Rev. 40:453—
Sle.
ON RUST FUNGI
Roelfs, A.P.; Bushnell, W.R., editors. 1985. The cereal rusts. Vol. II.
Diseases, distribution, epidemiology, and control. Academic
Alexopoulos, C.J.; Mims, C.W.; Blackwell, M. 1996. Introductory
Press, London. xxii + 605 p.
mycology. John Wiley & Sons, Inc., New York. 868 p. Has an
excellent chapter on rust fungi. Schumann, G.L. 1991. Plant diseases: their biology and social
impact. APS Press, St. Paul, MN. 397 p.
Azbukina, Z.M. 1974. [Rust fungi of Far East]. Nauka, Moscow.
527 p. (In Russian.) Introduction provides a good general Scott, K.J.; Chakravorty, A.K., editors. 1982. The rust fungi.
account of rust fungi. Academic Press, London. 288 p.
Swann, E.C.; Frieders, E.M.; McLaughlin, D.J. 2001. 2.
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East.] Nauka, Moscow. 288 p. (In Russian.)
Urediniomycetes. Pages 37-54 in D.J. McLaughlin and P.
Lemke, eds. The Mycota. Vol. VIIB. Systematics and evolution.
Bauer, R.; Begerow, D.; Oberwinkler, F.; Piepenbring, M.; Berbee,
Springer-Verlag, Berlin. The most up-to-date taxonomic review
M.L. 2001. Ustilaginomycetes. Pages 57-83 in D.J. of Urediniomycetes (including rusts) within Basidiomycota.
McLaughlin and P. Lemke, eds. Mycota. Vol. VIIB. Springer-
Verlag, Berlin. The most up-to-date information and discussion
regarding phylogenetic relationships of Urediniomycetes
(including rusts) and Ustilaginomycetes. DESCRIPTIVE MANUALS AND
Buller, A.H.R. 1950. Researches on fungi. Vol. VII. The sexual pro- REGIONAL FLORA
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Dietel, P. 1928. Uredinales. In A. Engler and K. Prantl: Die
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Ito, S. 1938. [Mycological flora of Japan.] Vol. 2. Basidiomycetes. 3. J. Bot. 67:3315-3365.
Uredinales—Pucciniaceae, Uredinales Imperfecti. Yokendo, Parmelee, J.A. 1996. A catalogue of types (Uredinales) in the
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183.
(1973-1990) Uredinales (rusts~rouilles). Natl. Mycol. Herb.,
Kaneko, S, 1981. The species of Coleosporium, the causes of pine Agric. Agri-Food Can., Ottawa. ON. A compilation of 89
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Roure, L.A. 1963. The rusts of Puerto Rico. Univ. Puerto Rico
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LoS
 APPENDIX 2
AUTHORS OF NAMES OF RUST FUNGI
AND SUGGESTED ABBREVIATIONS

“&

If: fungal nomenclature, citing the name of the Bacc.

author or authors who first described a fungus Baccarini, Pasquale
ensures precision in scientific publications. Such (1858-1919)
citations are important in tracking the history ofa Bagyan. (Bhagyan.)
name, and thus the full identity of the taxon. It is Bagyanarayana (Bhagyanarayana), Gaddam
often necessary to abbreviate authors’ names for (1953— )
convenience and to save space; however, there is B.K. Bakshi
great variability in the author abbreviations used by Bakshi, Bimal Kumar
different writers. This can lead to confusion when (1919- )
persons with the same or similar names have
Barclay
described new species. Therefore in the list below
Barclay, Arthur
we have used (with permission) the abbreviations
(1852-1891)
recommended by Kirk and Ansell (1992) for authors
Barthol.
of names of rust fungi. We have also added many
names not included in that list. The dates of birth
Bartholomew, Elam
and death of an author frequently help in tracing the (1852-1934)
origin of fungal names, and we have updated this J.W. Baxter
information when possible. When these dates are Baxter, John Wallace
not known, a year when the author is known to have (1918—_ )
published a description is indicated. Benth.
Bentham, George
(1800-1884)
Adans. Berk.
Adanson, Michel Berkeley, Miles Joseph
(1727-1806) (1803-1889)
Alb. R. Berndt
Albertini, Johannes Baptista von Berndt, Reinhard
(1769-1831) (1960— )
F.C. Albuq. Bert.
Albuquerque, Fernando Carneiro de Bertero, Carlo Luigi Guiseppe
(1932— ) (1789-1831)
F.W. Anderson Bethel
Anderson, Frederick William Bethel, Ellsworth
(1866-1891) (1863-1925)
H.W. Anderson Bisby
Anderson, Harvey Warren Bisby, Guy Richard
(1885-1971) (1889-1958)
Arthur Boedijn
Arthur, Joseph Charles Boedijn, Karel Bernard
(1850-1942) (1893-1964)
Azbukina Broome
Azbukina, Zinaida M. Broome, Christopher Edmond
(1925-— ) (1812-1886)

194
Bubak G. Cunn.
Bubak, Frantis ek Cunningham, Gordon Herriott (Heriot)
(1865-1925) (1892-1962)
Burdsall J.L. Cunn.
Burdsall, Harold H. Cunningham, John L.
(1940- ) (published 1964 )
Buritica M.A. Curtis
Buritica, Pablo Curtis, Moses Ashley
(1943-— ) (1808-1872)
E.J. Butler Curtis
Butler, Edwin John Curtis, William
(1874-1943) (1746-1799)

W.T. Dale
DC.
Dale, William Thomas
Candolle, Augustin Pyramus de
(1918- )
(1778-1841)
Davis
A.A. Carvalho
Davis, John Jefferson
Carvalho Junior, Anibal Alves de
(1852-1937)
(1960-_)
de Bary
A.O. Carvalho
de Bary, Heinrich Anton
Carvalho, Aldir de Oliveira de
(1831-1888)
(1959— )
De Toni
Castagne
De Toni, Giovanni Batista
Castagne, Jean Louis Martin
(1864-1924)
(1785-1858)
Desm.
M. Chen
Desmazieres, John Baptiste Henri Joseph
Chen, Mo Mei
(1786-1862)
(1930-— )
Dianese
Chevall. Dianese, José Carmine
Chevallier, Francois Fulgis (1940- )
(1796-1840) Dietel
Chou Dietel, Paul
Ghouls (1860-1947)
(published 1954) Doidge
Citi, Doidge, Ethel Mary
Ciferri, Raffaele (1887-1965)
(1897-1964) Duby
Clem. Duby, Jean Etienne
Clements, Frederick Edward (1798-1885)
(1874-1945) Dudley
Cooke Dudley, William Russel
Cooke, Mordecai Cubitt (1849-1911)
(1825-1914) Durrieu
Corda Durrieu, Guy
Corda, August Karl Joseph (1931-— )
(1809-1872)
P.E. Crane Eboh
Crane, Patricia Ellen Eboh, Dan O.
(1948— ) (1937-—_)
Cummins Ehrenb.
Cummins, George Baker Ehrenberg, Christian Gottfried
(1904— ) (1795-1876)

o>
Ellis Gaum.
Ellis, Job Bicknell Gaumann, Ernest (Ernst) Albert
(1829-1905) (1893-1963)
Engl. Gilb.
Engler, Heinrich Gustav Adolf Gilbertson, Robert Lee
(1844-1930) (1925— )
Espinosa Gjerum
Espinosa Bustos, Marcial Ramon Gjeerum, Halvor B.
(1874-1959) Ghee)
Gokhale
Farl. Gokhale, V.P.
Farlow, William Gilson (published 1951)
(1844-1919) Gonz. Frag.
Faull Gonzalez Fragoso, Romualdo
Faull, Joseph Horace (1862-1928)
(1876-1961)
Gray
F.A. Ferreira
Gray, Samuel Frederick
Ferreira, Francisco (Xiku) Alves
(1766-1828)
(1950—_ )
Grev.
Figueiredo
Greville, Robert Kaye
Figueiredo, Mario Barreto
(1794-1866)
(1933- )
Guo
E. Fisch.
Guo, L.
Fischer, Eduard
(published 1986)
(1861-1939)
J.C. Fisch.
Fischer, Johann Carl G. Hahn
Hahn, Gotthold
(1804-1885)
W.P. Fraser (published 1875-1911)
Fraser, William Pollock Hara
(1867-1943) Hara, Kanesuke
F.O. Freire (1885-1962)
Freire, Francisco Das Changas Oliveira Harada
(1947- ) Harada, Yukio
Fr. (1939-—_ )
Fries, Elias Magnus Hashioka
(1794-1878) Hashioka, Yoshio
Fromme (1911- )
Fromme, Fred Denton Hedge.
(1886-1966) Hedgcock, George Grant
Fuckel (1863-1946)
Fuckel, Karl Wilhelm Gottlieb Leopold R. Hedwig
(1821-1876) Hedwig, Romanus (Romanes) Adolf
Furlanetto (1772-1806)
Furlanetto, Cleber J.F. Hennen
(1966—_ ) Hennen, Joe Fleetwood
(1928- )
Galloway Henn.
Galloway, Beverly Thomas Hennings, Paul Christoph
(1863-1938) (1841-1908)
D.E. Gardner Hernandez
Gardner, Donald E. Hernandez, Jose
(1942— ) (1963- )

196
N. Hirats. J.R. Johnst.
Hiratsuka, Naoharu Johnston, John Robert
(1873-1946) (1880-1953)
Hirats. f. Jorst.
Hiratsuka, Naohide Jorstad, Ivar
(1903-2000) (1887-1967)
T. Hirats. Juel
Hiratsuka, Toshiko Juel, Hans Oscar
(1931-_ ) (1863-1931)
Y. Hirats.
Hiratsuka, Yasuyuki Kakish.
(1933- ) Kakishima, Makoto
Hobson (1949— )
Hobson (Major General) Kalchbr.
(published 1886) Kalchbrenner, Karoly (Karl)
Hodges (1807-1886)
Hodges, Charles S. Kamat
(1931- ) Kamat, M.N. (1897-1980)
Hohn. S. Kaneko
Hohnel, Franz Xaver Rudolf von Kaneko, Shigeru (1943-—__)
(1852-1920) Kaps.-Gotsi
Holw. Kapsanaki-Gotsi, Evangelia
Holway, Edward Willet Dorland (1950— )
(1853-1923) Karsten
Hori Karsten, Petter (Peter) Adolf
Hori, Shétar6 (1834-1917)
(1865-1945) Katsuya
Hornem. Katsuya, Keizo
Hornemann, Jens Wilken (1934— )
(1770-1841) Katumoto
Howe Katumoto (Katsumoto), Ken
Howe, Elliot Calvin (1927— )
(1828-1899) Kellerm.
B. Huguenin Kellerman, William Ashbrook
Huguenin, B. (1850-1908)
(published 1967) Kelsey
N.R. Hunt Kelsey, Francis Duncan
Hunt, Nicholas Rex (1849-1905)
(1885-1963) F. Kern
Kern, Frank Dunn
Imazu (1883-1973)
Imazu, Michio Khesw.
(1963-— ) Kheswalla, Kavasji Framaji
S. Ito (published 1941)
Ito, Seiya J.J. Kickx
(1883-1962) Kickx, Jean Jacques
(1842-1887)
H.S. Jacks. Kleb.
Jackson, Herbert Spencer Klebahn, Heinrich
(1883-1951) (1859-1942)
Jacz. Kom.
Jaczewski, Arthur Louis Arthurovié Komarov, Vladimir Leontjevich (Leontevich)
(1863-1932) (1869-1945)

OY,
Korn. Long
Kornicke, Friedrich August Long, William Henry
(1828-1908) (1867-1947)
J.G. Kithn
Kuhn, Julius Gotthelf Magnus
(1825-1910) Magnus, Paul Wilhelm
Kuntze (1844-1914)
Kuntze, Carl (Karl) Ernst (Eduard) Otto Mains
(1843-1907) Mains, Edwin Butterworth
Kunze (1890-1968)
Kunze, Gustav T. Majewski
(1793-1851) Majewski, Tomasz
Kusano (1940- )
Kusano, Shunsuke L. Marchand
(1874-1962) Marchand, Louis
(1807-1843)
Lagerh. Markova
Lagerheim, Nils Gustaf (von, de) Markova, J.
(1860-1926) (published 1987)
Laundon F.A. Mason
Laundon, Geoffrey Frank (Gillian Fiona) Mason, Francis Archibald
(1938-1984) (1878-1936)
Ledingham Massee
Ledingham, George Aleck Massee, George Edward
(1903-— ) (1850-1917)
Leon-Gall. Tak. Matsumoto
Leon-Gallegos (Leén Gallegos) Hector M. Matsumoto, Takashi
(published 1972) (1891-1968)
Leppik Mayor
Leppik (Lepik), Elmar Emil Mayor, Eugéne
(published 1932) (1875-1976)
Lév. Mayr
Léveillé, Joseph-Henri Mayr, Heinrich
(1796-1870) (1856-1911)
B. Li McAlpine
Li, Bin McAlpine, Daniel
(1952- ) (1849-1932)
J.C. Linda. J.W. McCain
Lindquist, Juan Carlos McCain, John William
(1899-1990) (1948-_)
Link Medeiros
Link, Johann Heinrich Friedrich Medeiros, Ricardo Brilhante de
(1767-1851) (1968—_ )
Liro Meinecke
Liro, Johan Ivar (né J.I. Lindroth) Meinecke, Emillio Pepe Michael
(1872-1943) (1869- ? )
Litwinow Merr.
Litwinow, Dimitri Ivanovich Merrill, Elmer Drew
(1854-1929) (1876-1956)
Lohsomb. Milesi
Lohsomboon, Pongvipa Milesi, Marco
(1962- ) (published 1904)

198
Miyabe Oerst.
Miyabe, Kingo Oersted, Anders Sandoe
(1860-1951) (1816-1872)
Mont. Olive
Montagne, Jean Pierre Francois Camille Olive, Edgar William
(1784-1866) (1870-1971)
Moore Y. Ono
Moore, Justin Payson Ono, Yoshitaka
(1841-1923) (1949-—_)
Mordue Onuma
Mordue, Janet Elizabeth Mary Onuma, Fusaji
(1936—") (published 1930)
Moug. Orish.
Mougeot, Jean Baptiste Orishimo, Yoshinobu
(1776-1858) (1881-_)
J.B. Mull. G.H. Otth
Muller, Jean Baptiste (Baptista) Otth, Gustav Heinrich
(1806-1894) (1806-1874)
Mundk.
Mundkur, Bhalchendra Bhavanishankar Paclt
(1896-1952) Paclt, Jiri Jiré)
Muray. (1925- )
Murayama, Daiki Padro-Cardona
(1911-1993) Padro-Cardona, Victor Manuel
(1947- )
P.C. Pandey
Nannf.
Pandey, Poorn Chandra
Nannfeldt, John Johan) Axel Frithiof
(1904-1985) (1938-_)
Parmelee
Nattras
Parmelee, John A.
Nattras (Nattrass), Roland Marshall
(1924— )
(1895-—_—)
Pass.
Naumov
Passerini, Giovanni
Naumov (Naumoff), Nikolai Alexsandrovich
(1816-1893)
(1888-1959)
B.V. Patil
Navashin
Patil, B.V.
Navashin (Nawashin), Sergei Gavrilovich
(published 1975)
(Sergius Gawrilowitsch)
Pat.
(1857-1930)
Patouillard, Narcisse Théophile
Neger
(1854-1926)
Neger, Franz (Friedrich) Wilhelm
Pazschke
(1868-1923)
Pazschke, Franz Otto
Nestl.
(1843-1922)
Nestler, Chrétien Géofroy (Christian Gottfried)
Peck
(1778-1832)
Peck, Charles Horton
Niessl
(1833-1917)
Niessl von Mayendorf, Gustav
Pers.
(1839-1919)
Persoon, Christiaan Hendrik
(1761-1836)
Oehrens R.S. Peterson
Oehrens, B. Edgardo Peterson, Roger Shipp
(published 1987) (1931-— )

199
Petr. P.A. Rodr.
Petrak, Franz Rodriguez, Paola A.
(1886-1973) (1958-_)
Plowr. Rohl.
Plowright, Charles Bagge Rohling, Johann Christoph
(1849-1910) (1757-1813)
Poepp. Rostr.
Poeppig, Eduard Friedrich Rostrup, (Frederik Georg) Emil
(1798-1868) (1831-1907)
K.V. Prasad Roussel
Prasad, K.V. Roussel, Henri Francois Anne de
(published 1993) (1747-1812)
Rudolphi
Rabenh. Rudolphi, (Israel) Karl (Carl) Asmund
Rabenhorst, Gottlob (Gottlieb) Ludwig (Asmunt, Asmus)
(1806-1881) (1771-1832)
Racib.
Raciborski, Marjan (Maryan, Marian, Maryjan) Sacc.
(1863-1917) Saccardo, Pier Andrea
Rajendren (1845-1920)
Rajendren, R.B. Saho
(published 1966) Saho, Haruyoshi
Ramachar (1927-— )
Ramachar, Penugonda T. Sanchez
(1926-1990) Sanchez, T.
K. Ramakr. (published 1994)
Ramakrishnan, K. Santos
(published 1949) Santos, Leila Terezinha Pereira dos
T.S. Ramakr. (1956-_)
Ramakrishnan, Taracad Subromania Sathe
(published 1928) Sathe, A.V.
Ramaley (1935- )
Ramaley, Annette W. S. Sato
(published 1987) Sato, Shoji
Ramsb. (1927- )
Ramsbottom, John Toy. Sato
(1885-1974) Sato, Toyozo
K.N. Rao (1954—_)
Rao, K. Nirnjan Savile
(published 1987) Savile, Douglas Barton Osborne
Rebent. (1909-2000)
Rebentisch, Johann Friedrich Sawada
(1772-1810) Sawada, Kaneyoshi (Kenkichi)
D.A. Reid (1888-1950)
Reid, Derek A. Schltdl.
(1927-— ) Schlechtendal, Diederich Franz Leonhard von
Rezende (1794-1866)
Rezende, Denise Vilela J.C. Schmidt
(Santiago, Denise Rezende) Schmidt, Johann Carl (Karl)
(1958-) (1793-1850)
Roberge Scholler
Roberge, Michel (Michael) Robert Scholler, Marcus
( -1864) (1961-—_ )

200
J. Schrot. I. Takah.
Schroter, Joseph Takahashi, Ikuo
(1837-1894) (IGEQ= ))
C. Schub. Teng
Schubert, Carl Teng, Shu Chun
(published 1820) (1902-1970)
Schultz Thaung
Schultz, Carl (Karl) Friedrich Thaung, Maung Mya
(1765 /6-1837) (published 1973)
Schwein. Thaxt.
Schweinitz, Lewis (Ludwig) David von Thaxter, Ronald
(1780-1834) (1858-1932)
Seym. Thirum.
Seymour, Arthur Bliss Thirumalachar, Mandayani Jeersannidhi
(1859-1933) (1914—_ )
C.G. Shaw C.H. Thomps.
Shaw, Charles Gardner, Jr. Thompson, Charles Henry
(1917-1998) (1870-1931)
C.G. Shaw III Thim.
Shaw, Charles Gardner, III (Terry) Thtimen, Felix (Karl Albert Ernst Joachim) von
(1948— ) (1839-1892)
Shirai Togashi
Shirai, Mitsutaro (‘Kotaro’) Togashi, Kogo
(1863-1932) (1895-1952)
Suj. Singh Tracy
Singh, Sujan Tracy, Samuel Mills
(1930-— ) (1847-1920)
Sotao Tranzschel
Sotao, Helen Maria Pontes Tranzschel, Woldemar (Andrejevitch)
(1963-) (1868-1942)
Speg. Traverso
Spegazzini, Carlo Luigi Traverso, Giovanni Battista
(1858-1926) (1878-1955)
F. Strauss Trotter
Strauss, Friedrich Karl (Carl) Joseph von Trotter, Alessandro
(1787-1855) (1874-1967)
Sullia Tubeuf
Sullia, Shanker Bhat Tubeuf, Carl (Karl) von
(1940-— ) (1862-1941)
Sulmont Tul.
Sulmont, Ph. Tulasne, Louis René (“Edmond’)
(published 1969) (1815-1885)
Syd.
Sydow, Hans S. Uchida
(1879-1946) Uchida, Seinosuke
P. Syd. (published 1960)
Sydow, Paul Ulbr.
(1851-1925) Ulbrich, Oskar Eberhard
(1879-1952)
Fala Underw.
Tai, Fung (Fang) Lan (L.Y.) Underwood, Lucien Marcus
(1893-1973) (1853-1907)

201
Unger X.B. Wu
Unger, Franz (Joseph Andreas Nicolaus) Wu, Xing Bang
(1800-1870) (published 1987)
Z. Urb. Wurth
Urban, Zdenék Wurth, Theophil
(1923-2000) (1875-1922)
Urries
Urries de Azara, M.J. de B.R.D. Yadav
(published 1932) Yadav, B.R. Dayaker
(published 1993)
Viégas G. Yamada
Viegas, Ahmés Pinto Yamada, Gentaro
(1905-1986) (1873-1943)
Vienn.-Bourg. J.M. Yen
Viennot-Bourgin, Georges Yen, Jo Min (published 1971)
(1906- ) Yohem
Yohem, K.H. (published 1985)
Woronin
Zhang
Woronin, Michael Stepanovitch
(1838-1903)
Zhang, Ning
Wallr.
(1972— )
Zhuang
Wallroth, Carl (Karl) Friedrich Wilhelm
Zhuang, Jian Yun
(1792-1857)
(1944— )
Y.C. Wang
Ziller
Wang, Yun Chang
Ziller, Wolf Gunther
(1906- )
(1910-1995)
Wei
Zinno
Wei, Shu-xia
Zinno, Y.
(1939-— )
(published 1972)
Westend.
H. Zogg
Westendorp, Gérard Daniel
Zogg, Hans
(1813-1869)
(published 1949)
Whetzel
Whetzel, Herbert Hice
(1877-1944) LITERATURE CITED
F.B. White Kirk, P.M.; Ansell, A.E. 1992. Authors of fungal names. Index of
White, Francis Buchanan Fungi Supplement. Int. Mycol. Inst., Cab Int., Kew, Surrey,
(1842-1894) UK. 95 p.

G. Winter
Winter, Heinrich Georg
(1848-1887)
 GLOSSARY
(terms as applied to rust fungi)

o&

aecial primordia: haploid structures usually pro- kind of sorus is called a telium and cells are called
duced with spermogonia; they develop into aecia teliospores, the resulting basidia are called inter-
upon dikaryotization (also called aecial initials) nal basidia.
(Fig. 12).
basidiospore: haploid spore produced on the
aecidioid: sorus type applied to aecia, uredinia, or basidium, usually as a result of meiosis; often
telia resembling the anamorphic genus Aecidium; binucleate (two nuclei in a spore) due to mitotic
not a recommended term. division in the spore.
aeciospore: a dikaryotic spore produced after basidium (pl. basidia): the structure that produces
dikaryotization. basidiospores following meiotic division of a
diploid nucleus; in rust fungi, basidia are equiva-
aecium (pl. aecia): the first dikaryotic sorus devel-
lent to metabasidia of some heterobasidio-
oping after dikaryotization; usually accompanied
mycetes, and teliospores are equivalent to
by spermogonia.
probasidia.
amphispore: a specialized urediniospore that has
bilaminate: two-layered, applied to spore walls
thicker walls and is usually more deeply
(e.g., teliospores of Phragmopyxis and Uropyxis).
pigmented than the ordinary urediniospores
(e.g., Puccinia vexans Farl.), but some species have caeomatoid: sorus having morphology like the
colorless amphispores (e.g., many species of anamorphic genus Caeoma (sensu lato), i.e., with
Hyalopsora). catenulate spores but no peridium.
anamorph: an asexual spore state; in rust fungi, all catenulate: spores produced in chains.
morphs besides the teleomorph (teliospore).
conidioma: a specialized multihyphal, conidia-
annellidic: repeated production of blastic spores in bearing structure.
a basipetal fashion, leaving ringlike scars at the
end of the sporophore; in rusts, spermatia forma- cysts: sterile, usually hyaline, hygroscopic cells
tion is considered annellidic. that subtend teliospores, likely involved in spore
release and dispersal (e.g., teliospores of Cysto-
apical cells: usually denotes cells at the distal end
myces, Spumula, Ravenelia, and Kernkampella).
of the pedicel and to which spores are attached
(e.g., teliospores of Diabole, Diorchidiella, and demicyclic: a life cycle comprising spermogonia,
Anthomyces). aecia, and telia but lacking uredinia (e.g., most
species of Gymnosporangium).
autoecious: describes a rust fungus requiring only
a single host to complete the life cycle. dikaryon: a pair of sexually compatible nuclei in a
cell, usually from different parents.
ballistospore: a spore that is discharged forcibly;
basidiospores of most rust species are ballisto- diorchidioid: two-celled teliospores with the sep-
spores. tum more or less vertical (e.g., Diorchidium and
Desmella).
basal cell: a cell that produces more than one spore
(e.g., found in telia of Edythea, Desmella, and echinulate: with sharp pointed spines, characteris-
Hemileia). tic of most urediniospores (Fig. 14A-C).
basidiosorus: sometimes refers to teleomorph of a endocyclic: a microcyclic life cycle in which aecio-
rust that lacks clearly defined telia or teliospores spore-like cells produce basidia and basidio-
(probasidia), but having cells that divide into spores upon germination (e.g., Endophyllum,
basidia without germination; in this manual this Endocronartium, and Endoraecium).

203
external basidium: basidium produced outside of ontogenic system: naming of spore states based on
teliospore by germination; occurs in most genera spore function or position in the life cycle rather
(cf. internal and semi-internal basidium). than on the basis of morphology (cf. morphologic
system).
fasciculate: applies to compound structure
composed of several adherent pedicels (e.g., ostiolar cells: cells that delimit the opening
Ravenelia). (ostiole) of a dome-shaped peridiate sorus (e.g.,
Milesia-type uredinia of Pucciniastrum, Milesina,
flabelliform: in the form of a half-circle, like a fan; or Melampsoridium, or aecia of Naohidemyces).
occurs in telia of Kweilingia.
paraphysis (pl. paraphyses): sterile, elongated cells
flexuous hypha: hyphal projection from spermo- within or at the edge of a sorus (cf. periphysis)
gonium or stoma and likely serving as a receptive (e.g., in uredinia of Melampsora, Aplopsora, and
structure for spermatia of compatible mating Olivea).
type (Fig. 12).
patelliform: a round plate having a well-marked
germ pore: a small, usually discrete, area of spore edge (e.g., teliospores of Kernkampella).
wall through which germination occurs (Fig. 15).
pedicel: stalk cell of a spore.
heteroecious: describes a rust fungus requiring
pedicellate: having a pedicel.
two unrelated hosts to complete its life cycle.
peridermioid: sorus type applied to aecia,
holomorph: the whole fungus, including all states
uredinia, or telia resembling the anamorph genus
and morphs, and the name based on the
Peridermium.
teleomorph.
peridiate: having a peridium.
hyaline: transparent or translucent, often used in
the sense of colorless. peridium (pl. peridia): a cellular layer surrounding
or enclosing the sporogenous area of sorus; the
hygroscopic: becoming soft and large by absorbing peridium of Aecidium, Peridermium, and other
water; may occur in spore walls, pedicels genera is composed of modified spores and is
(Gymnosporangium, Phragmopyxis, Uropyxis), or ontogenetically different from the membranous
cysts (Ravenelia, Uromycladium). peridium of uredinia of Miyagia, Corbulopsora,
intercalary cells: sterile cells occurring between and Batistopsora (Uredostilbe-type) and of
catenulate spores. Uredinopsis, Milesina, Hyalopsora, and Puccinia-
strum (Milesia-type).
internal basidium: a basidium that develops by
peripheral: around the margin of a sorus.
septation of the contents of a teliospore (e.g.,
Ochropsora, Coleosporium, and Achrotelium); peripheral paraphysis: elongated sterile cells
strictly speaking, these rusts can be considered to (paraphyses) surrounding a sorus; periphysis.
lack teliospores (i.e., probasidia or spores give
rise to metabasidia upon germination) (cf. periphysis: a peripheral paraphysis.
basidiosorus). receptive hypha: same as flexuous hypha and
trichogyne.
macrocyclic: describes a rust fungus having aecia,
uredinia, telia, and usually spermogonia; may be reniform: kidney-shaped (e.g., urediniospores of
autoecious or heteroecious. Diorchidium and Hemileia).
microcyclic: describes a rust fungus having sper- reticulate: netlike spore surface ornamentation
mogonia and telia or only the latter. (Fig. 14N).

mitospore: a spore produced by mitosis. roestelioid: having the appearance of the ana-
morph genus Roestelia.
morphologic system: naming of spore states on the
basis of spore morphology (cf. ontogenic system). semi-internal basidium: abasidium that formsina
manner somewhat intermediate between exter-
muriform: spores divided by septa in more than nal and internal basidia; the apical part of the
one plane. teliospore continues to grow and produces a

204
 basidium; occurs in Mikronegeria, Blastospora, teliospore: probasidium of rust fungi; a spore that
Chrysocelis, and Chrysocyclus. germinates to produce a basidium (metaba-
sidium); nuclear fusion usually occurs within the
seriate: arranged in a series or succession. spore.
sessile: spore produced without a pedicel. telium (pl. telia): the sorus that bears teliospores.
sorus: sporulating structure. trichogyne: same as flexuous hypha and receptive
hypha.
spermatium (pl. spermatia): noninfecting haploid
spermatizing cell produced in a spermogonium. triquetrous: three-cornered, three-edged (e.g.,
teliospores of Hapalophragmium).
spermogonium (pl. spermogonia): a structure con-
taining spermatia; also called a pycnium. urediniospore: usually dikaryotic repeating spore,
sometimes called a uredospore or urediospore.
sporogenous cells: cells that produce spores; often
also called basal cells. uredinium (pl. uredinia): sorus bearing uredinio-
spores, sometimes called a uredium.
suprastomatal: developing through and outside of
stomata (e.g., sori of Hemileia, Gerwasia, Blasto- uredinoid: sorus type applied to aecia, uredinia, or
spora, Zaghouania, Desmella, and Edythea). telia that resemble the anamorphic genus Uredo
(sensu lato); not a recommended term.
systemic infection: infection spreading internally
throughout the plant tissue; opposite of localized verrucose: wartlike spore surface markings; char-
infection. acteristic of most aeciospores (Fig. 14D-G).

teleomorph: sexual (perfect) form or morph of a

fungus; in rust fungi, the teliospores.

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 LIST OF GENERA

ollowing is a list of all described genera of rust Calyptospora J.G. Kuhn

fungi. Accepted holomorph and anamorph Capitularia Rabenh. =Uromyces
genera are in boldface type; synonyms and rejected Catenulopsora Mundkur =Cerotelium
genera are in regular type. Centridium Chevall. =Roestelia (anamorph)
Cephalotelium Syd. =Ravenelia
Ceraceopsora Kakish., T. Sato & S. Sato
Acervulopsora Thirum. =Maravalia Ceratitium Rabenh. =Roestelia or Aecidium
Achrotelium Syd. (anamorph)
Aecidiella Ellis & Kelsey =Pucciniosira Ceratocoma Buritica & J.F. Hennen
Aecidium Pers. ex Pers. (anamorph) Ceropsora B.K. Bakshi & Suj. Singh
Aeciure Buritica & J.F. Hennen =Caeoma (anamorph) Cerotelium Arthur
Agromyces Arthur =Puccinia Cerradoa J.F. Hennen & Y. Ono =Edythea
Agrotelium Arthur =Puccinia Chaconia Juel
Allodus Arthur =Puccinia Chardoniella F. Kern
Allopuccinia H.S. Jacks. =Sorataea Chnoopsora Dietel =Melampsora
Allotelium Syd. Chrysella Syd.
Alveolaria Lagerh. Chrysocelis Lagerh. & Dietel
Alveomyces Bubak =Uromyces Chrysocyclus Syd.
Ameris Arthur =Phragmidium Chrysomyxa Unger
Angiopsora Mains =Phakopsora Chrysopsora Lagerh.
Angusia Laundon =Maravalia Ciglides Chevall. =Roestelia (anamorph)
Anthomyces Dietel Cionothrix Arthur
Anthomycetella H. & P. Syd. Cleptomyces Arthur
Aplopsora Mains Coinostelium Syd. =Prospodium
Apra J.P. Hennen & F.O. Freire Coleoma Clem. =Gymnosporangium
Aregma Fr. =Phragmidium Coleopuccinia Pat. =Gymnosporangium
Argomyces Dietel =Puccinia Coleopucciniella Hara =Gymnosporangium
Argomycetella H. & P. Syd. =Maravalia Coleosporium Lév.
Argotelium Arthur =Puccinia Corbulopsora Cummins
Arthuria HS. Jacks. Coronotelium Syd. =Puccinia
Arthuriomyces Cummins & Y. Hirats. Cronartium Fr.
=Gymnoconia Crossopsora H. & P. Syd.
Atelocauda Arthur & Cummins Ctenoderma Syd. =Skierka
Cumminsiella Arthur
Baeodromus Arthur Cumminsina Petr.
Barclayella Dietel =Chrysomyxa Cutomyces Thm. =Puccinia
Batistopsora Dianese, Medeiros & Santos Cystingophora Arthur =Ravenelia
Bitzea Mains =Chaconia Cystomyces Syd.
Blastospora Dietel Cystopsora E.J. Butler =Zaghouania
Botryorhiza Whetzel & Olive Cystotelium Syd. =Ravenelia
Bubakia Arthur =Phakopsora
Bullaria DC. =Puccinia Dasturella Mundk. & Khesw. =Kweilingia
Caeoma Link (anamorph) Dasyspora Berk. & M.A. Curtis
Caeomurus (Link) Gray = Uromyces Dendroecia Arthur =Ravenelia
Calidion H. & P. Syd. (anamorph) Desmella H. & P. Syd.
Calliospora Arthur =Uropyxis Desmellopsis J.M. Yen =Puccinia

206
Desmotelium Syd. =Chaconia Holwayella H.S. Jacks. =Chrysocyclus
Diabole Arthur Hyalopsora Magnus
Diabolidium R. Berndt =Allotelium Hypodermium Link =Caeoma (anamorph)
Dicaeoma Gray =Puccinia Intrapes J.F. Hennen & Figueiredo (excluded, may
Dicheirinia Arthur not be rust)
Dichlamys H. & P. Syd. =Uromyces
Didymopsora Dietel Jacksonia J.C. Lindq. =Dietelia
Didymopsorella Thirum. Jacksoniella Sathe. =Dietelia
Didymosira Clem. =Pucciniosira Jackya Bubak =Puccinia
Dietelia Henn. Joerstadia Gjerum & Cummins
Diorchidiella J.C. Lindg.
Diorchidium Kalchbr. Kamatomyces Sathe =Masseeélla
Diphragmium Boedijn =Diorchidium Kernella Thirum.
Dipyxis Cummins & J.W. Baxter Kernia Thirum. =Kernella
Discospora Arthur =Pileolaria Kernkampella Rajendren
Kimuromyces Dianese, Santos, Medeiros &
Earlea Arthur =Phragmidium Furlanetto
Edythea HS. Jacks. Klastopsora Dietel =Pucciniostele
Elateraecium Thirum., F. Kern & B.V. Patil Klebahnia Arthur =Uromyces
(anamorph) Kuehneola Magnus
Endocronartium Y. Hirats. Kulkarniella Gokhale & B.V. Patil =Monosporidium
Endophylloides Whetzel & Olive =Dietelia Kunkelia Arthur =Gymnoconia
Endophyllum Lévy. Kweilingia Teng
Endoraecium Hodges & D.E. Gardner
Epitea Fr. =Caeoma (anamorph) Lecythea Lév. (anamorph)
Eriosporangium Bert. =Puccinia Leptinia Juel =Calidion (anamorph)
Esalque J.F. Hennen, Figueiredo & A.A. Carvalho Leptopuccinia (G. Winter) Rostr. =Puccinia
Leucotelium Tranzschel =Sorataea (see Savile Can.
Frommea Arthur =Phragmidium J. Bot. 67:2983. 1989)
Frommeélla Cummins & Y. Hirats. Lindrothia Syd. =Puccinia
Linkiella Syd. =Puccinia
Gallowaya Arthur =Coleosporium Lipocystis Cummins
Gambleola Massee =Pucciniosira Lipospora Arthur =Tranzschelia
Gerwasia Racib. Longia Syd. =Ravenelia
Goplana Racib. Lysospora Arthur =Tranzschelia
Groveola Syd. =Uromyces
Gymnoconia Lagerh. Macabuna Buritica & J.F. Hennen =Calidion
Gymnopuccinia T.S. Ramakr. =Didymopsorella (anamorph)
Gymnosporangium R. Hedwig Macalpinia Arthur =Uromycladium
Gymnotelium Syd. =Gymnosporangium Macruropyxis Azbukina
Mainsia H.S. Jacks. =Gerwasia
Hamaspora Korn. Malupa Y. Ono, Buritica & J.F. Hennen
Hamasporella Hohn. =Hamaspora (anamorph)
Hapalophragmiopsis Thirum. =Hapalophragmium Maravalia Arthur
Hapalophragmium H. & P. Syd. Masseeélla Dietel
Haplopyxis H. & P. Syd. =Uromyces Mehtamyces Mundk. & Thirum.
Haploravenelia (Dietel) Syd. =Ravenelia Melampsora Castagne
Haplotelium Syd. =Uromyces Melampsorella J. Schrot.
Hemileia Berk. & Broome Melampsoridium Kleb.
Hemileiopsis Racib. =Hemileia Melampsoropsis (J. Schrot.) Arthur =Chrysomyxa
Hennenia Buritica Mesopsora Dietel =Melampsora
Hiratsukaia Hara =Chrysomyxa Micropuccinia Rostr. =Puccinia
Hiratsukamyces Thirum., F. Kern & B.V. Patil Microravenelia Dietel =Ravenelia

207
 Microtriphragmium G. Winter =Triphragmium Polythelis Arthur =Tranzschelia
Mikronegeria Dietel Pomatomyces Oerst. =Peridermium (anamorph)
Milesia F.B. White (anamorph) Porotenus Viégas
Milesina Magnus Prospodium Arthur
Mimema HS. Jacks. Pseudopuccinia Hohn. =Stigmina, not a rust
Miyagia Miyabe ex H. & P. Syd. Puccinella Fuckel =Uromyces
Monosporidium Barclay =Endophyllum Puccinia Pers. ex Pers.
Pucciniastrum G.H. Otth
Naohidemycees S. Sato, Katsuya & Y. Hirats. Puccinidia Mayr =? Puccinia (on Abies concolor in
Necium Arthur =Melampsora North America)
Neoravenelia Long =Ravenelia Pucciniola L. Marchand =Uromyces
Nephlyctis Arthur =Prospodium Pucciniosira Lagerh.
Newinia Thaung Pucciniostele Tranzschel & Kom.
Nielsenia Syd. =Uromyces
Nigredo (Pers.) Roussel =Uromyces Ramakrishnania Ramachar & Bhagyan.
Nothoravenelia Dietel Ravenelia Berk.
Nyssopsora Arthur Reyesiella Sacc. =Anthomycetella
Nyssopsorella Syd. =Triphragmiopsis Roestelia Rebent. (anamorph)
Rostrupia Lagerh. =Puccinia
Ochropsora Dietel Rubigo (Pers.) Roussel =Uredo (anamorph)
Olivea Arthur
Ontotelium Syd. =Uromyces Santapauella Mundk. & Thirum. =Phragmidiella
Oplophora Syd. =Nyssopsora Sartvellia Berk. =Dasyspora
Scalarispora Buritica & J.F. Hennen
Peridermium (Link) J.C. Schmidt & Kunze Schizospora Dietel =Pucciniosira
(anamorph) Schroeterella Syd. =Puccinia
Peridiopsora Kamat & Sathe =Milesia (anamorph) Schroeteriaster Magnus =Uromyces
Peridipes Buritica & J.F. Hennen =Milesia Sclerotelium Syd. =Puccinia
(anamorph) Scopella Mains =Maravalia
Peristemma Syd. =Miyagia Scopellopsis T.S. Ramakr. & K. Ramakr. =Maravalia
Persooniella Syd. =Puccinia Skierka Racib.
Petersonia Cummins & Y. Hirats. (anamorph) Solenodonta Castagne =Puccinia
Phakopsora Dietel Sorataea Syd.
Phragmidiella Henn. Sphaerophragmium Magnus
Phragmidiopsis G. Winter =Phragmidium Sphenospora Dietel
Phragmidium Link Spirechina Arthur =Kuehneola
Phragmopsora Magnus =Pucciniastrum Spumula Mains
Phragmopyxis Dietel Stakmania Kamat & Sathe =Phakopsora
Phragmostele Clem. =Pucciniostele Stereostratum Magnus
Phragmotelium Syd. =Caeoma (anamorph) Stichopsora Dietel =Coleosporium
Physonema Lév. =Caeoma (anamorph) Stilbechrysomyxa M. Chen =Chrysomyxa
Physopella Arthur =Phakopsora Stomatisora J.M. Yen = Chrysocelis
Pileolaria Castagne Synomyces Arthur =Coleosporium
Pleomeris Syd. =Puccinia
Pleoravenelia Long =Ravenelia Tegillum Mains =Olivea
Podisoma Link =Gymnosporangium Teleutospora Arthur & Bisby =Uromyces
Podocystis Fr. =Uredo (anamorph) Teloconia Syd. = Phragmidium
Podosporium Lév. =Melampsora Telomapea Laundon =Maravalia
Polioma Arthur Telospora Arthur =Uromyces
Poliomella Syd. =Puccinia Thekopsora Magnus
Poliomopsis Ramaley =? Polioma (excluded: the Thirumalachariella Sathe =Dietelia
type and only reported specimen missing) Trachyspora Fuckel
Poliotelium Syd. =Uromyces Trachysporella Syd. =Trachyspora

208
Trailia Syd. =Puccinia Uredopeltis Henn.
Tranzschelia Arthur Uredostilbe Buritica & J.F. Hennen (anamorph)
Triactella Syd. =Hapalophragmium Uromyces (Link) Unger
Tricella Long =Phragmopyxis Uromycladium McAlpine
Trichopsora Lagerh. Uromycodes Clem. =Uromyces
Triphragmiopsis Naumov Uromycopsis Arthur =Uromyces
Triphragmium Link Uropyxis J. Schrot.
Trochodium Syd. =Uromyces
Wardia J.F. Hennen (anamorph)
Trolliomyces Ulbr. =Phragmidium
Tunicopsora Suj. Singh & P.C. Pandey =Kweilingia
Xanthotriphragmium Milesi & Traverso
; =Triphragmium
Uleiella J. Schrot. (not a rust according to Xenodochussenlidl
Thirumalachar and Mundkur (1950) Xenostele H. & P. Syd. =Puccinia
Uncol Buritica (excluded)
Uraecium Arthur =Milesia (anamorph) Ypsilospora Cummins
Uredendo Buritica & J.F. Hennen =Uredo
(anamorph) Zaghounia Pat.
Uredinopsis Magnus
Uredo Pers. (anamorph)

209
 INDEX
roe

ilies following index includes scientific names of Cumminsiella, 163

fungi and plants, common names of plants, and Dasturella, 80
common names of plant diseases caused by rust Endophyllum, 47
fungi. Entries for common names of plant diseases Gymnosporangium, 164
include the scientific names of the rust fungi that Hennenia, 135
cause them. Under each anamorph type (e.g., Kernkampella, 136
Aecidium-type), the names of the rust genera having Masseeélla, 189

that type are listed. Page numbers in bold type refer Miyagia, 169
Monosporidium, 43
to the main description of a family or genus.
Ochropsora, 100
Phakopsoraceae, 32, 43
Phragmidiella, 86
Puccinia, 171
A Pucciniaceae, 34, 47
Abies, 2,7, 8, 11, 31, 51,52, 53, 56, 58, 60 Pucciniostele, 87
alba, 39 Ravenelia, 139
lasiocarpa, 8 Raveneliaceae, 33
abnormal life cycles, 25 Sorataea, 115
Acacia, 118, 121, 122
Sphaerophragmium, 142
koa, 5, 7,118, 122 Spumula, 144
lebbeck, 142 Tranzschelia, 116
pycnantha, 121 Triphragmiopsis, 145
saligna, 7 Uromyces, 175
acacia rusts Uropyxidaceae, 33
Atelocauda angustiphylloda, 3, 118 Zaghouania, 177
Atelocauda bicincta, 118, 119
aecium, definition and morphology, 16, 20
Atelocauda digitata, 5 Aeciure, 37, 76
Atelocauda hyalospora, 118 Aegiphila, 161
Atelocauda koae, 3,7, 26, 118, 119 Agaricostilbum, 1,2
Endoraecium acaciae, 3, 25, 118, 122 Albizia, 121
Endoraecium hawatiense, 118, 122 lebbeck, 142
Ravenelia pringlei, 25 Alchemilla, 152, 156
Uromycladium tepperianum, 7 pedata, 152
Acanthaceae, 72 vulgaris, 156
Aceraceae, 58
xanthochlora, 156
Achrotelium, 33, 44, 90
Alchornea latifolia, 101
ichnocarpi, 90 alfalfa rust
Adenocalymma calderonti, 105 Uromyces striatus, 25, 175, 176
Aecidium, 37, 38, 40, 92, 178, 184, 185, 186
Allium, 11, 74
berberidis, 37
Allotelium, 34, 46, 123, 130
holboellii, 26
calliandrae, 123
mort, 3,5
mirabile, 123
nitens, 150 Alnus, 54
pampeanum, 26 Alternaria, 127
solani-argentei, 184
Alveolaria, 34, 48, 178, 179
Aecidium-type, 22, 36 andina, 179
Apra, 126 cordiae, 179
Ceraceopsora, 93
dugetiae, 179
Cerotelium, 78
Amelanchier alnifolia, 9
Chaconiaceae, 33
Amicia lobbiana, 115
Corbulopsora, 162
Amorpha herbacea, 117

210
 Ampelopsis, 87 Asarum caulascens, 78
cantoniensis, 88 Asclepiadaceae, 66, 90
Amphilophium, 112 Asian hard pine gall rust
Anacardiaceae, 33, 86, 107, 120, 153 Cronartium orientale, 3, 7,27, 66
anamorph genera, key and descriptions, 36-40 as paragus rust
Anaphalis margaritacea var. angustior, 169 Puccinia asparagi, 171
Anemia Aster ciliolatus, 9
fulva, 103 Asteraceae, 64,95, 158, 160, 162, 169,171, 175, 180, 182, 183 ,
tomentosa var. fulva, 103 184, 186
Anemone, 100, 116 Astilbe, 3, 87
berlandieri, 116 chinensis, 87
caroliniana, 116 microphylla, 5
flaccida, 93 Astilbe rust
Angiopsora, 84 Pucciniostele clarkiana, 3,5, 87
compressa, 85 Astronium, 105
divina, 80 fraxinifolium, 107
angiosperms, 1 Atelocauda, 33, 45, 118-119, 120, 121, 122
Angusia, 98 angustiphylloda, 3, 118
Annona, 77, 135 bicincta, 118, 119
nolosericea, 40 digitata, 5,118, 119
tomentosa, 77 hyalospora, 118
Annonaceae, 40, 77, 102, 135, 142, 143, 179, 181 incrustans, 118, 119
Anthomyces, 34, 45, 124 koae, 3, 7, 26, 118, 119
brasiliensis, 124 Attalea ceraensis, 106
Anthomycetella, 34, 45, 124, 125, 127 Auriculariaceae, 95
canaril, 125 Auriculariales, 2, 32, 81
Apiaceae, 34, 138 Al istrocedrus, 73
Aplopsora, 33, 44, 78, 91, 93 Avena sativa, 4,21
corni, 91
hennenii, 91
lonicerae, 91 B
nyssae, 91 Baeodromeae, 180
tanakae, 78, 91 Baeodromus, 34, 48, 180, 181
Apocynaceae, 90 dominicana, 180
apple rust holwayi, 180
Gymnosporangium juniperi-virginianae, 7,25, 164, 165 senecionis, 180
Apra, 34, 45, 126 tranzschelii, 180
bispora, 126 bamboo stem rust
apricot rust, Japanese Stereostratum corticioides, 2, 4, 174
Blastospora smilacis, 3,5, 7,10, 70, 71 Bambusa, 81
Caeoma makinoi, 3, 10, 70 Bambusoideae, 81, 161, 174
Aquifoliaceae, 62 Ba phia nitida, 147
Arabis, 7,26 Basidiomycetes, 1, 2
Arachis hypogaea, 4, 21 Basidiomycota, 2, 3
Araliaceae, 138 basidium, definition and morphology, 22
Araucaria, 73 Ba tistopsora, 32, 40, 43, 77
araucana, 39 crucis-filti, 77
Araucariaceae, 39 pistila, 77
Arctostaphylos uva-ursi, 8, 62 be an rust
Aregma, 154 Uromyces appendiculatus, 2,25, 175,176
Arrabidaea, 105 beet rust
Arthuria, 32, 37, 43, 76 Uromyces betae, 175
catenulata, 76 Be nsingtonia, 1
columbiana, 76 Berberidaceae, 37, 106, 145, 163, 186
glochidionis, 76 Be rberis, 11, 106, 163, 186
Arthuriomyces, 150 vulgaris, 4, 37
peckianus, 150 Be tula, 54, 70
Arundinaria, 174 verrucosa, 54
Betulaceae, 54, 58 Raveneliaceae, 33
Bignoniaceae, 11, 47, 82, 86, 89, 94, 105, 110, 112, 113 Uropyxidaceae, 33
birch-Thujopsis rust Xenodochus, 157
Blastospora betulae, 3, 5,7, 70,71 Zaghouania, 177
Caeoma deformans, 3,5, 7,70 Caesalpinia, 124, 133
black stem rust of wheat, grains and grasses Calidion, 36, 37, 38
Puccinia graminis, 2,4, 14, 20, 21, 25, 37,171 lindsaeae, 37
blackberry, 7 Calidion-type
Blastospora, 22, 32, 39, 42, 70-71, 73 Anthomycetella, 125
betulae, 3,5, 7,70, 71 Cerotelium, 78
itoana, 70, 71 Chaconia, 94
smilacis, 3,5, 7,10, 70, 71 Cumminsina, 127
Blennoria abietis, 62 Dasturella, 80
Boraginaceae, 53, 179, 187 Dicheirinia, 130
Botryorhiza, 33, 44, 90, 92, 98 Hapalophragmium, 134
hippocrateae, 92 Kimuromyces, 107
Bouteloua curtipendula, 21 Kweilingia, 81
Breynia rhamnoides, 136 Mimema, 109
brooms. See witches’ brooms Nothoravenelia, 83
brown rust of grains and grasses Phakopsora, 84
Puccinia recondita, 2, 4,19, 26, 171, 172 Phakopsoraceae, 43
Bubakia, 84 Phragmidiaceae, 34
Bulnesia, 139 Phragmidium, 154
Burseraceae, 83, 125, 190 Pucciniostele, 87
Raveneliaceae, 33
Scalarispora, 88
Cc Sphaerophragmium, 142
Caeoma, 37, 38, 39 Uredopeltis, 89
callianthum, 188 Uropyxidaceae, 33
deformans, 3,5, 7,70 Uropyxis, 117
makinoi, 3, 7, 10, 70 Calliandra
peltatum, 19, 32, 40 bijuga, 144
radiatum, 70 trinervia, 123
sanctae-crucis, 39 Calocedrus, 11
saxifragarum, 37 Calyptospora, 31, 39, 42, 51, 58
sorbi, 100 goeppertiana, 8, 25, 51
Caeoma-type Campanula, 62
Arthuria, 76 Campanulaceae, 62
Chrysomyxa, 62 Canada thistle, 7
Coleosporiaceae, 31 Canarium
Coleosporium, 64 commune, 190
Crossopsora, 79 villosum, 125
Gymnoconia, 150 Canavalia ensiformis, 78
Joerstadia, 152 cane blight, raspberry
Kernkampella, 136 Kuehneola uredinis, 15, 153
Masseeélla, 189 carnation rust
Melampsora, 74 Uromyces dianthi, 175
Melampsoraceae, 30, 32 Caryophyllaceae, 53
Nothoravenelia, 83 Cassia, 109, 133
Olivea, 101 versicolor, 109
Phakopsoraceae, 32 Catenulospora, 153
Phragmidiaceae, 34 cedar—apple rust
Phragmidium, 154 Gymmnosporangium juniperi-virginianae, 7,25, 164, 165
Phragmopyxis, 111 Celastraceae, 153, 188
Polioma, 170 Ceraceopsora, 33, 43, 93
Pucciniaceae, 34 elaeagni, 93
Pucciniostele, 87 Cerastium, 53
Ravenelia, 139 Ceratocoma, 34, 48, 181

212
 guineensis, 181 perlaria, 62
jacksoniae, 181 piceae, 61, 62
cereal rusts pirolata, 8, 62
black stem rust of wheat, etc. gilianensis, 62
Puccinia graminis, 2, 4, 14, 20, 21, 25, 37,171 rhododendri, 63
brown or leaf rust of wheat, etc. simplex, 62
Puccinia recondita, 2, 4, 19, 26, 171, 172 stilbae, 62
crown rust of oats, etc. succinea, 62, 63
Puccinia coronata, 21, 26, 171, 172 tsugae-yunnanensis, 62
Puccinia coronata var. avenae, 2, 4 weirtl, 62
Puccinia coronata var. coronata, 27 woroninil, 7, 8, 62
stripe or yellow rust of wheat, etc. Chrysomyxaceae, 30
Puccinia striiformis, 2, 171 Chrysopsora, 22, 34, 47, 159, 160
Ceropsora, 31, 42, 61, 62, 73 cestri, 160
piceae, 61, 62 gynoxidis, 160
smithiana, 61 mikaniae, 160
Cerotelium, 32, 43, 78, 86, 88, 91, 153 Cionothrix, 34, 48, 66, 178, 181, 183, 190
asar1, 78
praelonga, 183
canavaliae, 78
Cirsium arvense, 7
dicentrae, 78
classification of rust fungi, 30-35
fici, 78 Clematis, 9
hashiokai, 88
Cleptomyces, 34, 47, 107, 161, 174
tanakae, 78,91
lagerheimianus, 161
Cerradoa, 103 clover rust
palmaea, 106 Uromyces trifolii-repentis, 175
Cestrum megalophyllum, 159 Codiaeum, 185
Chaconia, 22, 33, 44, 72, 75, 93, 94,98, 147
Coffea, 4, 96
alutacea, 94 coffee leaf rust
ingae, 94 Hemileia vastatrix, 2, 4,96, 97
Chaconiaceae, 1, 22, 30, 32, 33, 34, 43-44, 61, 90-101
Coleopuccinia, 164
Chamaecyparis, 11 Coleopucciniella, 62, 164
Chardoniella, 34, 48, 182, 187 simplex, 62
andina, 182 Coleosporiaceae, 22, 30, 31, 42, 61-67, 160
gynoxidis, 182 Coleosporium, 7,11, 14, 22, 31, 39, 42, 64-65, 95
chickpea rust asterum, 9,19, 25, 65
Uromyces ciceris-arietinus, 2, 175 bletiae, 65
Chlamydomyces palmarum, 156 campanulae, 64
Chondrilla juncea, 7 clematidis, 9
Chrysella, 22, 34, 47, 158, 159 crowellii, 64, 187
mikaniae, 15, 158 himalayense, 64
Chrysocelis, 22, 32, 37, 43, 72, 73, 93, 98
incompletum, 64
geophilicola, 72 inconspicuum, 64
gymnostemmatis, 72 jonesii, 18
lupini, 15, 72 neocacaliae, 65
Chrysocyclus, 22, 34, 47, 158, 159, 160 phellodendri, 65
cestri, 159 pini-asteris, 65
Chrysomyxa, 7,11, 14, 22, 31, 39, 42, 61, 62-63, 76
pini-densiflorae, 64
abietis, 62 pint-pumilae, 64
albida, 153 pinicola, 25, 64
arctostaphyli, 7,8, 25, 62 pulsatillae, 65
bambusae, 81 reichei, 64
cassandrae, 63 rhinanthacearum, 64
deformans, 62 viguierae, 64
himalensis, 62 collection of rust fungi, 11
keteleeriae, 62 cone rusts
ledi, 62, 63 southern pine cone rust
ledicola, 8, 62, 63 Cronartium strobilinum, 7, 66
menziesiae, 62 southwestern pine cone rust

213
 Cronartium conigenum, 7, 66 Cryptostegia grandiflora, 7
spruce cone rust Cucurbitaceae, 72
Chrysomyxa pirolata, 8, 62 Cummiunsiella, 11,34, 47, 117, 163
Copaifera, 143 mirabilissima, 163
Corbulopsora, 34, 40, 47, 162, 169 sanguinea, 163
clemensiae, 162 standleyana, 163
cumminsit, 162 wootoniana, 163
gravida, 162 Cumminsina, 34, 46, 127
Cordia, 179 clavispora, 15, 127
corn rusts Cupressus, 11
Puccinia polysora, 2, 172 Cyathopsora, 113
Puccinia sorghi, 2,171, 172 Cydonia, 11
Cornaceae, 91 Cyperaceae, 171
Cornus controversa, 91 Cyperus esculentus, 7
correlated species and Tranzschel’s Law, 26 Cystomyces, 34, 45, 128
Corticium, 174 costaricensis, 128
Corydalis, 100 Cystopsora oleae, 177
lineariloba, 78 cytology and nuclear cycles, 26-28
Costaceae, 81
Costus pictus, 81
cotton rusts D
Phakopsora gossypii, 84 Dalbergiella, 143
Puccinia cacabata, 2,171, 172 Dasturella, 32, 43, 80, 81
Cracca virginiana, 139 divina, 80
Crassulaceae, 178 Dasyspora, 33, 45, 102
Crataegus, 11 foveolata, 102
Cronartiaceae, 22, 30, 31, 42, 66-69 gregaria, 15, 102
Cronartium, 11, 22, 26, 27, 32, 39, 42, 66-67, 68, 79, 182, Dendrocalamus, 80
183, 187 strictus, 80, 81
asclepiadeum, 66 Derris, 146
capparidis, 189 uliginosa, 134
coleosporioides, 3, 6, 66, 67 Desmella, 33, 40, 44, 103, 106
comandrae, 3, 6, 67 anemiae, 103
comptoniae, 3 Desmotelium, 72
conigenum, 7, 66 Diabole, 34, 45, 123, 126, 129
flaccidum, 3, 66 cubensis, 129
jacksoniae, 181 Diabolidium calliandrae, 123
keteleeriae, 62 Dicentra cucullaria, 78
orientale, 3, 7,27, 66 Dicheirinia, 34, 46, 123, 124, 126, 128, 129, 130, 131, 144
paraguayensis, 184 binata, 130
praelongum, 183 canariensis, 130
quercuum, 3, 7,27, 66 maderensis, 130
quercuum f.sp. fusiforme, 3, 16, 66 manaosensis, 130
ribicola, 3, 6, 14, 18, 19, 25, 66, 67 trispora, 130
strobilinum, 7, 66 viennotil, 130
verruciforme, 185 Didymopsora, 34, 48, 66, 178, 184
zizyphi, 79 africana, 15, 184
crop diseases, 2-3 chuquiraguae, 184
Crossopsora, 32, 37, 43, 66, 79, 109 macrospora, 184
malloti, 79 paraguayensis, 184
sawadae, 79 solani-argentei, 184
zizyphi, 79 toddaliae, 104, 184
Croton, 76, 89 triumfettae, 184
crown rust Didymopsorella, 33, 44, 104, 184
Puccinia coronata, 21, 26, 171, 172 lemanensis, 104
Puccinia coronata var. avenae, 2, 4 toddaliae, 104
Puccinia coronata var. coronata, 27 Dietelia, 25, 34, 48, 178, 179, 181, 185
Cruciferae, 26 codiaei, 185

214
 portoricensis, 15 pampeanum, 26
verruciformis, 15, 185 sempervivi, 25, 178
Diorchidiella, 34, 46, 126, 131 tuberculatum, 26
australis, 15, 131 Endoraecium, 22, 33, 45, 118, 122
verlandii, 131 acaciae, 3, 25, 118, 122
Diorchidium, 34, 46, 123, 130, 132, 143 hawatiense, 118, 122
amapaensis, 132 Eocronartium, 1,2
australe, 131 Epilobium angustifolium, 58
copaifera, 132 Enicaceaemllmolo77105) 59 02
koordersti, 132 Erythrina, 130
pallidum, 143 Erythrobasidium, 1
puiggarii, 132 Esalque, 46, 133, 135, 146
quadrifidum, 132 holwayi, 107, 133, 146
tetrasporum, 132 Eucalyptus, 5
woodii, 132 eucalyptus rust
Dioscorea, 143 Puccinia psidii, 3, 5
Dioscoreaceae, 34, 95, 143 Eupatorium, 180, 183
Diphragmium, 132 Euphorbia, 7
Dipyxis, 33, 44, 105, 108 exigua, 74
mexicana, 105 helioscopia, 40
viegasil, 105 Euphorbiaceae, 34, 40, 74, 76, 83, 89, 94, 95, 101, 136, 139,
diseases 175, 185, 189
agricultural and horticultural crops, 2-5 Euphragmidium, 154
forest trees, 3, 6-7 Euprospodium, 113
Dryopteris austriaca, 56
Duchesnea, 148 E
Fabaceae, 11, 33, 34, 39, 43, 44, 72, 78, 94, 98, 109, 111, 115,
E IZ, TANS), WAN), AL, 2, Wy A IS, IPAs) WS), iNSi0),
Earlea, 154 131, 132, 133, 134, 135, 137, 139, 142, 143, 144, 146,
eastern gall rust of hard pines 147, 161, 170, 175, 181
Cronartium quercuum, 3, 7,27, 66 Fagaceae, 32, 43, 58, 73
economic importance of rust fungi, 2-3, 7 families, key and descriptions, 30-35
Edythea, 33, 40, 44, 103, 106 fasciation, 7. See also witches’ brooms
palmaea, 106 fern rusts, 1, 2, 11, 52, 56, 60
quitensis, 106 Filipendula ulmaria, 146
tenella, 106 fone iebksusy toy, IIL, Day, Sy 5X6), ‘stsy, G10)
Eichhornia, 7 fir broom rust
Elaeagnaceae, 93 Melampsorella caryophyllacearum, 7,25, 39, 53
Elaeagnus fir needle rusts
macrophylla, 93 Calyptospora goeppertiana, 8, 25,51
umbellata, 93 Hyalopsora aspidiotus, 52
Elateraecium, 22, 36, 37, 38, 188 Pucciniastrum epilobii, 58
salicicola, 39 fir-fern rusts, 73
Elateraecium-type flax rust
Hiratsukamyces, 188 Melampsora lint, 2,3, 14, 22, 24, 25, 74,75
Empetraceae, 62 Flueggea virosa, 189
Endocronartium, 11, 22, 32, 42, 66, 68-69 forest tree diseases, 3, 6, 7
harknessii, 3, 6, 7, 25, 68 Fraxinus
pini, 3, 25, 68 lanaginosa, 108
sahoanum, 68 sieboldiana, 108
sahoanum var. hokkaidoense, 68 Frommea, 148
sahoanum var. sahoanum, 68 Frommeélla, 34, 47, 110, 148, 151, 157
yamabense, 68 mexicana, 148
Endophyllaceae, 25, 35, 178 tormentillae, 148
Endophylloides, 25, 66 Fumariaceae, 100
Endophyllum, 22, 25, 34, 35, 47, 178 fusiform gall rust
holwayi, 185 Cronartium quercuum f.sp. fusiforme, 3, 16, 66

ZS
 G ornatum, 134
Galium aparine, 84 haustoria, 24
gall symptom, 7. See also pine gall rusts Helicobasidium, 1, 2
Gambleola cornuta, 186 Helicogloea, 1
Geraniaceae, 170 Hemileia, 33, 40, 44, 96-97
Geraniales, 190 vastatrix, 2,4, 96, 97
Geranium, 170 Hemileopsis, 96
germ pores, 24 Hennenta, 133, 135
Gerwasia, 11, 34, 46, 149 ditelia, 135
rubi, 149 Herpobasidium, 1
standleyi, 149 Heterophragma sulphureum, 110
Glochidion, 76 Hippocratea volubilis, 92
Glycine max, 4 Hippocrateaceae, 39, 44, 92
Gompholobium latifolium, 181 Hiratsukaia tsugae, 62
Goodyera, 58 Hiratsukamyces, 35, 39, 48, 188
Goplana, 22, 33, 44, 73, 95 divinum, 188
dioscoreae, 15,95 salacicola, 188
micheliae, 95 hollyhock rust
grain and grass rusts. See cereal rusts Puccinia malvacearum, 25, 27,171
Grewia nenensis, 127 holomorph genera, keys by family, 42-48
Grossulariaceae, 95 Holwayella, 160
Guettarda, 89 host-rust relationship, 7, 11
Gymnoconia, 34, 37, 46, 47, 150 Hyalopsora, 2, 11, 31, 39, 42, 52, 73
alchemillae, 152 aspidiotus, 52
interstitiale, 150 chelianthis, 52
peckiana, 24, 25, 26, 150 polypodii, 52
Gymmnopuccinia, 104 Hymenomycetes, 2, 3, 24
Gymnosporangium, 7, 11,22, 34, 40, 47, 62, 164-167 Hyperium, 74
asiaticum, 9, 164 hypertrophy and hyperplasia (galls), 7. See also pine
biseptatum, 167 gall rusts
clavariiforme, 9, 165 hyphae, 24
clavipes, 9, 165
cunninghamianum, 167
ellisti, 165 I
exiguum, 167 Ichnocarpus volubilis, 90
externum, 166 identification of rust fungi, 11-12
fraternum, 167 Inga edulis, 147
fuscum, 164, 167, 171 Intrapes, 40, 136
gaeumannii spp. albertensis, 9, 164 Ixora, 173
hyalinum, 167
Juniperi-virginianae, 7,25, 164, 165
libocedri, 165 J
multiporum, 165 Jacksoniella, 185
nelsoni, 9 Japanese apricot rust
speciosum, 166 Blastospora smilacis, 3, 5,7, 10,70, 71
yamadae, 164, 167 Caeoma makinoi, 3, 10, 70
Gynoxys, 182 Jeffersonia dubia, 145
pulchella, 160 Joerstadia, 34, 46, 152
alchemillae, 152
keniensis, 152
H Jola, 1
Hamaspora, 11, 34, 46, 109, 151 June grass, 26
hashiokae, 151 Juniperus, 11, 164
longissima, 151 communis var. depressa, 9
Hapalophragmiopsis, 134 horizontalis, 9
Hapalophragmium, 34, 46, 133, 134, 135, 146 sabina, 164
derridis, 134
Juniperus—Rosaceae rusts, 7
holwayi, 133 Gymnosporangium, 7, 11, 164

216
K Lippia, 112
Kamatomyces, 189 Litsea, 168
Kernella, 34, 47, 168 Lonchocarpus, 118, 128
lauricola, 168 guatemalensis, 128
Kernia lauricola, 168 lucerne rust
Kernkampella, 34, 45, 83, 136, 139 Uromyces striatus, 25, 175, 176
appendiculata, 136 Lupinus, 72
breyniae, 136 Lycopodiophyta, 1
breyniae-patentis, 136
Keteleeria, 62 M
keys Macabuna, 37
anamorph genera, 36-37 Macruropyxis, 33, 44, 108
families, 30-31 fraxini, 108
holomorph genera by family, 42-48 Magnolia, 54
Kigelia africana, 110 Magnoliaceae, 95
Kimuromyces, 33, 44, 107 Mahonia, 11, 163
cerradensis, 107 Mainsia rubi, 149
koa rusts malformation symptom, 7
Atelocauda angustiphylloda, 3,118 Malupa, 36, 38, 39
Atelocauda koae, 3,7, 26, 118, 119 meibomiae, 39
Endoraecium acaciae, 3, 25, 118, 122 Malupa-type
Endoraecium hawatiense, 118, 122 Anthomyces, 124
Koeleria cristata, 26 Aplopsora, 91
Kriegeria, 1 Cerotelium, 78
Kuehneola, 22, 34, 46, 110, 153 Crossopsora, 79
albida, 153 Ochropsora, 100
loesneriana, 153 Olivea, 101
uredinis, 15, 153 Phakopsora, 84
Kulkarniella, 178 Phakopsoraceae, 32
Kweilingia, 32, 43, 81 Prospodium, 113, 114
americana, 81 Raveneliaceae, 33
bagchii, 81 Uropyxidaceae, 33
bambusae, 81 Malus, 11
divina, 81 Malvaceae, 25, 153, 185
Malvaceae-Stipa rust
L Puccinia interveniens, 25, 26
Lactuca, 162 Maravalia, 33, 44, 90, 92, 98-99
Lamiaceae, 101, 170 ascotele, 98
Laportea canadensis, 78 cryptostegiae, 7
Larix, 11,19, 54, 145 elata, 98
Lauraceae, 95, 168 kevorkianti, 99
leaf rusts of grains and grasses pallida, 98, 99
Puccinia recondita, 2, 4, 19, 26, 171, 172 pura, 98
Lecythea, 37, 38, 39 Markhamia, 89
Ledum, 62 sansibarensis, 86
groenlandicum, 8 Masseeélla, 35, 48, 189
legumes, 11, 175. See also Fabaceae capparidis, 189
Leptinia brasiliensis, 107 flueggiae, 189
Leucotelium, 115 narasimhanii, 189
padi, 115 Mehtamyces, 82, 86
prunti-persicae, 115 stereosperm1, 82, 86
life cycle variations, 24-25 Melampsora, 3, 11, 22, 32, 37, 40, 43, 74-75, 91
life cycles, abnormal, 26 albertensis, 19, 21
Liliaceae, 171, 175 chelidonii-pierotii, 75
Linaceae, 74 euphorbiae, 40, 74
Lindsaea, 37 fagi, 73
Lipocystis, 34, 45, 137 farlowti, 74
caesalpiniae, 15, 137 hypericorum, 74

PLATE
 hpi, 22,3), WD, Wh sy VAN 785) holwayi, 109
monticola, 75 venturae, 109
occidentalis, 5,75 Mimosa, 126, 129, 131, 137
punctiformis, 84 ceratonia, 137
vaccinil, 57 micracantha, 126
vernalis, 37 rocae, 131
Melampsoraceae, 30, 32, 43, 74-75 rufescens var. amnis-nigri, 126
Melampsorella, 31, 39, 42, 53 mint rust
aspidiotus, 52 Puccinia menthae, 2,171
caryophyllacearum, 7,25, 39,53 Mixia, 1,2
kriegeriana, 56 osmundae, 2
symphyti, 15, 53 Miyagia, 34, 40, 47, 162, 169
Melampsoridium, 31, 39, 42, 54-55 anaphalidis, 169
alni, 54 macrospora, 169
asiaticum, 54 pseudosphaeria, 169
betulinum, 54,55 Monosporidium, 22, 32, 43, 178
carpini, 54 Morus alba, 5
hiratsukanum, 54 mulberry rust
inerme, 54 Aecidium mori, 3,5
Melastromataceae, 184
Meliaceae, 138
Meliosmaceae, 95 N
Meliosoma myriantha, 19 Naohidemyces, 16, 31, 39, 40, 42, 57, 58, 59
Memora, 21, 112 fujisanensis, 57
glaberrima, 112 vaccinit, 57
Mesopsora hypericorum, 74 Nephlyctis, 113
Meum athamanticum, 138 Newinia, 33, 34, 45, 47, 110
Michelia velutina, 95 heterophragmae, 110
Microbotryum, 2 kigeliae, 110
Mikania hirsutissima, 158 thaiana, 21, 110
Mikronegeria, 22, 32, 39, 43, 73 Nothofagus
alba, 19, 22, 73 nervosa, 73
fagi, 39, 73 procera, 73
Mikronegeriaceae, 22, 30, 32, 42-43, 70-73, 159 Nothoravenelia, 32, 43, 83
Milesia, 22, 36, 38, 39, 40, 56, 58 commiphorae, 83
polypodii, 39 Japonica, 83
Milesia-type nuclear cycles and cytology, 26-28
Cerotelium, 78 nut sedge, 7
Cronartiaceae, 31 Nyssa aquatica, 91
Cronartium, 66 Nyssaceae, 91
Hiratsukamyces, 188 Nyssopsora, 34, 46, 138, 145, 146
Hyalopsora, 52 cedrelae, 138
Melampsorella, 53 clavellosa, 138
Melampsoridium, 54 echinata, 138
Milesina, 56
Naohidemyces, 42, 57 O
Phakopsora, 84
Ochropsora, 22, 33, 44, 91, 93, 98, 100
Pucciniastraceae, 31, 42
ariae, 100
Pucciniastrum, 58 daisensis, 100
Thekopsora, 59 kraunhiae, 100
Uredinopsis, 60 nambuana, 100
Milesina, 2, 11, 16, 31, 39, 42, 52, 56 sorbi, 100
dieteliana, 39, 56
Oleaceae, 94, 108, 177
kriegeriana, 56 Olearia, 162
pycnograndis, 56
Olivea, 33, 37, 39, 44, 73, 98, 101
vogesiae, 56 capituliformis, 101
Milletia, 132
fimbriata, 101
Mimema, 109, 115
Onagraceae, 58

218
 onion rust compressa, 85
Puceinia allii, 3 gossypii, 84
ontogenic system, 14 mexicana, 85
orange rust of Rubus pachyrhizi, 3, 4, 84, 85
Gymnoconia peckiana, 25 punctiformis, 84
Orchidaceae, 58 stereospermi, 82
orchids, 143 Phakopsoraceae, 22, 30, 32, 43, 76-89, 77
Phakopsoreae, 32
Phaseolus vulgaris, 175
P Phegopteris
Pachnocybe, 2 dryopteris, 52
Palmae, 106 polypodioides, 60
pea rust Phillyrea media, 177
Uromyces pisi, 7, 175 Phleogena, 2
peach rust Phragmidiaceae, 22, 31, 34, 46-47, 148-157
Tranzschelia discolor, 3, 116 Phragmidiella, 32, 43, 82, 86
peanut rust markhamiae, 15, 86
Puccinia arachidis, 3, 4,21, 172 Phragmidium, 11, 34, 37, 39, 47, 88, 127, 148, 151, 154-155,
pear rusts 156
Gymmosporangium asiaticum, 164 barnardi, 18
Gymnosporangium juniperi-virginianae, 7, 25, 164, 165 biloculare, 155
Gymnosporangium yamadae, 164, 167 boreale, 155
Peridermium, 37, 38, 39 fusiforme, 155
elatinum, 39 longissimum, 151
filamentosum, 68 mucronatum, 25, 39, 154
harknessii, 25, 68 peckianum, 155
pini, 25 potentillae, 148
Peridermium-type, 22, 30 rosae-multiflorae, 3, 5, 21
Calyptospora, 51 sanguisorbae, 18
Chrysomyxa, 62 tormentillae, 148
Coleosporiaceae, 31 violaceum, 7
Coleosporium, 64 Phragmopyxis, 33, 45,111
Cronartiaceae, 31 acuminata, 111
Cronartium, 66 deglubens, 111
Crossopsora, 79 leonensis, 111
Hyalopsora, 52 noelit, 111
Melampsorella, 53 Phragmotelium, 154
Melampsoridium, 54 Phyllocladus trichomanides, 19
Milesina, 56 phylogeny of rust fungi, 2
Naohidemyces, 57 Physopella, 40, 80, 82, 84, 89
Pucciniastraceae, 31, 42 meibomiae, 39
Pucciniastrum, 58 mexicana, 85
Thekopsora, 59 pachyrhizae, 84
Uredinopsis, 60 [eno sy. Maly, 3311), Sy, sts}, SiS),
Peridiopsora, 39 abies, 62
Peridipes, 39 glauca, 8
Periplocaceae, 90 smithiana, 61
Petersonia, 37, 38, 39 Pileolaria, 33, 45, 118, 120
sanctae-crucis, 39 brevipes, 15, 19, 120
Petersonia-type effusa, 120
Blastospora, 70 klugkistiana, 120
Chrysocelis, 72 shiraiana, 120
Mikronegeria, 73 terebinthi, 25, 120
Mikronegeriaceae, 32 Pileolariaceae, 22, 31, 33, 45, 118-122
Phragmidiaceae, 34 Pinaceae, 39, 61, 62, 68, 145, 164
Trachyspora, 156 pine gall rusts, 7
Phakopsora, 32, 39, 40, 43, 77, 84-85, 89, 91 Asian hard pine gall rust
ampelopsidis, 19, 85 Cronartium orientale, 3, 7,27, 66

219
 Rosaceae, 11, 34, 39, 40, 47, 58, 59, 100, 109, 115, 116, 146, Sclerotium betulinum, 54
148, 149, 150, 151, 152, 153, 154, 156, 157, 164 Scopella, 98
rose rust Securinega
Phragmidium mucronatum, 25, 39, 154 ramiflora, 83
Phragmidium rosae-multiflorae, 3, 5,21 suffruticosa, 83
rubber-vine weed, 7 Selaginella, 1
Rubiaceae, 59, 80, 84, 89, 95, 96, 173 Sempervivum globiferum, 178
Rubus, 7, 11, 25, 109, 149, 150, 153, 154 Sempervivum rust
fruticosus, 153 Endophyllum sempervivi, 25, 178
rigidus, 151 Senecio cinerarioides, 180
rush skeletonweed, 7 Septobasidiales, 2, 95
rust fungi, general information Septobasidium, 1
anamorph genera, key and descriptions, 36-40 Sida flavescens, 185
classification, 30-35 skeletonweed, rush, 7
collection, 11 Skierka, 35, 48, 183, 190
correlated species and Tranzschel’s Law, 26 canarii, 190
cytology and nuclear cycles, 26-28 cristata, 190
diseases, agricultural and horticultural crops, 2-5 holwayi, 190
diseases, forest trees, 3, 6-7
robusta, 190
economic importance, 2-3, 7
Smilacaceae, 70
families, key and descriptions, 30-35 Smilax sieboldii, 5, 70
germ pores, 23-24
smuts, 1, 2
haustoria, 24
snapdragon rust
holomorph genera, keys by family, 42-48
Puccinia antirrhini, 171
host-rust relationship, 7, 11
Solanaceae, 159, 184, 186
hyphae, 24
Solanum argenteum, 184
identification, 11-12
Sorataea, 11, 33, 44, 107, 109, 112, 115
key to anamorph genera, 36-40
amiciae, 115
key to families, 30-31
cerasi, 115
keys to holomorph genera, 42-48
padi, 115
life cycle variations, 24-25
Sorbus, 11
life cycles, abnormal, 26
aucuparia, 100
phylogeny, 2
sorghum rust
preservation, 11
Puccinia purpurea, 171
relationship to other groups, 1-2
southern pine cone rust
spore states, definitions and morphology, 14-24
Cronartium strobilinum, 7, 66
symptoms, 4-5, 7
Rutaceae, 104
southwestern pine cone rust
Cronartium conigenum, 7, 66
soybean rust
S Phakopsora pachyrhizi, 3, 4, 84, 85
safflower rust spermogonium, definition and morphology, 14, 16
Puccinia calcitrapae, 171 Sphacelotheca, 2
Sakaguchia, 1 Sphaerophragmiaceae, 22, 30, 33
Salacia, 188 Sphaerophragmium, 34, 46, 135, 142
prinoides, 39, 188 acaciae, 15, 142
Salicaceae, 11 artabotrydis, 142
Salix, 11, 74 Sphenospora, 34, 46, 143
Salvia purpurea, 170 copaiferae, 132, 143
Sanguisorba officinalis, 157 dalbergiellae, 143
Sapindaceae, 138 pallida, 143
Sapindales, 190 saphena, 143
Sapotaceae, 44, 90, 98, 101 smilacina, 143
Saxifraga, 11,74 xylopiae, 143
aizoides, 37 Spiraea ulmaria, 146
Saxifragaceae, 37, 87 Spirechina, 153
Scalarispora, 32, 43, 87, 88 spore states, definitions and morphology, 14-24
hashiokai, 88 aecium, 16, 20
Schizaeaceae, 103 basidium, 22

222,
 lagerheimiana, 161 R
longipedicellata, 172 Ramakrishnania, 47, 173
malvacearum, 25, 27, 171
ixorae, 173
melanocephala, 2,171, 172
Randia, 80
menthae, 2, 171
Ranunculaceae, 93, 100, 115, 116, 145, 180
mesnieriana, 26
Ranunculus, 180
monoica, 7, 26
raspberry rusts
mucronata, 154
Gymmoconia peckiana, 25, 26, 150
nivea, 170
Kuehneola uredinis, 15, 153
pampeana, 26, 178
Ravenelia, 11, 34, 45, 83, 128, 129, 136, 137, 139-141, 144
peckiana, 150 bella, 140
polysora, 2,172 bezerrae, 141
prosti, 172 breyniae, 136
pruni-persicae, 115 caesalpiniae, 137
psidii, 3,5
corbula, 140
punctiformis, 7 cumminsii, 140
purpurea, 171
dieteliana, 141
recondita, 2, 3, 4,19, 26, 171, 172
echinata, 139
rubigo-vera, 26 epiphylla, 139
scirpi-ternatani, 172
fragrans var. fragrans, 139
scleirae, 172
glandulosa, 139
sherardiana, 26
hermosa, 140
solidipes, 172 holwayi, 140
sorghi, 2,171, 172
lonchocarpi, 141
spegazziniana, 172
lonchocarpicola var. mera, 15
striiformis, 2,171
mera, 15
tanaceti, 172
mexicana, 140
thlaspeos, 26
pileolarioides, 141
ulmariae, 146
pringlei, 25
vexans, 14, 21
sarmentoi, 139
wolgensis, 172
spegazziniana, 140
xanthit, 27
stevensti, 140
Pucciniaceae, 22, 30, 31, 33, 34, 47, 107, 158-177
taxensis, 144
Pucciniastraceae, 30, 31, 42, 51-60
Raveneliaceae, 22, 30, 31, 33-34, 45-46, 123-147
Pucciniastrum, 31, 39, 42, 51, 58
Raveneliae, 33
epilobii, 58
resin top rust
goeppertianum, 7 Endocronartium pini, 3, 25, 68
Pucciniosira, 25, 34, 48, 184, 186 Rhamnaceae, 79
cornuta, 186 Rhamnales, 190
pallidula, 186 Rhamnus, 26
triumfettae, 186 Rhododendroideae, 62
Pucciniosiraceae, 25, 30, 34-35, 48, 179-187 Rhodosporidium, 2
Pucciniosireae, 25, 34, 178 Rhodotorula, 1
Pucciniostele, 32, 43, 87, 88 Rhus, 19, 120
clarkiana, 3, 5, 87 Ribes, 11, 74
hashiokai, 87, 88 rice, 2
mandschurica, 87 Roestelia, 22, 37, 38, 40, 164, 166
Pyrola asarifolia, 8 cancellatum, 40
Pyroloideae, 62 distorta, 166
Pyrus, 11
Roestelia-type
communis, 40 Gymnosporangium, 164
pyrifolia, 9 Rosa, 11, 154
centifolia, 154
Q eglanteria, 39
Qualea multiflora, 91 multiflora, 5, 21

221
 Rosaceae, 11, 34, 39, 40, 47, 58, 59, 100, 109, 115, 116, 146, Sclerotium betulinum, 54
148, 149, 150, 151, 152, 153, 154, 156, 157, 164 Scopella, 98
rose rust Securinega
Phragmidium mucronatum, 25, 39, 154 ramiflora, 83
Phragmidium rosae-multiflorae, 3, 5,21 suffruticosa, 83
rubber-vine weed, 7 Selaginella, 1
Rubiaceae, 59, 80, 84, 89, 95, 96, 173 Sempervivum globiferum, 178
Rubus, 7,11, 25, 109, 149, 150, 153, 154 Sempervivum rust
fruticosus, 153 Endophyllum sempervivi, 25, 178
rigidus, 151 Senecio cinerarioides, 180
rush skeletonweed, 7 Septobasidiales, 2, 95
rust fungi, general information Septobasidium, 1
anamorph genera, key and descriptions, 36-40 Sida flavescens, 185
classification, 30-35 skeletonweed, rush, 7
collection, 11 Skierka, 35, 48, 183, 190
correlated species and Tranzschel’s Law, 26 canarii, 190
cytology and nuclear cycles, 26-28 cristata, 190
diseases, agricultural and horticultural crops, 2-5 holwayi, 190
diseases, forest trees, 3, 6-7 robusta, 190
economic importance, 2-3, 7 Smilacaceae, 70
families, key and descriptions, 30-35 Smilax sieboldii, 5,70
germ pores, 23-24 smuts, 1, 2
haustoria, 24
snapdragon rust
holomorph genera, keys by family, 42-48 Puccinia antirrhini, 171
host-rust relationship, 7, 11
Solanaceae, 159, 184, 186
hyphae, 24
Solanum argenteum, 184
identification, 11-12
Sorataea, 11, 33, 44, 107, 109, 112, 115
key to anamorph genera, 36-40 amiciae, 115
key to families, 30-31
cerasi, 115
keys to holomorph genera, 42-48
padi, 115
life cycle variations, 24-25
Sorbus, 11
life cycles, abnormal, 26
aucuparia, 100
phylogeny, 2
sorghum rust
preservation, 11
Puccinia purpurea, 171
relationship to other groups, 1-2
southern pine cone rust
spore states, definitions and morphology, 14-24
Cronartium strobilinum, 7, 66
symptoms, 4-5, 7
southwestern pine cone rust
Rutaceae, 104
Cronartium conigenum, 7, 66
soybean rust
S Phakopsora pachyrhizi, 3, 4, 84, 85
safflower rust spermogonium, definition and morphology, 14, 16
Puccinia calcitrapae, 171 Sphacelotheca, 2
Sakaguchia, 1 Sphaerophragmiaceae, 22, 30, 33
Salacia, 188 Sphaerophragmium, 34, 46, 135, 142
prinoides, 39, 188 acaciae, 15, 142
Salicaceae, 11 artabotrydis, 142
Salix, 11, 74 Sphenospora, 34, 46, 143
Salvia purpurea, 170 copaiferae, 132, 143
Sanguisorba officinalis, 157 dalbergiellae, 143
Sapindaceae, 138 pallida, 143
Sapindales, 190 saphena, 143
Sapotaceae, 44, 90, 98, 101 smilacina, 143
Saxifraga, 11,74 xylopiae, 143
aizoides, 37 Spiraea ulmaria, 146
Saxifragaceae, 37, 87 Spirechina, 153
Scalarispora, 32, 43, 87, 88 spore states, definitions and morphology, 14-24
hashiokai, 88 aecium, 16, 20
Schizaeaceae, 103 basidium, 22

222,
 germ pores, 23-24 Toddalia aculeata var. gracilis, 104
spermogonium, 14, 16-20 Tormentilla erecta, 148
surface sculpture, 22—23 Tournefortia, 187
telium, 15, 16, 22 Trachyspora, 34, 46, 156
uredinium, 16 alchemillae, 156
Sporidiomyces, 1 intrusa, 15, 156
Sporobolomyces, 1 keniensis, 156
spruce broom rust vestita, 156
Chrysomyxa arctostaphyli, 7, 8, 25, 62 Tranzschelia, 33, 37, 45, 93, 116
spruce bud rust cohaesa, 116
Chrysomyxa woroninit, 7, 8, 62 discolor, 3, 116
spruce needle, bud and cone rusts pruni-spinosae, 116
Chrysomyxa, 7,8, 11, 62-63 thalictri, 25
Spumula, 34, 45, 144 Tranzschel’s Law and correlated species, 26
bottomleyae, 144 Triactella, 133, 146
heteromorpha, 144 holwayi, 133, 146
quadrifida, 144 pulchrum, 133
Stellaria uliginosa, 53 Trichopsora, 34, 48, 182, 187
Stereospermum tournefortiae, 187
cylindricum, 21, 110 Triphragmiopsis, 34, 46, 138, 145, 146
suaveolens, 82 isopyrl, 145
Stereostratum, 34, 47, 161, 174 Jeffersoniae, 145
corticioides, 2,4, 174 laricinum, 145
Stereum, 174 Triphragmium, 34, 46, 133, 138, 145, 146
Stilbechrysomyxa, 31, 61, 62 acaciae, 142
Stipa rust binatum, 130
Puccinia interveniens, 25, 26 deglubens, 111
Stomatisora geophilicola, 72 echinatum, 138
stripe or yellow rust of wheat pulchum, 146
Puccinia striiformis, 2, 171 ulmariae, 146
sugarcane rusts Triticum
Puccinia kuehnii, 2,171 aestivum, 4, 21,171
Puccinia melanocephala, 2,171, 172 vulgare, 171
sunflower rust Triumfetta, 186
Puccinia helianthi, 3, 14,25, 27,171 Tsuga, 31, 40, 57, 58, 59, 62
Symphytum, 53 canadensis, 74
symptoms of rust fungi, 4-5, 7 diversifolia, 59
Tunicopsora, 81
T
Taphrina, 2 u
Taphrinales, 2 Uraecium, 39, 40
Tegillum fimbriatum, 101 Uredendo, 40
Teliomycetes, 1 Uredinales, 1, 2, 3, 95
telium, definition and morphology, 16, 22 Urediniomycetes, 1, 2, 3, 24
Tephrosia virginiana, 139 uredinium, definition and morphology, 16
Thalictrum, 19, 25 Uredinopsis, 2, 11, 16, 31, 39, 42, 56, 60, 73
Thalictrum rust filicina, 60
Tranzschelia thalictri, 25 longimucronata, 60
Thekopsora, 16, 31, 39, 42, 51, 57, 58, 59 osmundae, 60
areolata, 59 pteridis, 60
minima, 59 Uredo, 30, 36, 38, 40, 52, 56, 58, 59, 60, 64, 84
Thermopsis, 170 appendiculatus, 175
thistle, Canada, 7 cupressicola, 164
Thuja, 7 eichhorniae, 7
Thujopsis, 7 euphorbiae-helioscopiae, 40
dolabrata, 5 ingae, 147
Thymeleaceae, 90 lindsaeae, 37
Tiliaceae, 34, 127, 184, 186 miniata, 39

DLS
 obtusa, 148 Triphragmiopsis, 145
rhinanthacearum, 64
Triphragmium, 146
sempervivi, 178 Uromyces, 175
tremellosa var. campanulae, 64
Uromycladium, 121
vetus, 1 Uropyxidaceae, 33
Uredo-type, 22, 36 Ypsilospora, 147
Achrotelium, 90 Zaghouania, 177
Atelocauda, 118 Uredopeltis, 32, 43, 80, 89
Blastospora, 70 congensis, 89
Ceraceopsora, 93 dominicana, 89, 180
Chaconia, 94 guettardae, 89
Chaconiaceae, 33 Uredostilbe, 36, 38, 40
Chrysocelis, 72 pistila, 40
Cumminsiella, 163 Uredostilbe-type
Dicheirinia, 130 Batistopsora, 77
Didymopsorella, 104 Corbulopsora, 162
Diorchidium, 132 Miyagia, 169
Dipyxis, 105 Phakopsoraceae, 43
Esalque, 133 Pucciniaceae, 34
Frommeélla, 148 Uromyces, 7,11, 16, 22, 34, 37, 40, 47,98, 118, 139, 162, 169,
Gerwasia, 149
175-176, 182
Goplana, 95 aleuropodis-repentis, 176
Gymnosporangium, 164 amurensis, 176
Hamaspora, 151
appendiculatus, 2,25, 175, 176
Hapalophragmium, 134 betae, 175
Kernkampella, 136 bicinctus, 118
Kuehneola, 153 bidenticola, 176
Lipocystis, 137 ciceris-arietinus, 2,175
Maravalia, 98 commelinae, 176
Masseeélla, 189 corollocarpi, 176
Mehtamyces, 82 cucullatus, 176
Melampsora, 74 dactylidis, 26, 176
Melampsoraceae, 32 dianthi, 175
Mikronegeria, 73 digitatus, 118
Mikronegeriaceae, 32 eragrostidis, 176
Newinia, 110 erythronii, 176
Nyssopsora, 138 gemmatus, 176
Phakopsora, 84 glycyrrhizae, 176
Phakopsoraceae, 32
halstedii, 176
Phragmidiaceae, 34
koae, 118
Phragmidiella, 86
linearis, 176
Phragmidium, 154
pisi, 7,175
Phragmopyxis, 111
seditiosus, 176
Pileolaria, 120
socius, 176
Pileolariaceae, 33
sonorensis, 176
Polioma, 170
striatus, 25, 175, 176
Porotenus, 112
trifolii-repentis, 175
Prospodium, 113
viciae-fabae, 176
Puccinia, 171
vignae, 176
Pucciniaceae, 34
Uromycladium, 33, 45, 118, 121, 129
Ravenelia, 139
aritimum, 121
Raveneliaceae, 33
cubense, 129
Skierka, 190 simplex, 121
Sorataea, 115
fepperianum, 7
Sphenospora, 143
Uropyxidaceae, 22, 31, 33, 44-45, 102-117
Spumula, 144
Uropyxioideae, 33
Stereostratum, 174
Uropyxis, 11, 33, 44, 107, 108, 117, 163
Tranzschelia, 116 amorphae, 117

224
 witches’ brooms, 7
fraxini, 108
Acacia broom rust
heterospora, 117
Ravenelia pringlet, 25
holwayt, 117
Acacia koa broom rust
quitensis, 106
Atelocauda koae, 3,7, 26, 118, 119
sanguinea, 163
fir broom rust
Urtica laetevirens, 180
Urticaceae, 90, 180
Melampsorella caryophyllacearum, 7, 25, 39, 53
Ustilaginales, 1, 2 spruce bud rust
Chrysomyxa woroninii, 7, 8, 62
Ustilaginomycetes, 3
spruce yellow witches’ broom
Chrysomyxa arctostaphyli, 7,8, 25, 62
V Thuja and Thujopsis rusts
Vaccinioideae, 62 Blastospora betulae, 3, 5,7, 10), 7
Vaccinium, 7,51, 57
myrtillus, 57
vitis-idaea, 8, 51 xX
Verbenaceae, 11, 101, 112, 113, 153, 161 Xenodochus, 34, 47, 110, 157
Vitaceae, 87, 88, 95 carbonarius, 157
Vochysiaceae, 91 clarkiana, 87
minor, 157
Ww Xyloma
areolatum, 59
Wardia, 36, 38, 40
euphorbiae, 74
vastatrix, 40
Wardia-type Xylopia, 102
Chaconiaceae, 44 aethiopica, 181
Desmella, 103
Edythea, 106 Y
Hemileia, 96 yellow or stripe rust of wheat
Uropyxidaceae, 33 Puccinia striiformis, 2,171
water hyacinth, 7 yellow witches’ broom of spruce
weed control, 7 Chrysomyxa arctostaphyli, 7, 8, 25, 62
western gall rust of hard pines Ypsilospora, 34, 45, 147
Endocronartium harknessii, 3, 6, 7, 25, 68 africana, 147
wheat rusts baphiae, 147
wheat leaf rust tucumanensis, 147
Puccinia recondita, 2, 4,19, 26, 171, 172
wheat stem rust
Puccinia graminis, 2, 4,14, 0), Pl, Py, 3, WAM Z
wheat stripe or yellow rust Zaghouania, 34, 47,177
Puccinia striiformis, 2, 171 oleae, 177
white pine blister rust phillyreae, 177
Cronartium ribicola, 3, 6, 14, 18, 19, 25, 66, 67 Zingiberaceae, 72,98
willow rusts Ziziphus oenopolia, 79
Melampsora, 3, 74-75 Zygophyllaceae, 139

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