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A TAXONOMIC ANALYSIS OF THE SPECIES OF UROMYCES ON LEGUMES IN BRAZIL

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Authors Almeida, Rogerio Tavares de, 1941-

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Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 76-1603 ALMEIDA, Rogerio Tavares de, 1941- A TAXONQMXC ANALYSIS OF THE SPECIES OF ' URGMYEES ON LEGUMES IN BRAZIL.

The University of Arizona, Ph0D0? 197S Agriculture, plant pathology

Xerox University Microfilms t Ann Arbor, Michigan 48106

THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. A TAXONOMIC ANALYSIS OF THE SPECIES OF UROMYCES ON LEGUMES IN BRAZIL

by Rogerio Tavares de Almeida

A Dissertation Submitted to the Faculty of the DEPARTMENT OF PLANT PATHOLOGY In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA

19 7 5 THE UNIVERSITY OF ARIZONA

GRADUATE COLLEGE

I hereby recommend that this dissertation prepared under my

direction by Rogerio Tavares de Almeida

entitled A Taxonomic Analysis of the Species of Uj^ongces

on Legumes in Brazil

be accepted as fulfilling the dissertation requirement of the

degree of Doctor of Philosophy

bf-u?hr Dissertation Director Date

After inspection of the final copy of the dissertation, the

following members of the Final Examination Committee concur in its approval and recommend its acceptance:""

AT 7

i < ht

"This approval and acceptance is contingent on the candidate's adequate performance and defense of this dissertation at the final oral examination. The inclusion of this sheet bound into the library copy of the dissertation is evidence of satisfactory performance at the final examination. STATEMENT BY AUTHOR

This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the Univer sity Library to be made available to borrowers under rules of the Libraryo Brief quotations from this dissertation are allow able without special permission, provided that accurate acknowledgment of source is made0 Requests for permission for extended quotation from or reproduction of this manu script in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his judgment the proposed use of the material is in the interests of scholarship., In all other instances, however, permission must be obtained from the author®

SIGNED> ch MLjufic* DEDICATION

The Author Dedicates This Dissertation To His Parentsj Moises Almeida and Francisca T. de Almeida and To His Wife and Sont Conceicao C. de Almeida and Rogerio T. de Almeida Filho

* « • 111 ACKNOWLEDGMENTS

The author wishes to express his sincere gratitude to Dr. Re Le Gilbertson and Dr0 Go B0 Cummins, under whose guidance and supervision this work was done, and for their assistance and counselo Thanks are given also to Dr0 S0 M.

Alcorn, Dr0 R0 L0 Caldwell, and Dr0 M0 R0 Nelson for their advice and help in preparation of this manuscript; to Dre E. L. Nigh Jr., Read of the Department of Plant Pathology of The University of Arizona, for his assistance and encourage ment; to Profe Jose Ilo da Ponte Vasconcelos and Prof. J. Julio da Ponte Filho of the Universidade Federal do Ceara, Brazil, for their encouragement and orientation in my ca reer, and to the people in the various Institutions in Brazil, the United States, Sweden, Argentina, and Germany for the loan of specimens that made possible this study. Finally, the author is grateful to the Universidade Federal do Ceara, the Agency for International Development (AID), and the Institute of International Education (IIE) for fi nancial support.

iv TABLE OP CONTENTS

Page

LIST OF ILLUSTRATIONS vi LIST OP TABLES ix

ABSTRACT o.ooo.ooooe.oooooo.oeoooooooeoo.eooooo.oo.oo.o. X INTRODUCTION 000000000.0.00000000000000000000000000.0000 1 LITERATURE REVIEW00000000.000000000.00.0000.oooooeoo... 3 MATERIALS AND METHODS o«.o.oooo.»oe..ooioo.oo..ooao...o« 8

GENERIC DESCRIPTION. 000000.00000.00088000000.0000.00.0. IJL KEY TO SPECIES OP UROMYCES ON LEGUMES IN BRAZIL BASED ON UREDINIA AND TELIA 15 DESCRIPTION OF SPECIES OF UROMYCES ON LEGUMES IN BRAZILo .00..0000.00 .00.00.000000000000. 000.0000000000 32 SPECIES OF UROMYCES ON LEGUMES IN BRAZIL NOT VALID OR TRANSFERRED TO OTHER GENERA..0». 68 DISCUSSION AND CONCLUSIONS 69 LITERATURE CITED 86

v LIST OF ILLUSTRATIONS Figure Page 1. Spermogoniums schematic (after Cummins, 1959) • 13 2. Aecium: schematic (after Cummins, 1959)««•»°9»•••«•• 13 3. Urediniums schematic (after Cummins, 1959)• ° ®»<>»•«« 13

4« Teliums SChematiCo ooooeoocoooooooooooooooooooooooeoo 13 5. Leaf spots on Bauhinia forficata with spermogonia and telia of Uromyces viegasii (X *5)° •>»0»0 0 0 0 0 B 0 0 0 e 14 6. Aecia of Uromyces orbicularis on leaves of DeSmodjum Sp 0 (X 05 )°QO<,O0OO°ooooooooooooooooeoooooo l4

7. Uredinia of Uromyces appendiculatus var0 appendiculatus on leaf of Phaseolus vulgaris (X <>5)»»0 B 0 •> 14 8. Telia of Uromyces dietelianus var. nervicola on leaf of Bauhinia forf icata (X .5)*>*«» 14

9. Teliospores of Uromyces appendiculatus var0 appendiculatus with few to many verrucae (X 750)e>o®a 23 10. Smooth teliospores of Uromyces appendiculatus var © "bra siliensis (X 750) 000000000000000000000000000 23 11. Verrucose-striate teliospores of Uromyces StriatUS (X 750)oo8oooeoooeoeoooeoe«ooooeeosooo80oer 23 12. Reticulate teliospores with wall bilaminate and one urediniospore (center) of Uromyces ba uhmiae (X 75O)oooooooo0e0OooooooooQO0QooeoQooGooa 23 13. Teliospores (top) and urediniospores (bottom) of Uromyce s belemensis® Qooooooeooooooooaoooeeooooeoo 24 14. Teliospores (top) and urediniospores (bottom) of Uromyces neuro c a r p i»ooosoooooooooooeooooooooooooo 24 15. Teliospores (top) and urediniospores (bottom) °? Uromyces appendiculatus var» appendiculatus.«..B, 24

vi vii

LIST OF ILLUSTRATIONS—Continued Figure Page 16. Teliospores (top) and urediniospores (bottom) of Uromyces appendiculatus var. brasiliensis...... 2b 17. Teliospores (top) and urediniospores (bottom) of

Uromyces lathyrinus0qoooooooooooooooooqoooooqoooooqo 23 18. Teliospores (top) and urediniospores (bottom) of Uromyces viciae-^abae0oooooooooooooooooooooooooocooo 23 19• Teliospores (top) and urediniospores (bottom) of Uromyces trifolii var0 tnfoliio 00»«»».«»««0»»0 0 5»o 25 20. Teliospores (top), urediniospores (bottom)„ and one paraphysis (right) of Uromyces unionensiso0000o0 26 21. Teliospores (top and bottom at left)„ uredinio spores (bottom at right), and paraphyses (top at right) of Uromyces hedysari-paniculatio„0„„ce.0000a. 26 22. Teliospores (top) and urediniospores (bottom) of Uromyces bradburyae000000000000000000000000000000000 26 23. Teliospores (top) and urediniospores (bottom) of

UrOmyc e S S tria tUS O OOOOOOOOOOOOOOOOQOOOOOOOOOOOOOQQOQ 27

2b, Urediniospores of Uromyces crotalariae e 9 c 0«, 0 0». 0 a e c e 27 25« Teliospores (top) and urediniospores (bottom) of Uromyces pisi.000000000.0000000000000000000.00.0000. 26. Teliospores (top) and urediniospores (bottom) of Uromyces anthy11idiSooooooooooooooooooo&oooooooooooo 28 27. Teliospores (top and left) and aeciospores (right) of Uromyces orbicularxso0000000000000000000000000000 28

28. Teliospores of Uromyces nerviphilus0„0000Be000ae00e8 28

29* Teliospores of Uromyces goyazensis0 ooooooooooooooooo 29

30. Teliospores of Uromyces floraliso 0 0 0 0 •>. 0.8«».»..«0 o 0 29

31. Teliospores of Uromyces viegasii...... ao... 29 LIST OF ILLUSTRATIONS—Continued Figure Page 32o Teliospores (top) and urediniospores (bottom) of Uromyces bauhiniaeo eeooooaaooaoeoescovoooeoosveoooQe 29

33* Teliospores (right and left) and urediniospores (middle) of Uromyces foveolatus0cooeoeeeoooooQooo»s« 30 3^. Teliospores (top) and urediniospores (bottom) of Uromyces dietelianus var e dietelianuso0 0 o 0« 0 0 0 o 0«o so 30 35. Teliospores (top) and urediniospores (bottom) of Uromyces dietelianus var a nervicolae0000000000000000 30 36. Teliospores (top) and urediniospores (bottom) of Uromyces vicosensisa#o«oo««oooooeo«»ooooooe88oooo®o® 30 37. Teliospores (top) and urediniospores (bottom) of UromyceS ca staneus 9 ooocoesocoooooooooocoooooeoooeooo 31 38. Teliospores (top) and urediniospores (bottom) of Uromyces desmodiicola var® desmodiicola» ° ..<>«.• 31 39. Teliospores (top) and urediniospores (bottom) of Uromyces desmodiicola varo desmodiioooeoeooeoeoeooeo 31 *K>. Geographical distribution of the species of Uromyces on legumes m Brazil®•o*«*o«o*oo«aaa**«o*«* 83 LIST OF TABLES Table Page 1. Summary of the results of a taxonomic study of the species of Uromyces reported on legumes in Brazil#• a****>*eo**o*«*«a«s*«o*e«**********oo****** 72 2. Life cycles and spore stages of the species of Uromyces on legumes in Brazil* •*** •••*•>• 0 •*•••••••• 78

ix ABSTRACT

The genus Uromyces„ established by Unger in I833, is the second largest genus of the rust fungi„ being exceeded only by Puccinia. Taxonomically, it is placed in the family

Pucciniaceae9 order Uredinales, and in the class Basidiomyceteso In Brazil over one hundred species of rust fungi par asitize legumes and most of these belong to the two genera Uromyces and Raveneliao Even without considering the large number of species, Uromyces is the most important genus of rusts on legumes in Brazil because of the number of species of economic impor

tance e Among the species that attack cultivated plants are

Uo anthyllidis (Grev0) Schroet0, U0 viciae-fabae (Pers0)

Schroete (broad bean rust), U. appendiculatus (Pers0) Ung. var. appendiculatus (bean rust), U® striatus Schroeto (al

falfa rust), Uo pisi (DCe) Otth (pea rust), and Uo trifolii

(Hedwe f0 ex DC«) Wint0 var0 trifolii and U0 nerviphilus

(Grog.) Hotson (clover rusts)0 Bean rust is the most impor tant because of its wide distribution and the losses it

causes to this crop0

x A review of the literature indicated that 37 species of Uromyces, including two varieties of U. appendiculatus (Pers.) Ungo, occur on legumes in Brazil; a taxonomic analy sis was considered a necessity in order to verify the iden tity of these species and apply the correct nomenclaturee The species of Uromyces reported on legumes in Brazil were analyzed taxonomically» The basis for identi fication of species included studies of the wall thickness and ornamentation* color, size, and shape of teliospores and urediniospores, pedicel features of teliospores, and the number and position of germ pores in urediniospores0 Pa- raphyses, spore forms present in the life cycle, host symp toms or signs, and host specificity were additional charac ters that aided in species identificatione Analytical keys and complete descriptions were pre pared to facilitate identification of Uromyces species0 Nomenclatural problems were resolved through study of type specimens in all cases possible. Illustrations of species were made by drawings and photomicrographs.. As a result of this study, the author concluded that:

1. Twenty-nine species of Uromyces occur on legumes in Brazil. Among them are two varieties of U. appendiculatus (Pers.) Ung., two varieties of U. desmodiicola Jorst., two varieties of U. dietelianus xii

Paz., and one variety of U. trifolii (Hedw. f ex DC.) Wint. 2* U« bauhiniicola Arth., U. corru^atus Spego, U.

johowii Diet. & Neger, U. imperfectus Arth0, U.

nordensk.joldii Diet0, U. occidentalis Diet., U. superfixus Vesto , and U. trifolii-megalanthi (Diet. & Neger) Jacks. & Holw., probably do not occur in Brazil. 3* U* verus Jacks. & Holw. is identical to U„ floralis Vest., and U. pannosus Vest, and U. dietelianus Paz. represent a single species. 4. Among the taxa recognized, U. vicosensis, U. viegasii, U» desmodiicola Jorst. var« desmodii, and U. dietelianus Paz. var. nervicola are considered to be new, 5. U. phaseoli longepedunculati Viegas nom. nudum was recognized as a distinct variety and named U. appendiculatus (Pers.) Ung. var. brasiliensis var. nov. INTRODUCTION

Uromyces is the second largest genus of the rust fungi, being exceeded in number of species only by Fuccinia (Cummins, 1959) <> In Brazil over one hundred species of rust fungi par asitize legumes and most of these species belong to the two

genera Uromyces and Ravenelia0 Thirty-seven species of Uromyces have been reported on legumes in Brazil® Among them two varieties of U® appendiculatus have been recognized® Even without consider ing the large number of species, Uromyces is the most impor tant genus of rusts on legumes in Brazil because of the

number of species of economic importance0 Among the species that attack cultivated plants in Brazil, are Uromyces

anthyllidis (Grev«) Schroeto, U0 viciae-fabae (Pers«)

Schroeto (broad bean rust), U0 pisi (DC®) Otth (pea rust),

Ua appendiculatus (Pers») Ung„ varQ appendiculatus (bean

rust), Uo striatus Schroeto (alfalfa rust), and U0 trifolii

(Hedwe f ex DCo) Wint® vare trifolii and U0 nerviphilus

(Grog0) Hotson (clover rusts)© Bean rust is the most impor tant because of its wide distribution and the losses it

causes to this cropa According to Galli et alo (1968), bean rust occurs in all regions where beans (Phaseolus

1 2

vulgaris L.) are grown in Brazilo Because of the high vari ability of this species„ many surveys to identify physio logical races have been made in several states of Brazil (Menezesp 1952s Dias and Costa, 1968; Junqueira Netto, Athow and Vieira, 19&9? Augustin and Costa, 1971)® These studies were begun to provide a more intelligent control of this rust fungus through the use of resistant varieties0 A search of the literature indicates that a taxonomic analysis of the species of Uromyces is needed since there is no work on this aspect which includes all species re ported on legumes in Brazilo Also9 such an analysis should be useful for plant pathologists, mycologists and others interested in legumes in Brazil® Therefore, the objectives of this research ares a) to describe and illustrate the species of Uromyces reported on legumes in Brazil and to de termine if their nomenclature is correct; b) to prepare keys to facilitate identification of these Uromyces species; c) to analyze the status of species of rusts on legumes in Brazil that were initially described as Uromyces species but later were transferred to different genera of rust fungij d) to report the geographical distribution of the species of Uromyces on legumes in Brazilo LITERATURE REVIEW

The genus Uromyces was established by Unger in I833 (Cummins, 1971s Wilson and Hendersons 1966)0 Guyot (1938) pointed out that the concept of this genus originated with Link in 1816 but at that time Uromyces was considered a sub genus of Pucciniaa Taxonomicallyp it is placed in the fami ly Pucciniaceaes order Uredinaless and in the class Basidio- mycetes# Over the years a number of articles have been writ ten concerning Uromyces species on legumes., These include:

Hariot (1892)f Vestergren (1905) (a monograph of the genus Uromyces on legumes of the genus Bauhinia in which 17 spe cies are described with illustrations of teliospores and ure< diniospores), P0 Sydow and Sydow (1910)„ Guyot (1957) (infor mation on the world-wide distribution of species of Uromyces that occur on legumes)® Guyot also made an extensive review of literature and synonymy of these specieso Species of Uromyces on legumes that occur in North America and Britain were described by Arthur (1907-1927» 1962) and Wilson and

Henderson (1966)9 respectively.,

There are many papers referring to the occurrence

0;f Uromyces on legumes in Brazil but without descriptions of speciese With exception of Viegas (19^58? 19^5b) and

3 4-

Albuquerque and Figueiredo (1971)all papers describing species of Uromyces on legumes in Brazil were published by non-Brazilians researcherso A survey of the literature indicates that 37 Uromyces species,, including two varieties of U. appendiculatus, occur on legumes in Brazilo Uo dietelianus

Pazschke (1891)5the first species described in Brazils, was

found on Bauhinia sp0 collected at Tubarao0 Santa Catarinae U, appendiculatus was reported in Brazil on Phaseolus sp® by Pazschke (1892), since then it has been widely disseminated

in this country (Galli et alo9 1968)0

Hennings (1895) described two new species, U.

goyazensis (from material obtained in Goias) and Ua bauhiniae (from a specimen collected at Paranaiba, Minas

Gerais) on Bauhinia sp0 In the same year, Dietel (1895) de

scribed Uo neurocarpifl a new species on legume, from materi

al' collected by Lhotsky in Bahia0 Hennings (I896) described Aecidium desmodii on

Desmodium sp0, from a specimen collected in Rio de Janeiro, This fungus is considered (Lindquist and Costa Neto, 1963)

the aecial stage of U0 hedysari-paniculati (Schw,) Farl., a species widely distributed in Brazilo

Dietel (1897) described two new species, U. clavatus

on Lathyrus magellanicus Lam0 and U« orbicularis on

Desmodium spe, from specimens collected at Serra Geral. 5

In the same year, according to P. Sydow and Sydow (1910), Juel described a new species, U. foveolatus on Bauhinia hirsuta Vog0» from a specimen collected by Malme at Cuiaba, Mato Grosso. H° viciae-fabae was reported by Hennings (1902b) on

Vicia faba L0 in Sao PaulOo Vestergren (1905)9 described six new species of Uromyces„ (U0 anthemophilusB Uo floralis.

U. hemmendorffii„ U0 pannosus,, Uc perlebiae, and Uo regius), all occurring on Bauhinia 0 Also„ he reported and rede- scribed Uo praetextus Vest0 nomQ nov0 (originally described by Hennings (1895) as Uo bauhiniae) and Ue foveolatus Juelo

Sydow and Sydow (1907) reported U0 striatus in Sao

Paulo on Medicago sativa L0 and P« Sydow and Sydow (1910) described U, castaneus on Desmodium incanum DC„, from Rio de

Janeiro.

A new species, U0 peraffinisfl was described by Dietel (1922) from material obtained in Rio de Janeiro on

Bauhinia sp0 Jackson (I93I) described as new U„ bradburyae

Jacks0 & HolWo (occurring on Bradbur.ya virginiana (L0) Kuntze and Bo pubescens (Bentho) Kuntze at Campos do Jordao,

Sao Paulo) and U# verus Jacks0 & Holw0 (on Bauhinia rufa

Steud® at Belo Horizonte9 Minas Gerais)0 Alsop Jackson re ported the occurrence of Haplop.vxis crotalariae (Artho) Syd. on Crotalaria vitellina Ker. at Silvestre, Rio de Janeiro. 6

This fungus is now placed in the genus Uromyces by Baxter (1962).

Thurston (19^0) reported U0 imperfectus Arth® on

Bauhinia mirandina Pit'tier at Uberaba p Minas GeraiSo This species was originally described from Nicaragua by Arthur (1920)o

Viegas (19^5a) described a new speciese U®

unionensis„ from material collected at Uniaop Alagoas9 on Desmodiurn (Meibomia) sp® and reported the occurrence on Bra zilian legumes of U® bauhiniicola (originally described from

Mexico by Arthurs 1905), U® superfixus (described from Boli via by Vestergren, 1905)» and U. occidentalis (described from the United States by Dietel 1903)° Viegas (19^5b) also

briefly described a new variety of U0 appendiculatuss Us phaseoli longepedunculati, but without the required latin

diagnosis® Silveira (1951) reported U. pisi on Pisum sativum L. in Brazil®

Jorstad (1959) described as new U® desmodiicolafl on

Desmodiurn albiflorum Salzm® in Rio Grande do Sul8 Lindquist and Costa Neto (1963) reported U® anth.yllidis„ U® ner-

viphilusB and U0 trifolii also from Rio Grande do Sulo Later* Fontoura and Nowacki (1967-70) observed U® trifolii

on Trifolium spp® in Parana0 Recently, Albuquerque and Figueiredo (1971) de scribed a new species, U, belemensis, on Ormosia nobilis Tul. at Belem, Para. In the same year, Watson (1971) re ported U. corrugatus Speg. , U. .johowii Diet. & Neger, U. nordensk.joldii Diet., and U. trifolii-megalanthi (Diet. &

Neger) Jacks. & Holwe but there is no reaffirmation of these records in Brazil. The above citations indicate that many species of Uromyces occur on legumes in Brazil, especially on species of Bauhinia; a taxonomic analysis was considered a necessity in order to verify the identity of these species and apply the correct nomenclature. MATERIALS AND METHODS

Taxonomic analysis was accomplished primarily by microscopic observation of important fungal structures * These structures were mounted either in lactophenol (made by combining one part each of lactic acid, phenol crystals and waters with two parts glycerin) or in a saturated aqueous so lution of chloral hydrate (sometimes heated) to give a better visualization of the important features, especially the germ pores of urediniosporeso Leaf section were made in order better to detect spermogonia and paraphyses of uredinia# The basis for identification of species included studies of the wall thickness and ornamentation, color, size and shape of teliospores and urediniospores, pedicel features of teliospores, and the number and position of germ pores in urediniosporeso Paraphyses,, spore forms present in the life cycle, host symptoms or signs, and host specificity were ad ditional characters that aided in species identification,) Analytical keys and complete descriptions were pre pared to facilitate identification of the Urom.yces species. Nomenclatural problems were resolved through study of avail able type specimens<> Illustrations of species were made by drawings and photomicrographs. Drawings were executed by the author with

8 9 the advice of Dr. Go B, Cummins and photomicrographs were taken with the help of Dr0 R» L0 Gilbertson0 The figures of spores were drawn on a graph paper0 Measurements were made according to Cummins (1971). Thus, spore sizes might be given as (20-)25-30(_3*O x (15-) 20-25(-28) jUo In this example» 25-30 x 20-25 describes the size of the majority of the spores0 Figures in parenthesis are outside of the typical size but are not uncommon,, In a few cases„ descriptions of spermogonia and aecia were adapted from other sources when these structures were not present on specimens available to the author® This also was true for descriptions of teliospores of Uromyces crotalariae (Arthe) Jo Baxters Type specimens of U0 peraffinis Dieta„ U»- regius Vest,,, U0 hemmendorfii Vest09 U„

•perlebiae Vests, and U0 anthemophilus Vests also were un available for comparisons8 Abbreviations of most authors of rust species are according to the USDA (1970) and most host names,, including abbreviations of authorst are in the style of Viegas (1961). Herbarium citations follow Holmgren and Keuken (197*0«

Type specimens are cited only for those Uromyces species described originally from Brazilo

The materials representing the species of Uromyces on legumes in Brazil were obtained from Herbaria of the

United States and other countries0 These include: 10

Mycological Herbarium of The University of Arizona, Tucson, AZ, National Fungus Collections, Beltsville, MD, Arthur Herbarium of Purdue University, V/est Lafayette, IN, Herbar ium of Plant Pathology of Cornell University, Ithaca, NY, Berlin Museum, Germany, Herbarium of the Swedish Museum of Natural History, Stockholm, Sweden, Museo-Instituto Spegazzini, La Plata, Argentina, and Herbario Micologico of the Instituto Agronomico, Campinas, Sao Paulo, Brazil. GENERIC DESCRIPTION

UROMYCES Ung.

Exantheme Pflanzen p0 277o 1833. Type species: Uromyces appendiculatus (Pers.) Ung.

Cycle of development includes spermogonia, aecia, uredinia, and telia? all sori subepidermal in origin and erumpent except in telia of some species that remain covered

by epidermis? species autoecious or heteroecious0 Spermogonium (Pigs 1, 5) deep-seated in the tissue of the host, globoid, with ostiolar filaments and flexuous hyphae often prominent? functional but without feasible taxonomic characters to distinguish species? spermatia very small, hyaline, globoid, or ellipsoido

Aecium aecidioid (Fig0 2, 6) with peridium composed of polygonal cells and with catenulate and usually verrucose aeciospores or aecium uredinoid (Figo 3) with aeciospores borne singly on pedicels? germ pores usually indistincte

Uredinium (Fig0 3, 7) with urediniospores borne sin gly on pedicels, mostly echinulate but sometimes verrucose. Germ pores in urediniospores may differ in number and posi

tion in different species0 Paraphyses in a minority of spe

cies.

11 12

Telium (Pig. 8) with teliospores borne singly on pedicels, l-celled9 with 1 germ pore generally apical? wall mostly obviously pigmented and from smooth to variously sculpturedo Paraphyses in a minority of species. Basidium external bearing hyaline9 ovoid, or reniform basidiospores« Uromyces species attack a great variety of monocot- yledonous and dicotyledonous plants® Besides the Legumi- nosaee the families containing the most host species are the

Gramineae, Liliaceaep Euphorbiaceae, and Compositae0 Uromyces differs from Puccinia only in having uni cellular teliosporeso According to Arthur (1962), Cummins

(1959)f and Wilson and Henderson (1966) this genus is main tained as a matter of convenience and historical precedent. Figure 1. Spermogoniums schematic (after Cummins, 1959). Figure 2« Aecium: schematic (after Cummins, 1959)• Figure 3« Urediniums schematic (after Cummins, 1959). Figure k, Teliumi schematic. Figure 5* Leaf spots on Bauhinia forficata with spermogonia and telia of Uromyces viegasii (X »5)« Figure 6, Aecia of Uromyces orbicularis on leaves of Desmodiura sp0 (X .5)» Figure 7» Uredinia of Uromyces appendiculatus vare appendiculatus on leaf of Phaseolus vulgaris (X »5)c Figure 8« Telia of Uromyces dietelianus var# nervicola on leaf of Bauhinia forficata (X .5). Ik KEY TO SPECIES OF UROMYCES ON LEGUMES IN BRAZIL BASED ON UREDINIA AND TELIA

1. Teliospores smoothsoooooooooooo Section A la Teliospores ornamented <» 0 © o o ° o 0 <> (2)

2. Teliospores verrucose0800sooooo Section B

2e Teliospores reticulate 0 0 0 0 0 o0oo Section C

Section A0 Teliospores smooth 1. Urediniospores with constant number of germ pores (2)

1. Urediniospores with variable number of germ pores (5) 2. Urediniospores with 1 germ pore near the hilumj germ pore, not distinct..oso».oo.e«ale U. belemensis 2. Urediniospores with 2 germ pores (3)

3. Germ pores slightly below the equator0 without caps..

««oooo©ooooogodoooooooooooooooo2o U0 neurocarpi 3. Germ pores equatorial or slightly above the equator, with caps (4) k, Germ pores equatorial or slightly above the equator, indistinct? teliospores mostly with few to many verru-

cae, rarely smootho <> 0 o o o o o. o o. ® 3* U0 appendiculatus

var0 appendiculatus 4. Germ pores slightly above the equator, distinct? telio spores mostly smooth, rarely with few inconspicuous ver rucaeoa..o.....®o#ooo0®®fl®oo*®®^'o uo ap-pendi c u1 atus

var9 brasiliensis 15 16

5. Urediniospores with (5)6-8 scattered germ pores.•••«•••

#«e«coooooooooooooooooooooooo®o5o Uo 5. Urediniospores with 2-5 germ pores (6)

6. Germ pores (3)M5) equatorial or nearly equatorial9 less frequently scattered, or with 3~^ equatorial and 1 at

apexj teliospores smooth0e0000o60 Uo viciae-fabae 6, Germ pores 2-3(^0 equatorial or slightly above the equa tor; teliospores with few to many verrucae or smooths»«•

#»8eoo8oooooooooooooooooo0oeooo7o Ufl var. trifolii

Section B. Teliospores verrucose 1. Uredinia formed (2) 1. Uredinia unknown (11) 2. Uredinia paraphysate (3) 2. Uredinia aparaphysate (4) 3. Urediniospores with k-5 equatorial or approximately equatorial» or less frequently with 3-^ equatorial or approximately equatorial and 1 near or at apex germ poreSo©oooooo©0ooo0oooooooooQ®o8e IJ © umonensxs 3. Urediniospores with (3)^-7(8) scattered germ pores

•«©©OOGOOOOOOOOOOOOOOOOOOOOOOOO 9° U° hedysari-paniculati 4. Urediniospores with 2 germ pores (5) *1-. Urediniospores with variable numbers of germ pores (7) 17

5. Germ pores equatorial or slightly above the equator,, in distinct; teliospores mostly with few to many verrucae,

rarely smoo FCHO oooooo&oobqooo&oo^o IJ © appe ndICUIQ tus

varQ appendiculatus 5# Germ pores slightly above the equator,, distinct (6)

60 Teliospores mostly smooth^ rarely with few inconspicuous

VerrUCa eooooooooGoooooooooooooo^'o varo brasiliensis 6. Teliospores appearing smooth but actually verrucoses with numerous verrucae distributed evenly or irregularly

throughout the surface of spore and umbo®e0o©oooo«aoo»»• eoesooooooeoooooooooooocoooooo 109 TJo bradburyae 7. Germ pores equatorial or slightly above the equator (8) 7. Germ pores scattered (9) 8, Germ pores (2)3-M5)s teliospores verrucose-striate.• • • •

«oooooooooooooooooooooooococeollo Ue StST 13 tUS 8. Germ pores 2-3(4-); teliospores with few to many verrucae or smootho ©ooo©qqo©©©oooqqo©og©7® trifolii var® trifolii 9. Urediniospores with 6-8, mostly 8„ germ pores? telio

spores verrucose-echinulateBool2o U0 crotalariae 9. Urediniospores with 3-8 germ pores (10) 10. Germ pores 3-6; teliospores densely and minutely verru-

COS6oooooooooooooqoooooooooooo13° H® PIS J 10. Germ pores 4-8, mostly 6-8; teliospores sparsely and

strongly verrucoses... 0 e ® o«o <> »1^» U. anth.yllidis 18

11. Teliospores varying from verrucose, with irregular warts sometimes variously united or arranged to form a kind of

labyrinth^ to irregularly reticulate0e00o»seaoe0o®ooe®•a • ®oooooooooooooogooooooooooooo15o JJ.° orbicularis

11. Teliospores minutely and inconspicuously verrucoset ap pearing smootho oooooooooooooo ol6o IJ o nerv lT^hi lus

Section C0 Teliospores reticulate 1, Uredinia unknown (2) 1. Uredinia formed (6) 2. Teliospores varying from verrucose» with irregular warts sometimes variously united or arranged to form a kind of

labyrinth^ to irregularly reticulate00oo.0©.ooc0.ooooo..

«.e»8oeoooooociooooooooooooooool5o JZ® 2. Teliospores not as above (3) 3. Teliospores minutely reticulate» with meshes about 0.5- 1.5 ji diam (U) 3. Teliospores with larger meshes (5)

Teliospores pale brown# slightly and minutely reticulate, appearing smooth* with meshes about 0.5-1 p diam; pedi

cel mostly to 80-100 ju long..0170 U. goyazensis *+. Teliospores golden or light cinnamon-brown, minutely and

uniformly reticulateg with meshes about 1-1„5 p diamj

pedicel usually breaking near the hilume rarely to about 80 p long..00000000.0.00000.0.18. U. floralis 19

5. Telia on flowers, without spots, 3-10 mm long; teliospores 18-25 x 18-22 ]i» mostly without a papilla at apexj wa11 3"™3 ° 5 J-i thick at sideSoooooGoooosooooeooeooo* eooooooooooooooooooooooooooooo190 o anthemophilus 5. Telia on leaves, minute, with brown spots, 1-5 mm diam,

teliospores (15-)18-22(-24) x (13-)l6-l8(-20) jiQ with a

papilla at apex; wall 105-205 p thick at sides*, 0«. e e e»o«•

«C0000000000000000000000000000200 0 vie g3 S i i

60 Uredinia paraphysate (7) 6. Uredinia aparaphysate (8) 7. Urediniospores with 4-5 equatorial or approximately equa torial, or less frequently with 3-^ equatorial or ap proximately equatorial and 1 near or at apex germ pores#•

«oooooooooooooooooooooooooooooo8o U0

7. Urediniospores with (3)4-7(8) scattered germ pores®®„0e.«

o«eoeoooooooooooooooooooooooooo9o Uo 8. Teliospores with wall bilaminate (9) 8. Teliospores with wall single (10) 9. Teliospores distinctly and uniformly reticulate with meshes to about 2-2<>5 ^ diami outer wall distinct, some times thicker at apex but without an umbo? pedicel usu

ally breaking near the hilumo ooooooooooooooooqoooooooqo*

•••••••eeoeeoooeoeeec«oeeoeeeo21a Uo bauhiniae 20

9. Teliospores reticulate, with meshes variable in shape and size, to about 2-2o5 ft diam; outer wall not dis tinct, mostly with an umbo at apex; pedicel to about 20-

25 p. long, basally rugose»». 0..220 U. foveolatus 10, Teliospores with a broad and distinct papilla or corona at apex (11) 10. Teliospores not as above (15) lie Urediniospores with constanct number of germ pores (12) 11. Urediniospores with variable numbers of germ pores (13) 12. Urediniospores 16-23 x 16-20 ja, conspicuously echinulate, with 4 germ pores; teliospores reticulate with meshes variable in shape and sizeooeoo.eooooooooooeoooo. .00.90009.00000000000000000000o23^ Uo reglUS 12. Urediniospores 24-28 x 18-21 }A, sparsely echinulate, with 3 germ pores; teliospores indistinctly and minutely

reticulate.oooe ooooooooooo oo 0 002^0 U_o peraffinis 13. Teliospores wall 3-3«5 ^ thick at sides; urediniospores

with 4, rarely 5« germ poresoeo25« U° perlebiae

13, Teliospores wall 105~3 M "thick at sides (14) 14. Telia scattered; teliospore wall lo5-2o5 ^ thick at sides; urediniospores with (3)^-50 rarely 6 or 7> equa torial or nearly equatorial germ pores00000.0.00.00.o...

...0.00000000.00.00.0000000.000260 Uo var. dietelianus 21

I**, Telia or uredinia mostly arranged linearly along or on the veins? teliospore wall 2-3 P thick at sides?

urediniospores with 3-4 „ rarely 20 5 or 6„ equatorial germ poreso oooooooooeoooooooo ©27 o Uo die te lianus

var0 nervicola 15« Urediniospores with 4 germ poresj teliospores yellow-

brown,, 18-22 x 16-18 p.r, minutely reticulate0»o 0 0 0 a ®»»® o o

oooooooooooooooooooooooooooooo28 0 U 0 15. Urediniospores with variable numbers of germ pores (16) 16. Urediniospores with 4-5 equatorial or approximately equatorial,, rarely 4 equatorial and 1 at apex germ pores, 22-29 x 20-24 ji(, conspicuously echinulateg teliospores

pale yellow to golden brown, 17-22 x 15-20 }i9 wall 2-3

ja thick at sides® ©©©ogooooooo©29° J2° vicosensis 16. Urediniospores mostly with 3-4 germ pores (17)

17. Teliospores (18-)20-24 x (15-)18-22 p? wall 2»5-4 )x thick at sides; pedicel to 60-80 rarely to 100 long OOOOOOOOOOOOO0OOOO0OOOOOOOOOOO 30 ® Uo

17. Teliospores 18-26 x 13-18 p; wall lo5-2 p. thick at sidesj pedicel usually breaking near the hilum (18)

18. Teliospores minutely reticulate9 appearing minutely ver rucoses with meshes to about 1 p diam; urediniospores

with 3-4 equatorial or superequatorial germ pores0oooo«« • #oeoooo®ooooooooooooooi»ooooo#3io U© desrnod 11cola

var0 desmodiicola 22

Teliospores reticulate, with meshes to 1.5-2 diam; urediniospores with 3(^)» rarely 2, equatorial or slight ly superequatorial germ pores#32o U. desmodiicola vr • desmodii Figure 9. Teliospores of Uromyces appendiculatus vara appendiculatus with few to many verrucae (X 750)» Figure 10. Smooth teliospores of Uromyces appendiculatus var. brasiliensis (X 750). Figure 11. Verrucose-striate teliospores of Uromyces striatus (X 750). Figure 12. Reticulate teliospores with wall bilaminate and one urediniospore (center) Uromyces bauhiniae (X 750). 24

22 M

Figure 13. Teliospores (top) and urediniospores (bottom) of Uromyces belemensis*

Figure 14. Teliospores (top) and urediniospores (bottom) of Uromyces neurocarpio

Figure 150 Teliospores (top) and urediniospores (bottom) of Uromyces a-ppendiculatus var« appendiculatusB

Figure 16. Teliospores (top) and urediniospores (bottom) of Uromyces appendiculatus var. brasiliensis. Figure 17» Teliospores (top) and urediniospore (bottom) of Uromyces lathyrinus. Figure 18„ Teliospores (top) and urediniospore (bottom) of Uromyces viciae-fabae, Figure 19* Teliospores (top) and urediniospore (bottom) of Uromyces trifolii var. trifolii. 21 22

Figure 20® Teliospores (top), urediniospores (bottom) and one paraphysis (right) of Uromyces unionensis» Figure 21« Teliospores (top and bottom at left), urediniospores (bottom at right), and paraphyses (top at right) of Uromyces hedysari--paniculati9 Figure 22. Teliospores (top) and urediniospores (bottom) of Uromyces bradburyae. 27

C V 24 0 o i

Figure 23. Teliospores (top) and urediniospores (bottom) of Uromyces striatus. Figure Urediniospores of Uromyces crotalariae. Figure 25* Teliospores (top) and urediniospores (bottom) of Uromyces pisi. 28 . zzu ,

Figure 26. Teliospores (top) and urediniospores (bottom) of Uromyces anthyllidis. Figure 27. Teliospores (top and left) and aeciospores (right) of Uromyces orbicularis. Figure 28. Teliospores of Uromyces nerviphilus. Figure 29. Teliospores of Uromyces goyazensis. Figure 30® Teliospores of Uromyces floralis.

Figure 31. Teliospores of Uromyces viegasii. Figure 32. Teliospores (top) and urediniospores (bottom) of Uromyces bauhiniae. 30

/.i

y u" \

h 36 35

Figure 33„ Teliospores (right and left) and ure diniospores (middle) of Uromyces foveolatusa Figure 3*U Teliospores (top) and urediniospores (bottom) of Uromyces dietelianus var. dietelianuse

Figure 35« Teliospores (top) and urediniospores (bottom) of Uromyces dietelianus var» nervicola 0 Figure 360 Teliospores (top) and urediniospores (bottom) of Uromyces vicosensis. 31

22 M

;>v,

O V*

38 39

Figure 37. Teliospores (top) and urediniospores (bottom) of Uromyces castaneus.

Figure 38. Teliospores (top) and urediniospores (bottom) of Uromyces desmodiicola var. desmodiicola«

Figure 39« Teliospores (top) and urediniospores (bottom) of Uromyces desmodiicola var, desmodii. DESCRIPTION OP SPECIES OF UROMYCES ON LEGUMES IN BRAZIL

1, UROMYCES BELEMENSIS Albuq«> & Figue„, Pesq. Agropec. Bras. Ser. Agron. 6s 1^5-1^6o 1971 (Fig® 13)• Spermogonia and aecia unknown® Uredinia mostly hypophyllous, scattered or in small groups or densely aggregated, mostly along the veins? numerous, rounded, 0ol~0o5 nim diam, first covered by epidermis, later pulverulent, cinnamon- browm Urediniospores asymmetrical, triangular in one view, obovoid or broadly ellipsoid when turned 90°, (20-)2^-32(-33) x (18-)22-28 }i, wall 1 }i thick, cinnamon-brown,, conspicuously echinulate, germ pore 1 near the hilum, not distinct. Telia dark cinnamon-brown or reddish brown, otherwise as the ure dinia, or teliospores mostly mixed with urediniospores0 Teliospores subgloboid and rounded at apex, lateral views (20-)22-28(-29) x (24-)26-32(-33) pi, and top views (26-)29-

33(-35) x (2^-)26-30(-33) ji, dark cinnamon-brown, wall uni formly 1 p. thick, smooth, with 1 germ pore at apex; pedicel hyaline, some collapsing, 6-8 ja wide, to 30 long, less frequently to 60 ji long. Hosts and distribution in Brazils Ormosia nobilis Tul., Belem, Para (Albuquerque and Figueiredo, 1971)®

Types on Ormosia nobilis Tul0, Belem, Para, F.C. Albuquerque (IAN No. 1035? holotype; PUR F18258, isotype).

32 33

2. UROMYCES NEUROCARPI Diet,, Hedwigia 3^j 292, 1895 (Fig. 14-)« Uromyces rostratus Po Henne, Hedwigia 35s 22? o I8960 Uromyces insularis Arth0, Bullo Torrey Bot0 Club 33s 515» 1906o Spermogonia chiefly epiphyllous, a few among or on opposite side of aecia0 Aecia chiefly hypophyllous, in groupsi, sometimes in circles around spermogonia0 Aecia ure- dinoidp sometimes associated with veins,, spores similar to the urediniosporesc Uredinia chiefly hypophyllous„ scat- tered9 dark cinnamon-brown, 0ol-0o5 mm across, ruptured epi dermis noticeable, somewhat pulverulento Urediniospores triangular or obovoid in side view, broadly globoid in top view, (19-)23-25(-28) x (18-)20-23(-25) jx, wall cinnamon- brown, 1-1.5 ^ thick, echinulate except around the pores, with 2 slightly below the equator germ pores, without caps. Telia chiefly hypophyllous, scattered or in groups, some times associated with veins? compact, pulvinate, yellow- brown, later greyish.. Teliospores mostly fusiform or oblong-ellipsoid, (25-)29-38(-*H) x (10-)12-18 ji, wall smooth, pale yellowish, 0o5-l ^ thick at sides, 1.5-^ P- at apex; pedicel hyaline, slender, short, to length of spore, less frequently to 60 /a long0 Hosts and distribution in Brazil: Clitoria ca.ianifolia Benth., Bahia (Dietel, 1895), Para (H. Sydow and Sydow

1910), Rio de Janeiro (Hennings, I896; Sydow and Sydow, 34-

1907; Jackson, 1931)5 C0 rubiginosa Jusse, Minas Gerais

(Viegas, 19^5a)5 Clitoria spB, Para (He Sydow and Sydowp1910). Types on Clitoria ca.ianifolia Benth„ , Bahia , Lhotsky

(S, holotype)0

3. UROMYCES APPENDICULATUS (Perso) Ungo, Eifl. d» Bodens, p. 216, 1836a var„ APPENDICULATUS (Figo 9. 15). Uromyces phaseoli (Pers0) Winto, Hedwigia 19, p<> 37<> 1880e Urom.yces -phaseoli typica Artho , Man0 Rusts U0S0 and Canada pQ 2960 19620 Spermogonia epiphyllous, in groups0 Aecia hypophyllous, in groupse peridium cupulate, whitish, margin eroses aeciospores mostly polygonoid-globoid, minutely verrucoses mostly I8-36 x 16-2^ colorless® Uredinia amphigenous, cinnamon-brown9 on indistinct spots, scattered or in small groups, 0o2-l mm across, soon naked, surrounded by the rup tured epidermis? urediniospores mostly obovoid or broadly el

lipsoid (18~)2^-28(-32) x (18-)20-25(-28) Hp wall 1-2 ]is golden-brown, echinulate, with 2 equatorial or slightly above the equator germ pores, indistinct when seen through the wall, caps smooth or rarely with echinulations0 Telia amphigenous s, blackish brown, scattered or circinate around uredinia, ear ly exposed, pulverulent? teliospores globoid, obovoid or broadly ellipsoid, (24-)28-32(-35) x (19-)22-26(-29) ja, wall 2„5-3a5 p at sides, 5-8 jj. thick at apex, chestnut-brown, with a pale umbo and few to many verrucae scattered or sometimes in lines, rarely smooths pedicel colorless, collapsing, short to 50 p longe 35

Hosts and distribution in Brazils Phaseolus caracalla L0» Santa Catarina (Hennings8 1896)? Pa ovatus

Benth®d Rio Grande do Sul (Lindquist and Costa Neto* 1963)5

P. vulgaris L0„ Bahia (Carvalho and Vieira, 1972s Ponte,

197^)9 Ceara (YasconcelosB 1971) 5 Minas Gerais (Thurston,,

1940)e Parana (Fontoura and Nowacki 1967-70)p Pernambuco

(Krutman, 1968) 9 Sao Paulo (Hennings, 1902a; Viegas, 19^-5a);

P„ vulgaris L0 var0 mulatinhofl Sao Paulo (Viegasp 19^5a)i

Phaseolus sp09 Rio de Janeiro (Jacksonp 1931)» Sao Paulo

(Sydow and Sydow, 1907)5 Vigna sinensis Savi (Vo sinensis

Endl.), Para (Albuquerque9 1971)« This species probably oc curs in all regions where beans (P. vulgaris L.) are grown in Brazil®

4. UROMYCES APPENDICULATUS (Pers.) Ung. var. BRASILIENSIS var. nov, (Figo 10, 16). Urom.vces phaseoli longepedunculati Viegas9 Bragantia 5s 566. 1944. nom« nudum, Urediniis longe epidermide tecti, postea pulverulentis; sporae plerumque 20-29 x 18-26 p, poris germinationis 2, leviter superequatorialibus, distinctis0 Teliis conformibust sporae plerumque 29-35 x 19-26 levibus, raro paucis inconspicuis verrucae instructis0 Uredinia long covered by epidermis, later pulveru lent; urediniospores mostly 20-29 x 18-26 with 2 slightly above the equator germ pores, distinct. Telia as the uredi- 36 nia; teliospores mostly 29-35 x 20-26 yi, smooth, rarely with few inconspicuous verrucae. Hosts and distribution in Brazils Phaseolus longepedunculatus Mart0, Campinas, Sao Paulo (Viegas, 19^5b).

Types on Phaseolus longepedunculatus Mart0 , Campinas,

Sao Paulo, A0 Po Viegas (IACM No0 5071s holotype).

This variety differs from U0 appendiculatus var. appendiculatus because of the slightly superequatorial dis tinct germ pores of the urediniospores and the mostly smooth teliosporeso

5. UROMYCES LATHYRINUS Speg., Anal0 Soc. Cient. Argent, 12: 71. 1881 (Fig. 17). Uromyces clavatus Dieto, Hedwigia 36s 27. 1897. Uromyces chilensis Diet0 & Neger, Bot. Jahrb. 24s 15*K

Spermogonia hypophyllous or caulicolouss a few among the aecia0 Aecia densely aggregated, peridium whitish to yellowish, margin somewhat erose? aeciospores broadly glo boid to broadly ellipsoid, mostly with angulate edges? 15-26 x 13-20 ji, wall colorless, about 0.5-1 thick, minutely verrucose. Uredinia amphigenous or caulicolous, scattered, minute, ruptured epidermis conspicuous, pulverulent, yellow- brown to light cinnamon-browns urediniospores globoid, obovoid or broadly ellipsoid, 22-29 x 20-25 P-s wall golden- brown, 1.5-2 }a thick, minutely echinulate, with (5)6-8 scat tered germ pores, with hyaline, minutely echinulate caps. 37

Telia amphigenous or caulicolous, to 1 mm across, scattered, compact, cinnamon-brown becoming greyish? teliospores most ly fusiform or clavate, 30-^5 x 13-20 p, wall pale yellow- brown, about 0.5-1 P- thick at sides, smooth, to 8 at apex? pedicel colorless, thin-walled, to 10 ^ wide, breaking short

or to 60-70 ja longo Hosts and distribution in Brazils Lathyrus mage11an-

icus Lam0, Serra Geral (Dietel, 18975 Silveira, 1951)? Lathyrus multiceps Clos„, Vicia tenuifolia Roth„ (Silveira, 1951).

6. UROMYCES VIGIAE-FABAE (Pers.) Schroet,, Hedwigia l^s l6l. 1857 (Pig. 18). Uredo viciae-fabae Pers., Syn. Meth. Fung. p0 221. 1801. Uromyces fabae (Pers.) dBy., Ann0 Sci. Nat0 Bot„ ser. A-, 20: 80. 1863.

Spermogonia chiefly epiphyllous, in groups0 Aecia mostly hypophyllous, in groups, peridium cupulate, whitish, margin erosej aeciospores mostly broadly ellipsoid, 18-26 x

15 x 21 }i, wall colorless, lo0-1.5 ^ thick, densely and

finely verrucosea Uredinia amphigenous, yellow-brown or light cinnamon-brown, scattered, mostly rounded, 0,1-1 mm across, surrounded by the ruptured epidermis, pulverulent; urediniospores ellipsoid, obovoid or globoid, (20-)22-30 (-32) x (18-)19-2M-26) wall 1.5-2 ji, light golden-brown, echinulate, with (3)M5) equatorial or nearly equatorial, less frequently scattered germ pores, or with 3-^ equatorial 38

and 1 at apex germ pores with hyaline, echinulate caps. Telia backish brown, compact, otherwise as the urediniaj teliospores ovoid, obovoid, subgloboid or broadly ellipsoid,

(22-)2*4—35(~38) x (18-)20-26 ji, wall l05-2e>5 M thick at sides, 5-10 p at apex, chestnut-brown, smooth? pedicel yel lowish brown or only near the spore, mostly collapsing, to

40 ]x, less frequently to 65 jj. long0 Hosts and distribution in Brazils lens culinaris Medic., Rio Grande do Sul (Lindquist and Costa Neto, 1963).

Vicia faba L0 0 Minas Gerais (Thurston, 19*1-0), Parana (Fontou- ra and Nowacki 1967-70), Rio Grande do Sul (Lindquist and Costa Neto, 1963), Sao Paulo(Hennings, 1902bj Viegas, 1945a);

sativa L., Sao Paulo (Viegas, 1945a)j Vicia sp0, Sao Paulo (Viegas, 1945a)®

7. UROMYCES TRIFOLII (Hedw0 f. ex DCo) Wint0, Hedwigia 19; 37o I880e var6 TRIFOLII (Fig. 19). Puccinia trifolii Hedw0 f® ex DC®, Flore Fr. 2s 225. 1805« Uromyces trifolii (Hedw. f0) Lev®, Ann. Scio Nats III, 8s 371. 1847. Uromyces trifolii-repentis Liro, Acta Soca Fauna Flora Fenn. 29t no 6, p„ 15o 19060 Uromyces h.ybridi W0 H0 Davis, Mycologia 10s 2120 1924. Spermogonia chiefly epiphyllous „ in groups« Aecia amphigenous, in groups, peridium cupulate, margin erose, yellowish? aeciospores mostly globoid or with angulate edges

15-21(-24) x 13-17(~20) p.9 wall 1 ju thick, finely verrucose,

colorless8 Uredinia amphigenous, light cinnamon-brown, scattered, rounded, 0.1-0.5 mm across, surrounded by the 39

ruptured epidermis, pulverulentj urediniospores globoid to broadly ellipsoid, (21-)2^-28(~30) x (l8-)20-23 Us wall 1»5~

2 thick9 golden-brown,, echinulate, 2-3(^0 equatorial or slightly above the equator germ pores, with hyaline, echi-

nulate capso Telia chiefly hypophyllous9 chestnut-brown, otherwise as the uredinia; teliospores globoid to broadly el

lipsoid, (20-)22-28(-30) x (l6-)l8-2^ pv wall 1.5-2 ]i thick, cinnamon-brown, with a low hyaline papilla over the pore^ and few to many verrucae, scattered or frequently in lines, or smooths pedicel colorless, thin-walled, short, rarely to

30 n long. Hosts and distribution in Brazils on Trifolium spp., Parana (Pontoura and Nowacki, 19&7-70)» Rio Grande do Sul (Lindquist and Costa Neto, 1963)0

This variety differs from U0 trifolii (Hedw0 f ex

DC.) Winto varB fallens (Artho) Artho which often has 3 or k equatorial or approximately equatorial, or less frequently 6 or 7 somewhat scattered germ pores in the urediniospores and teliospores mostly smooth.

8. UROMYCES UNIONENSIS Viegas, Bragantia 5s 73-7^» 19^5 (Fig. 20).

Spermogonia and aecia unknown0 Uredinia chiefly hy pophyllous, light cinnamon-brown, round, minute, scattered

or confluent, first covered by epidermis, later pulverulent, with paraphyses inconspicuous, capitate or clavate, thin- 4o walled, collapsing, colorless, 10-20 }i wide apically? urediniospores globoid, obovoid or broadly ellipsoid, 18-28(-29) x (l6-)18-22(-24) )iB wall 1-1<>5 M "thick, pale yellow or pale yellow-brown, echinulate, with 4-5 equatorial or approxi mately equatorial germ pores, or less frequently with 3-4 equatorial or approximately equatorial and 1 near or at apex germ poreso Telia blackish brown, otherwise as the uredinia or teliospores sometimes mixed with urediniosporess teliospores ovoid or globoid-ellipsoid, 24-29(-32) x 22-28 ji, wall 3-4 p. thick at sides, 5-7 p thick at apex, dark chestnut-brown, with a paler, not clearly differentiated pa pilla at apex, varying from verrucose, with round, elongate and irregular warts more prominent toward the apex or these united to form a kind of labyrinth, to irregularly reticu late f pedicel colorless, mostly collapsing, to 40 p. long8 Hosts and distribution in Brazils Desmodium

(Meibomia) spe, Uniao, Alagoas (Viegas, 1945a)«

Type: on Desmodium (Meibomia) sp0, Uniao, Alagoas, Deslandes, No„ 436 (IACM No* 38I6, holotype? PUR F11571, isotype)• hi

9. UROMYCES HEDYSARI-PANICULATI (Schw.) Pari., Ellis, N. Amer0 Fungi 246. 1879 (Fig® 21)• Puccinia hedysari-paniculati Schw., Schr0 Nat. Ges0 Leipzig Is 7^o 1822. Aecidium desmodii P0 Henn0, Hedwigia 35s 239° 1896. Uredo desmodii-leiocarpi Po Henn0, Hedwigia 4ls 107. 1902. Uromyces desmodii-leiocarpi P0 Henn., Hedwigia ^8: 1. 1909 • Spermogonia epiphyllous or lackingo Aecia hypophyllous, in small groups9 peridium cupulate, whitish, with erose margin? aeciospores globoid or broadly ellipsoid, mostly with angulate edges 15-22(-2^-) x 13~18(-23) wall colorless, about 1 p thick, minutely verrucose0 Uredinia chiefly hypophyllous, scattered, minute, with ruptured epi dermis noticeable, pulverulent, light cinnamon-brown? with paraphyses colorless, mostly clavate, many 12-20 p. wide api- callys urediniospores broadly ellipsoid, obovoid or globoid, 19-2M-26) x 16-22 ji„ wall echinulate, 1-2 p thick, golden- brown, with (3)^-7(8) scattered germ pores, with hyaline, echinulate caps. Telia chiefly hypophyllous or epiphyllous, darkish brown, otherwise as the uredinia? teliospores ovoid, ellipsoid, broadly ellipsoid or subgloboid, 20-29(~32) x 15- 22 wall (2-)2.5-3 )i at sides, chocolate-brown or chestnut-brown, with a paler umbo, k-7 p. thick at apex, varying from verrucose, with irregular warts and ridges forming a kind of labyrinth, to irregularly or regularly reticulate? pedicel colorless, but mostly brownish yellow near hilum, some collapsing, to 40-60 jj. long. 42

Hosts and distribution in Brazils Desmodium leiocarpum G. Don„, Sao Paulo (Hennings, 1902a, 1909);

Desmodium sp0, Minas Gerais (Jackson, 1931)» Rio de Janeiro (Hennings, I896? Dietel, 1899; Jackson, 1931)t Ri° Grande do Sul (PUR P17739)9 Santa Catarina (Lindquist and Costa Neto,

1963)» Sao Paulo (Viegas, 1945a)s Do uncinatum DC®, Rio de Janeiro (Jackson, 1931)» Rio Grande do Sul (Lindquist and Costa Neto, 1963), Sao Paulo (Jackson, 1931)°

10. UROMYCES BRADBURYAE Jacks0 & Holw0p Mycologia 23s 352. 1931 (Fig. 22)o Spermogonia and aecia unknown,, Uredinia amphigenous or chiefly epiphyllous, scattered or gregarious, small, round or irregular, 0.1-0.5 mm across, light chestnut-brown, somewhat tardily naked, surrounded by the ruptured epider mis, pulverulent? urediniospores obovoid, ellipsoid or broadly globoid, 23-28(-31) x 18-24 wall pale brown to cinnamon-brown, 1.5-2 p. thick, often 2.5-3 P thick at apex and base, echinulate, 2 slightly above the equator germ pores, distinct, with smooth, hyaline caps. Telia dark chestnut-brown, otherwise as the uredinia5 teliospores most ly ovoid to pear-shaped or broadly ellipsoid, (27-)31-37 x

(l6—)20—26 p., wall 1,5-2 ji thick, light chestnut-brown, with a paler and clearly differentiated umbo, 5-7(-8) p. thick at apex, appearing smooth but actually verrucose, with numerous verrucae distributed evenly or irregularly throughout the H-3

surface of spore and umbo; pedicel mostly collapsing, color less, to 20 )i long, less frequently to 40 p long. Hosts and distribution in Brazils Centrosema (Bradbur.ya) -pubescens Benth. and Co virginianum Benth., Campos do Jordao, Sao Paulo (Jackson, 1931)® Types on Centrosema virginianum Benth. (Bradburya

virginiana (La) Kuntze), Campos do Jordao, Sao Paulo, Holway,

No. 1801 (PUR F3210, holotype)0

11. UROMYCES STRIATUS Schroet0, Abha Schles„ Ges. Vaterl. Cult. Nat. Abth0 1869-72; 11. 1870 (Fig. 11, 23). Uromyces medicaginis Pass0, Thuem0 Herb. Mycol. Oecon0 156. I8960 Spermogonia hypophyllous, numerous, scattered. Aecia hypophyllous among spermogonia, peridium cupulate, white; aeciospores polygonoid or ellipsoid mostly 18-23 p diam, wall

1 ]x thick, colorless, minutely verrucose0 Uredinia chiefly hypophyllous, cinnamon-brown, scattered or in small groups, minute, 0.1-0.5 mm across, surrounded by the ruptured epi dermis, pulverulent; urediniospores globoid, subgloboid or broadly ellipsoid, 18-2M-26) x (16-)18-22 ja, wall (1-)1.5- 2(-2•5) M "thick, golden-brown or light cinnamon-brown, echinulate, with (2)3-4(5) equatorial, less frequently slightly above the equator germ pores, with hyaline, echinulate caps. Telia about dark cinnamon-brown, otherwise as the uredinia; teliospores ellipsoid, obovoid or globoid to subgloboid, 18-

26(-29) x 14-21 p., wall 1-2 }i thick, cinnamon-brown or light chestnut-brown, 2.5-4 p. thick at apex with a paler papilla verrucose-striate, with ridges or elongated warts disposed longitudinally, sometimes united, on the surface of spore; pedicel colorless,, thin-walled, to 30 long, rarely to 40 p. long8 Hosts and distribution in Brazils Medieago sativa

L., Belo Horizonte, Minas Gerais (Thurston, 1940), Campina

Sao Paulo (Viegas, 1945a)9 Parana (Pontoura and Nowacki,

1967-70), Piracicaba, Sao Paulo (Viegas, 1945a), Sao Paulo

(Sydow and Sydow, 1907)» Southern Brazil (Jackson, 1931)•

12. UROMYCES CROTALARIAE (Arth.) J. Baxter, Mycologia 54': ^37-^39o 1962 (Fig. 24). Uropyxis crotalariae Arth. , Amer. J. Bot® 5? 4-29® 1918 Haplop.yxis crotalariae (Arth.) Syd.» Ann. Mycolo 17: 105. 1919. Spermogonia and aecia unknown,, Uredinia mostly hypophyllous, light cinnamon-brown, ruptured epidermis noticeable, pulverulent; urediniospores broadly ellipsoid, obovoid or globoid, 22-30 x 20-24 ji, wall yellowish to golden-brown, l«5-2(-2.5) M thick, echinulate, with 6-8, mostly 8, scattered germ pores, with hyaline, echinulate caps. Baxter (1962, p» 437) sayss Telia hypophyllous, black or chocolate-brown, pulverulent; teliospores oblate-spheroid, bluntly conoid or ovoid, assuming a globoid shape when turned to present the apical pore or point of ped icel attachment in surface view, oblate-spheroid teliospores 27-32 /a broad x 20-27 p. high, conoid or ovoid teliospores 20-27 p broad x 24-30 p. high; 45

wall bilaminate, inner layer 2-2.5 thinner and papillate at the point of pedicel attachment, chest nut-"brown , pore apical, inconspicuous; outer layer swelling in water to a thickness of 3-10 p, sharply indented at the point of pedicel attachment, hyaline, yellow or pale cinnamon-brown, verrucose- echinulate, with short conical tubercles; pedicel thin-walled, hyaline, usually breaking away at the point of attachment to the spore, occasionally per sistent, 12-22 ji longo Hosts and distribution in Brazils Crotalaria vitellina Ker0, Rio de Janeiro (Jackson, 1931)«

13. URCMVIYCES PIS I (DCS) Otth, Mitt. Naturfo Ges„ Bern, I863: 87o 1963 (Figo 25). Puccinia pisi DCe, Flore Fr0 2s 224® 1805® Uromyces pisi (Pers0) Wint., Pilze Deutschl, 163. 1884. Spermogonia hypophyllous, many, scattered among the aecia. Aecia hypophyllous, systemic, scattered all over the leaf surface, peridium cupulate, margin revolute, deeply la- ciniate, whitish; aeciospores globoid to ellipsoid, mostly with angulate edges, 15-24(-26) x 13-l8(-20) ja, wall color less, about 1 p thick, minutely verrucose. Uredinia chiefly hypophyllous, to 1mm diam, scattered, surrounded by the ruptured epidermis, light cinnamon-brown, pulverulent; urediniospores mostly globoid to ellipsoid, 19-26 x 17-24 p, wall golden-brown, 1.5-2 p thick, minutely echinulate, with 3-6 scattered germ pores, with hyaline, echinulate caps® Telia amphigenous, dark brown, surrounded by the ruptured epidermis, scattered or in groups, coalescent, to 2-3 mm diam, pulverulent} teliospores mostly globoid to ellipsoid,

20-30 x 15-22 p, wall light chestnut-brown with a low, paler papilla at apex, 105-2.5 ^ thick at sides, densely and minutely verrucose, with warts mostly round, evenly distrib uted on surface of spore and papilla, about 1 p. apart; pedicel colorless, thin-walled, some collapsing, breaking near the hilum or to about 50 p long0

Hosts and distribution in Brazil; Pisum sativum L0, Silveira, 1951? Rio Grande do Sul (Luz, 1970)»

1&. UROJYIYCES ANTHYLLIDIS (Grev.) Schroet., Hedwigia 1^: 162. 1875 (Pig. 26). Uredo anthyllidis Grev. in Smith, Engl. Fl» 5(2) s 383» 1836. Urom.yces renovatus Syd., Mo nog. Ured. 2s 113® 1910. Spermogonia and aecia resembling those of Uromyces pisi. Uredinia amphigenous, scattered, minute, roundish, ruptured epidermis noticeable, pulverulent, cinnamon-brown; urediniospores mostly globoid or subgloboid, 16-25 p diam, wall golden-brown, (1o5-)2-3(-3®5) p thick, finely echinulate, with ^-8P mostly 6-8, scattered germ pores, with hyaline, echinulate caps0 Telia about chocolate-brown, otherwise as the uredinia; teliospores globoid or broadly ellipsoid, 17-22 x 15-20 p., wall about dark, brown, 47

(l,5-)2-2.5(-3) P thick, verrucose, with well developed and sparsely disposed verrucae, with no or a low, paler papilla at apexj pedicel colorless, thin-walled, breaking near the hilum. Hosts and distribution in Brazils Lupinus albus L«, Porto Alegre, Rio Grande do Sul (Lindquist and Costa Neto,

1963)5 Lupinus sp0, Campinas9 Sao Paulo (Viegas, 1945a)0

15, UROMYCES ORBICULARIS Dieto, Hedwigia 36s 28„ 1897 (Fig. 27). Spermogonia and uredinia unknown,, Aecia hypophyllous, in groups, whitish or pale yellowish* densely aggre gated, forming very distinct spots mostly round, 1-4 mm across, peridium cupulate, with margin upright, somewhat erose; aeciospores globoid or broadly ellipsoid mostly with angulate edges, 15-22 x 13-17 }x, wall colorless, about 1 ji thick, minutely verrucoseo Telia epiphyllous„ opposite the aecia, frequently in circles, confluent, chocolate-brown, minute, with ruptured epidermis noticeable, pulverulent; teliospores ellipsoid, obovoid, globoid or subgloboid, 22-31

(-33) x 18-2M-26) ji, wall 2-3 ;i thick at sides, chestnut-

brown or dark golden-brown, with a paler umbo 3-7 p. high, varying from verrucose, with irregular warts sometimes vari ously united or arranged to form a kind of labyrinth, to

irregularly reticulates pedicel colorless, thin-walled, many

collapsing, to 35 }i long, rarely to 50 p long8 48

Hosts and distribution in Brazils Desmodium sp,,Belo Horizonte, Minas Gerais (Thurston, 1940)? Linhares, Espirito

Santo, collected by Dr6 Re B0 Hine from the Department of Plant Pathology of The University of Arizona? Serra Geral (Dietel, 1897). Types on Desmodium sp«,Serra Geral, E« Ule, No. 1647

(B, holotype)0

16„ UROMYCES NERVIPHILUS (Groga) Hotson, Publ„ Puget Sound Biol0 Sta0 Univo Washe 4s 368, 1925 (Pig® 28)» Uromyces flectens Laghe, Svensk Bot« Tidskr» 3' 3^o 1909o Spermogonia chiefly hypophyllous or mostly on peti oles and veins, among the aecia® Aecia in groups, cupulate, margin somewhat erose, yellowish? aeciospores globoid to broadly ellipsoid, mostly with angulate edges, mostly 18-24 x 13-20 p, wall colorless, about 1 ji thick, minutely ver rucoses, Uredinia unknown0 Telia chiefly hypophyllous, chocolate-brown, minute, scattered or in groups, coalescent, frequently elongated, causing hypertrophy on the veins and petioles, long covered by the epidermis, then pulverulent; teliospores ellipsoid, obovoid, globoid or ovoid, (l8-)20- 28(-30) x 15-22(-25) JA, wall l«>5-2 JI thick, cinnamon-brown or light chestnut-brown, with a low, round papilla over the pore, paler than spore, minutely and inconspicuously verrucose, appearing smooth? pedicel colorless, thin-walled, breaking near the hilum. 49

Hosts and distribution in Brazils Trifolium repens L., Porto Alegre, Rio Grande do Sul (Lindquist and Costa Neto, 1963)«

17. UROMYCES GOYAZENSIS P. Henn., Hedwigia 34: 89o 1895 (Fig, 29). Spermogonia, aecia and uredinia unknown,, Telia on branches of inflorescence and on fruits, minutes, round, or in

groups, confluent, 0o2-0o5 cm to 2 cm across, cushion-shaped, ruptured epidermis conspicuous, pulverulent, dark cinnamon- brown? teliospores ellipsoid, obovoid or globoid, (l8-)20-30 (-32) x (15-)18-22(-24) y1, wall pale brown, 2-3 y thick, slightly thickened at apex with a paler, low papilla not

abruptly differentiated to 2 p. high, slightly and minutely reticulate, appearing smooth, with meshes about 0*5-1 M diam; pedicel colorless, thin-walled, breaking near the hilum, or

mostly to 80-100 ji long. Hosts and distribution in Brazil! Bauhinia sp«, Es- pirito Santo do Pinhal, Sao Paulo (Viegas 1945a); Goias

(Hermings, I895). Type; on Bauhinia sp., Goias, E. Ule, No. 190? (B.

holotype). 50

18. UROMYCES FLORALIS Vest*, Ark. Bote 4(15): 23. 1905 (Fig. 30). Uromyces verus Jacks. & Holw„, Mycologia 32s 3^° 1931* Aecia and uredinia unknown,, Spermogonia epiphyllous, few, punctiform, among or surrounded by telia, or lacking®

Telia amphigenous, on flowers, caulicolous, or on leaves9 cinnamon-brown, scattered or in groups, coalescent, 1-10 mm to 15 mm long, long covered by the epidermis, later pulver ulent, ruptured epidermis conspicuous" teliospores obovoid, globoid or ellipsoid, (17-)20-2M-26) x (15-)17-20 ji, wall golden or light cinnamon-brown, (1.5-)2-2.5(-3) M thick, mi nutely and uniformly reticulate with meshes about 1-1.5 p diam, slightly thickened at apex with a paler, low papilla to 2 }x high; pedicel colorless, thin-walled usually breaking near the hilum, rarely to about 80 p long. Hosts and distribution in Brazils Bauhinia cuyabensis Steud., Mato Grosso (Vestergren, 1905); B. hiemalis Malme, Mato Grosso (Yestergren, 1905). B. holophylla Steudo, Minas Gerais (Vestergren, 1905)5 B« rufa Steud., Belo Horizonte, Minas Gerais (Jackson, 1931), Sao Simao, Sao Paulo, IACM No.

3317 (Viegas, 19^5a)» Bauhinia sp0, Campinas, Sao Paulo

(Sydow and Sydow, 1907)0 Franca, Sao Paulo, (Viegas, 19^5a). Type: on Bauhinia hiemalis Malme, Cuiaba, Mato Grosso, Malme, VV» 1902 (S, lectotype). 51

19. UROMYCES ANTHEMOPHILUS Vest., Ark. Bote Ml5)s 2^0 1905«

Spermogonia, aecia and uredinia unknown0 Telia de veloped on flowers„ without spots, 3-10 mm long, pulverulent, dark brown? teliospores globoids subgloboid or ovoidt apex rounded» not or barely papillate, distinctly and regularly reticulate, pale brown, 18-25 x 18-22 wall 3-3®5 p thick at sides? pedicel slender9 colorless,, deciduousQ Hosts and distribution in Brazils Bauhinia longifolia

D. Dietr0, Cuiaba, Ma to Grosso (Vestergren, I905)o

Types on Bauhinia longifolia Do Dietr0, Cuiaba, Mato Grosso, Malme (S„ holotype). This species, according to Vestergren, is closely related to Us floralis Vest8 but differs from it because of the larger meshes of the reticulate wall and absence of a papilla at apex in the teliospores„

20. UROMYCES VIEGASII sp8 nov. (Fig. 31), Aeciis et urediniis ignotis. Spermogoniis epiphylis, paucis. Teliis cinnamomeo-brunneis, amphigenis9 minutis, sparsis vel gregariis, aliquando circulariter dispositis, inter vel circiter spermogoniis, longe epidermide tectis, postea pulverulentis, rupta epidermis conspicua, maculis brunneis, 1-5 mm diam, flavido halo cinctisj teliosporae globoideae vel ellipsoideae„ (15-)18-22(-2*0 x (13-)16-18(-20) membrana aurea ad aureo-brunnea, 1.5-2o5 M crassa, reticu lata, reticulationes variabilibus usque ad 2-2®5 p diam; 52 apice reticulo prominenti et diluto, distincta papilla usque ad 2.5(-3) }A altag pedicello, hyalino, fragili, brevi„

Aecia and uredinia unknown0 Spermogonia epiphyllous, few. Telia cinnamon-brown, amphigenous, minute„ scattered or in groups, sometimes in circles around the spermogonia, long covered by the epidermist later pulverulent, ruptured epi dermis conspicuous5 with brown spots, 1-5 mm diam, sur rounded by a yellow halog teliospores globoid or ellipsoid, (15-)18-22(-24) x (l3~)l6~18(-20) wall golden to golden- brown, 1o5-2o5 H thick at sides, reticulate, with meshes of variable size and shape to about 2~2<>5 M diami at the apex, the reticula are more salient and paler, forming a low and distinct papilla to 2o5(-3) J1 highs pedicel colorless, thin- walled, breaking near the hilum to 10-15 p longe Hosts and distribution in Brazils Bauhinia forficata L., Sao Jose do Rio Pardo, Sao Paulo (Viegas 19^53)0 Types on Bauhinia forficata L«, Sao Jose do Rio

Pardo, Sao Paulo, A« So Costa <, (IACM No9 1361, holotype)0 This microcyclic species is closely related to U# floralis and U» anthemophilus but differs from the former especially because of the larger meshes and salient reticu la of the teliospore wallsB It differs from U0 anthemophilus mainly because of the thinner wall of the teliospores and presence of a papilla® Also this species forms distinct brown spots surrounded by a yellow halo# Such spots are not produced by any other species of Uromyces found on Bauhinia. 53

21. UROMYCES BAUHINIAE P. Henn„, Hedwigia 34s 90, 1895 (Fig. 32)o Uromyces praetextus Vest0, Ark0 Bot. 4(15)s 19® 1905s Uromyces guatemalensis Vest0, Ark0 Bot0 4(15)s 20® 1905® Spermogonia and aecia unknown,, Telia blackish brown, chiefly hypophyllous, minute, scattered or sometimes in

groups,, confluent, O05-le5 mm across, pulverulent, ruptured epidermis inconspicuous« Uredinia amphigenous, minute, . scattered or aggregated, cinnamon-brown or urediniospores mixed with teliospores; urediniospores mostly globoid or el lipsoid (,l8-)22-26 x 17-22(-26) JI, wall golden-brown or dark golden-brownj, echinulate lo5-3 P- thick, 3-5(6) equatorial, nearly equatorial or somewhat scattered germ pores, with no or small, hyaline, echinulate caps; teliospores globoid, el lipsoid, obovoid or ovoid, (20-)22-28(-30) x (l8-)22-25(-27) ]a, wall conspicuously bilaminate, distinctly and uniformly reticulate with meshes to about 2-2«,5 V- diam; inner wall chestnut-brown, 1.5-3 ^ thick; outer wall distinct about yellowish brown, (l-)l05-2o5(-3) H thick at sides, thinner at base, to 6 JI thick at or near the apex but without an umbo; pedicel colorless, thin-walled, to 40-50 long, but usually breaking near the hilurcu Hosts and distribution in Brazils Bauhinia bongardii Steudo* Bo cuyabensis Steudo, B. hiemalis Malme, Cuiaba, Mato Grosso (Vestergren, 1905)5 Bauhinia sp.» Areia, Paraiba IACM No. 3829 (Viegas, 1945a), Mogi das Cruzes, Sao Paulo, Reliq. Holw. No. 305 = Holw. 1998 (Jackson, 1931). Niteroi, 54

Rio de Janeiro,, Reliq. Holw. No. 275 = Holw. No* 1120

(Jackson, 1931)t Paranaiba, Minas Gerais (Hennings, 1895)*

Types on Bauhinia sp0, Paranaiba, Minas Gerais, E.

Ule, No. 19068 (B, holotype)0

22. UROMYCES FOVEOLATUS Juel, Bih. Till K. Svenska Vet.- Akad. Handlo Bb0 23* Afd. II, No. 10, p. 16. 189? (Fig. 33)o

Spermogonia and aecia unknown0 Uredinia chiefly hypophyllousB 0.1-0.5 mm diam, scattered, pulverulent, cinnamon-brown; urediniospores globoid to broadly ellipsoid or obovoid, 22-26(-28) x 20-25 wall 2-205 M thick, golden-brown to light cinnamon-brown, echinulate, with 4(5)» rare 6 or 3 equatorial germ pores, with hyaline, echinulate caps. Telia blackish brown, otherwise as the uredinia; teliospores globoid to broadly globoid or ellipsoid, 22-29 x 20-24 wall dark chestnut-brown, bilaminate, distinctly reticulate with meshes to 2-2.5 P- diam, inner wall 1-1.5 H thick, outer wall not distinct, paler, 1.5-2.5 p thick, thinner toward the base, and thicker at apex, usually forming a paler and distinct umbo, also reticulate, to 3-5 high; pedicel colorless, thin-walled, to about 20-25 M long, basally rugose. Hosts and distribution in Brazils Bauhinia hirsuta Vog,, Cuiaba, Mato Grosso (Vestergren, 1905); B. mirandina Pittier, Minas Gerais, Muller, No. 1077 (Thurston, 1940); 55

Bauhinia sp., Minas Gerais, Muller, No. 296 (Thurston, 19^0),

Petropolis, Rio de Janeiro, Reliq. Holw0 No. 284 = Holw0 No. 1270 (S). Type: on Bauhinia hirsuta Vog., Cuiaba, Mato Grosso, Malme (S, isotype).

23. UROMYCES REGIUS Vest., Ark® Bot„ 4(15) s 31» 1905o Spermogonia and aecia unknown. Telia amphigenous, without spots, scattered, punctiform, about chestnut-brown, pulverulent! urediniospores mixed with teliospores, globoid, subgloboid or ovoid, 16-23 x 16-20 wall yellow-brown, 2*0

}x thick, conspicuously echinulate, with 4 germ pores? telio spores globoid, subgloboid or ovoid, 20-27 x 17-20 ji, wall pale chestnut-brown, 2o5 p thick, reticulate, meshes variable in shape and size, with a yellow-brown corona to 6a0 ja high at apex; pedicel colorless, slender, same length of spore0 Hosts and distribution in Brazils Bauhinia candicans Benth., Rio Grande do Sul (Vestergren, 1905)» Types on Bauhinia candicans Benth., Rio Grande do Sul, Lindman (S, holotype). This species according to Vestergren is closely re lated to Uo dietelianus and U. pannosus but differ from those especially because of the smaller, conspicuously echinulate, urediniospores. 56

24. UROMYCES PERAFFINIS Diet,, Ann. Mycol. 20: 294. 1922. Spermogonia and aecia unknown,, Sori mostly epiphyllous, scattered, to 008 mm across, naked, pulverulent9 chestnut-brown. Urediniospores mixed with teliospores, el lipsoid or obovoid, 24-28 x 18-21 ja, pale brown,, sparsely echinulate, with 3 germ pores0 Teliospores obovoid- ellipsoid, rarely oblong, 25-33 x 18-23 with a paler, broad, obtuse„ tuberculate,, papilla to 6 p. high at apex, yellow-brown, wall indistinctly and minutely reticulate; ped icel slender, fragile. Hosts and distribution in Brazil: Bauhinia sp., Rio de Janeiro (Dietel, 1922). Type: on Bauhinia sp„ Rio de Janeiro, specimen not found, Eo Ule, 1895®

25. UROMYCES PERLEBIAE Vest. Ark. 4(15)s 26, 1905o Spermogonia and aecia unknown. Uredinia amphigenous, without spots, scattered, minute, pulverulent, cinnamon- brown; urediniospores globoid, subgloboid or ellipsoidD 22-26 x 21-24 n, wall pale yellow, 1,5-2 ja thick, loosely echinulate, with 4, rarely 5, germ pores. Telia about chestnut- brown, otherwise as the uredinia; teliospores globoid, sub globoid or broadly ellipsoid, 22-26 x 20-23 p, wall about chestnut-brown, 3-3.5 p. thick, apex rounded and yellow-brown corona, 2.5-3 rarely 4 p, high, distinctly reticulate; 57 pedicel colorless, subpersistent, moderately thick, same length as spore or a little longer0 Host and distribution in Brazils Bauhinia pentandra

Vog., Cuiaba, Mato Grosso (Vestergren, 1905)o

Type ; on Bauhinia pentandra Vog0„ Cuiaba, Ma to

Grosso, Malme0 (S, holotype)0

26. UROMYCES DIETELIANUS Paz0, Hedwigia 30s 199° 1891. var. DIETELIANUS (Figo 3^)° Urom.yces pannosus Yest0 Arko Bot0 ^(15) '• 3°° 1905s Spermogonia epiphyllous, few, in groups„ Aecia uredinoid, hypophyllous, opposite spermogonia, light cinnamon- brown, in groups, confluent, spores similar to urediniospores. Uredinia amphigenous or chiefly hypophyllous, scat tered, minute or in small groups, coalescent, 0ol-0o5 mm diam, pulverulent, yellow-brown to cinnamon-brown; urediniospores obovoid, ellipsoid or broadly globoid (18-)20-27(-29) x (15-)17-22 ji, wall golden-brown, 1-2 ja thick, minutely echinulate, with (3)^-5» rarely 6 or 7? equatorial or nearly equatorial germ pores, with small, hyaline, echinulate caps8 Telia about chestnut-brown or darkish brown, otherwise as the uredinia? teliospores mostly obovoid, ellipsoid or broadly globoid 22-28(-31) x (15-)17-21(-2*0 wall pale chestnut-brown to chestnut-brown, l05-2o5 M thick at sides, reticulate with meshes variable in size and shape to 2-3(-^) p. diam, or sometimes forming a kind of labyrinth, with a paler, distinct and broad papilla or corona to 3-5 ja high, 58

often irregularly undulate or serrate at apex; pedicel color less, thin-walled, breaking near the hilum or to about 25 M long, less frequently to 35 p long, basally rugose® Hosts and distribution in Brazils Bauhinia candicans

Bentho, Gramado, Rio Grande do Sul, LPS No0 3019^ (Lindquist and Costa Neto, 1963) j Bauhinia forficata Link,, Sao Joao, Sao

Paulo, HolWo No0 l652=Reliq0 Holws No0 296 (Jackson, 1931)?

Bauhinia pruinosa Vog0, Passo do Cerro Chato, Rio Grande do Sul, LPS No„ 30193 (Lindquist and Costa Neto, 1963)? Bauhinia

sp.( Espirito Santo do Pinhal, Sao Paulo, IACM No0 3^59

(Viegas, 19^5a)» Mandaqui, Sao Paulo, Holw„ No0 I883 (Jackson, 1931)» Rio de Janeiro, Reliq. Holw. No® 267=Holw. No. 1025 (Jackson, 1931)» Tijuca, Rio de Janeiro, Ule No,

2387 (B), Noack No0 512 (S), Tubarao, Santa Catarina, No, 12*4-0 (B), Nos 3719 (S). Types on Bauhinia sp»,Tubarao, Santa Catarina, E. Ule (B, holotype; S, probably isotypes Rab.-Wint. f. eur. No.

3719).

27. UROMYCES DIETELIANUS Paz» var. NERVICOLA var. nov. (Fig. 35)• Spermogoniis epiphylli.s, paucis, gregariis. Aeciis uredinoideis„ epiphyllis, gregariis, inter vel circiter spermogoniis; aeciosporae urediniosporae similibus. Ure- diniis amphigenis vel plerumque hypophyllis, minutis vel ple-

rumque gregariis, confluentibus, plerumque in folia nervis 59 dispositis vel sparsis, pulverulentis, cinnamomeo-brunneis vel urediniosporae paucis, teliosporae intermixtisj urediniosporae plerumque ellipsoideae vel obovoideae, 21-27 x (13)15- 19 p., membrana aureo-brunnea, lo5-2 p. crassa, subtiliter echinulata ad echinulata, poris germinationis 3-^8 rare 2, 5 or 6, parv'is, hyalinis, echinulatis umbonibus praeditiso Teliis castaneo-brunneis, aliter urediniis conformibusj teliosporae plerunque ellipsoideae, obovoideae vel late globoideae, 22-31(-33) x 18-2^ ]xe membrana pallide cinnamomeo- brunnea ad cinnamomeo-brunnea, 2-3 p crassa, papilla lata, distincta, pallida praedita usque ad 3-^ P altaj pedicello plerumque brevi hyalino, fragili usque ad 20-30 p, rare usque ad kO p. longo, non basaliter rugoso#

Spermogonia epiphyllous, few, in groups,, Aecia uredinoid, epiphyllous, in groups, among or surrounding the spermogonia; aeciospores similar to urediniospores0 Ure- dinia amphigenous or chiefly epiphyllous, minute or mostly in groups, coalescent, mostly arranged linearly along or on the veins or scattered,, pulverulent, cinnamon-brown, or urediniospores not numerous, mixed with teliosporesj urediniospores mostly ellipsoid or obovoid, 21-27 x (13-)15-19 p, wall golden-brown, 1.5-2 p thick, minutely echinulate to echinulate, with 3-^, rarely 2, 5 or 6, equatorial germ pores, with small, hyaline, echinulate capse Telia chestnut- brown, otherwise as the uredinia; teliospores mostly ellip 6o

soid, obovoid or broadly globoid, 22-31(-33) x 18-24 p, wall light chestnut-brown to chestnut-brown, 2-3 p thick at sides, with a paler, distinct and broad papilla to 3-4 ja high at apex, reticulate with meshes variable in size and shape to 3.0 n diam? the apex of the papilla usually appearing almost smooth in silhouette; pedicel colorless, thin-walled, many

collapsing, breaking near the hilum or to 20-30 jj. long, less

frequently to 40 p. long, not basally rugose0 Hosts and distribution in Brazils Bauhinia forficata

Link, Campinas, Sao Paulo, IACM No* 3094 and IACM No0 4222

(Viegas, 1945a); B0 forficata Link var. latifolia Bentho,

Prata, Sao Paulo, Reliq. Holw. No„ 299=Holw0 No0 1?02 (Jackson, 1931); Bauhinia sp., Petropolis, Rio de Janeiro,

Reliq. Holw„ No„ 280=Holwo No„ 1225 (Jackson, 1931)f Quinta da Boa Vista, Rio de Janeiro, Hohnel No. 131 (S), Rio de Ja neiro, Ule Noo 2156 (B)o Types on Bauhinia forficata Link, Joaquim Egidio, Campinas, Sao Paulo, A. P. Viegas and C. P. Viegas, (IACM No.

3094, holotype).

This variety differs from U. dietelianus var0 dietelianus mainly because of distribution of uredinia and telia mostly along on or the veins; number of germ pores; teliospores wall slight thicker and pedicel not basally ru gose. 61

28. UROMYCES HEMMENDORFFII Vest., Ark. Bot. 4(15)s 32. 1905. Spermogonia and aecia unknow. Telia epiphyllous, scattered, punctiform, pulverulent, chestnut-brown; urediniospores (only a few seen), ellipsoid or ovoid, 23-28 x 1.5-17 p., wall yellow-brown, 2 thick, sparsely echinulate, with 4 germ pores. Teliospores globoid, subgloboid or broadly el lipsoid, apex rounded, 18-22 x 16-18 wall yellow-brown,

2.5 thick, minutely reticulate, with a papilla to 205 M high or absent; pedicel short, deciduous. Hosts and distribution in Brazils Bauhinia forficata Link, Sao Paulo (Vestergren, 1905)• Types on Bauhinia forficata Link, Sao Paulo, Hemmendorff (S, holotype).

29. UROMYCES VICOSENSIS sp0 nov. (Fig. 36). Spermogoniis epiphyllis, paucis vel numerosis, gregariis, maculis usque ad 5 mm diam. Aeciis uredinoideis, gregariis, confluentibus, cinnamomeo-brunneis, spermogoniis op- positis; aeciosporae urediniosporae similibus. Urediniis plerumque hypophyllis, minutis, sparsis vel parvis gregariis, confluentibus, cinnamomeo-brunneis, pulver'ulentis; uredinio sporae obovoideae, late globoideae vel late ellipsoideae, 22- 29 x 20-24 ja, membrana flavido-brunnea ad cinnamomeo-brunnea, 2-2.5(-3) M crassa, conspicue echinulata, poris germinationis 4-5 equatorialibus vel quasi equatorialibus, sine vel parvis, hyalinis, echinulatis umbonibus praeditis. Teliis epiphyl- 62 lis, minutis, sparsis vel plerumque gragariis, confluentibus, rupta epidermis conspicua, inter vel circiter spermogoniis, cinnamomeo-brunneis ad rubello-brunneis; teliosporae ple rumque globoideae vel late ellipsoideae, 17-22 x 15-20 ix, membrana diluto flavida ad aureo-brunnea, 2-3 M crassa, re ticulata, reticulum prominens, reticulationes variabilibus usque ad 2 p. diam, apice diluto umbone usque ad 2oO-2„5 M alto; pedicello, hyalino, fragili, brevio

Spermogonia epiphyllous, few to many, in groups, on spots to 5 mm diarru Aecia uredinoid, in groups, coalescent, cinnamon-brown, opposite the spermogonia0 Uredinia chiefly hypophyllous, minute, scattered, or in small groups, coales cent, cinnamon-brown, pulverulent; urediniospores obovoid, broadly globoid or broadly ellipsoid, 22-29 x 20-2^ p.„ wall yellow-brown to cinnamon-brown, 2-205(-3) M thick, conspicu ously echinulate, b-5 equatorial or approximately equatorial, rarely b equatorial and 1 at apex germ pores, without or with small, hyaline, echinulate caps0 Telia epiphyllous, minute, scattered or mostly in groups, coalescent, ruptured epidermis conspicuous, among or surrounding the spermogonia, cinnamon- brown to reddish brown; teliospores mostly globoid or broadly ellipsoid, 17-22 x 15-20 p.9 wall pale yellow to golden-brown, 2-3 p thick at sides, reticulate, with salient reticula and meshes variable to about 2 p. diam, with a paler umbo to 2-2.5 63 ji high at apex; pedicel colorless, thin-walled, breaking near the hilum. Hosts and distribution in Brazils Bauhinia sp., Vi- cosa, Minas Gerais (Thurston, 19^0).

Type; on Bauhinia sp0, Vicosa, Minas Gerais, A. S, Muller, No. 7^1 (CUP, holotype). The present species does not correspond to any of the rust species described on Bauhinia« The combination of the larger, cinnamon-brown urediniospores which are con spicuously echinulate and have 4—5 equatorial germ pores and the smaller, pale yellow to golden-brown teliospores makes this species distinct.

30. UROMYCES CASTANEUS Syd., Monog. Ured. 2: 9^. 1910 (Fig. 37). Spermogonia and aecia unknown. Uredinia chiefly hypophyllous, throughout the entire leaf surface, more or less scattered or almost uniformly distributed, 0.1-0.5 mm, ruptured epidermis inconspicuous on the lower leaf surface but noticeable or long covered on the upper leaf surface, light cinnamon-brown, pulverulent; urediniospores broadly ellipsoid to subgloboid or globoid, 15-20 x 14—18(-20) p., wall l-l,5(-2) thick, golden-yellow to golden-brown, echi nulate, with 3-4- slightly above the equator germ pores, with hyaline, echinulate caps. Telia dark brown, otherwise as the uredinia or teliospores mixed with urediniospores; 64 teliospores globoid to subgloboid or broadly ellipsoid, (18-) 20-24 x (15-)18-22 p, wall chestnut-brown, minutely and uni formly reticulate with meshes about 1 p. diam, 2<>5-^ p thick at sides, slightly thickened at apex to 5 J-U with a paler, not abruptly differentiated umbos pedicel colorless, thin- walled, many collapsing, to 60-80 long, rarely to 100 p. long. Hosts and distribution in Brazils Desmodium incanum DC., Paraiba (Viegas, 1945a), Pernambuco (Viegas, 1945a),

Rio de Janeiro (Jackson, 1931)J Desmodium sp0, Rio de Janei ro (Jackson, 1931)® Types on Desmodium incanum DC., Rio de Janeiro, E. Ule, No. 664(PUR F2932, isotype).

31. UR0MYCES DESM0DIIC0LA Jorst., Ark0 Bot. 4(5): 75-76. 1959. var, DESM0DIIC0LA (Pig. 38). Spermogonia epiphyllous, opposite the aecia. Aecia hypophyllous, in groups, round, peridium cupulate, margin somewhat erose, whitish; aeciospores globoid or broadly el lipsoid, mostly with angulate edges, 13-19 x 11-17 wall colorless, minutely verrucose, about 1 p. thick® Uredinia chiefly hypophyllous, aparaphysate, minute, scattered or in groups, coalescent, ruptured epidermis noticeable, light cinnamon-brown, pulverulent; urediniospores globoid to broadly ellipsoid, 15-19 x 13-I8 jw, wall pale golden-brown, 65

1-1.5 M thick, densely echinulate, with J-k equatorial or superequatorial germ pores, without or with small, hyaline, echinulate caps0 Telia chestnut-brown, otherwise as the uredinia; teliospores ellipsoid or broadly globoid, 18-26 x 13-

1? ji, wall about light chestnut-brown lo5-2 p. thick, minute ly reticulate, appearing minutely verrucose, with meshes to about 1 p. diam, with a paler to hyaline papilla to k- p high at apex? pedicel colorless, thin-walled, mostly breaking near the hilum or to about 10 p. long0 Hosts and distribution in Brazils Desmodium albiflorum Salzm., Santo Angelo, Rio Grande do Sul (Jorstad, 1959). Type: on D. albiflorum Salzm. (now considered to be 2* affine Schlecht. by Schindler), Santo Angelo, Rio Grande

do Sul, C.A.M. Lindman (S, holotype)8

32. UROMYCES DESMQDIICOLA Jorst. var. DESMODII var. nov. (Fig. 39). Spermogoniis et aeciis ignotis,, Urediniis apara- physatis, flavidis, plerumque hypophyllis, rninutis, sparsis, mox nudis, pulverulentis, urediniosporae globoideae ad late

ellipsoideae, 15-19 x 13-16 membrana pallide flaveo- brunnea, 1-1.5 H crassa, denseque equinulata, poris germinationis 3(4), rare 2, equatorialibus vel leviter superequatorialibus, sine umbonibus vel parvulis, hyalinis, echinulatis, umbonibus praeditis. 66

Teliis quasi atro-brunneis aliter urediniis confor- mibus, teliosporae late globoideae, ovoideae8 ellipsoideae vel obovoideae, 18-26 x 13-18 membrana quasi pallide castaneo-brunnea, 1.5-2 }i crassa, reticulata, reticulationes usque ad lo5~2 diam, apice papilla quasi hyalina usque ad

4-5 p. altas pedicello plerumque brevi, hyalino, fragili vel rare 20-35 M longo.

Spermogonia and aecia unknown,, Uredinia aparaphysate, yellowish, chiefly hypophyllous, minute, scattered, soon naked, pulverulentj urediniospores globoid to broadly ellipsoid, 15-19 x 13-16 }*, wall pale golden-brown, 1-1.5 M thick, densely echinulate, with 3(4), rarely 2, equatorial or slightly above the equator germ pores, without caps or with small, hyaline, echinulate caps0 Telia about darkish brown, otherwise as the uredinia; teliospores broadly glo boid, ovoid, ellipsoid or obovoid, 18-26 x 13-18 ji, wall about light chestnut-brown, 1.5-2 thick at sides, reticu late with meshes to 1.5-2 p. diam, with a paler papilla at apex to 4-5 )i highs pedicel colorless thin-walled, mostly breaking short or near the hilum, less frequently to about 20-35 long. Hosts and distribution in Brazil: Desmodium incanum DC. (D. supinum DC.), Vicosa, Minas Gerais (Thurston, 1940). Type: on Desmodium incanum DC, (D. supinum DC.), Mi nas Gerais, A. 5. Muller, No. 541 (CUP, holotype). 6?

This variety differs from Uo desmodiicola Jorst. var. desmodiicola mostly because of the larger meshes of the teliospore walls to 1*5-2 diam, and by slight differences in germ pores of the urediniospores. SPECIES OF UROMYCES ON LEGUMES IN BRAZIL NOT VALID OR TRANSFERRED TO OTHER GENERA

1. Uromyces albescens Syd„» Ann# Mycolo 14!.668 1916, on

Pithecellobium glomeratum Benth® (Silveira, 1951)® syn0

Maravalia albescens SycL» Ann« Mycolo 22s 270o 1924„

valid name; Chaconia alutacea Juel, Bihang K9 Svenska

Veto Akado Handlo 23? 12o 189?0

2. Uromyces arachidis P0 Henn„, Hedwigia 35s 22**. 1896.

syn. Uredo arachidis Lagho* Uredo herbo El® Fries9 p« 106. 1895® valid names Puccinia arachidis Speg., Analo Soc. Cient#

Argente 17s 69-960 1884, on Arachis hypogaea L» (Fontou

ra and Nowackip 1967-70)0

3* Uromyces ingicola P» Henn„, Hedwigia 43» 157® 1904, on Inga sp« (Silveira, 1951)•

{ syn. Bitzea ingae (Syd9) Mains, Mycologia 3l 38® 1939*

on Inga edulis Marte and Inga sp. (Thurston, 1940)„ valid names Chaconia ingae (Sydo) Cumm., Mycologia 48:

602. 1956.

68 DISCUSSION AND CONCLUSIONS

Identification of species of Urom.yces on legumes in Brazil was based on a study of several characteristics, espe cially those of teliospores and urediniosporesa In the key based on uredinia and telia the species were divided in three sections based on wall ornamentation of teliosporeso To separate species9 emphasis was given to the number and position of germ pores of the urediniospores., In Section A„ teliospores smooth* six species were recognized,, Among them two varieties of U0 appendiculatus and one variety of U„ trifolii with wall of teliospores show ing variation from smooth to verrucosee Most of the changes in the species of this section were in nomenclature in which

U. lath.yrinus is the valid name for U® clavatusa a synonym also admitted by Jackson Uo viciae-fabae replaces U. fabae; U8 appendiculatus var« appendiculatus replaces U«

•phaseoli typica „ and U0 appendiculatus var0 brasiliensis re

places Uo phaseoli longepeduncula ti» Alsos the macrocyclic form, Uo trifolii (Hedw0 f0 ex DC0) Wint09 reported on Trifo- lium repens L0 and other Trifolium species by Lindquist and

Costa Neto (1963) is considered to be U0 trifolii var0 trifo lii. This variety differs from U0 trifolii (Hedw0 f0 ex DC.)

Wint. var® fallens (Artho) Arth0 by the number and position

69 70 of germ pores of the urediniospores and a slight difference in the wall ornamentation of teliosporesB Fontoura and

Nowacki (1967-70) reported U0 trifolii on Trifolium spp0 in Parana» As I have not been able to study these specimens I can not assume that all of them are U„ trifolii var0 trifolii, since U0 trifolii var. fallens and the demicyclic form

U. nerviphilus also can occur on Trifolium sppOQ In Section B with verrucose teliospores, 11 species were recognized,, Among them three that occur on Desmodium were also classified in Section C because the wall of teliospores showed variation from verrucose to irregularly or reg ularly reticulate, and two varieties of Uo appendiculatus and one variety of Ue trifolii also classified in Section Ao U. pisi reported on Pisum sativum in Brazil (Silveira, 1951; Luz, 1970) was studied only from European material since I was unable to get the Brazilian specimens,, It is important to mention that U« viciae-fabae can attack P. sativum (Arthur, 1962), and urediniospores can be produced on this host after inoculation with U„ striatus (Wilson and Henderson, 1966). In Section C, 17 species were recognized, with 12 species on Bauhinia, which includes two varieties of U. dietelianus„ two species on Desmodium, U« castaneus and U. desmodiicola var0 desmodiicola and U, desmodiicola var. desmodii, and three other species on Desmodium that were also classified in Section B. 71

U. castaneus and U. desmodiicolat originally de scribed as having verrucose teliospores, are included in this section because I interpret the teliospore walls as re ticulate * A summary of the results of the study of the species reported on legumes in Brazil are shown on Table 10 U. bauhiniicola Arth» , U„ imperfectus Arth0, U„ occidentalis

Diet*, and U0 superfixus Vesto, based on specimens analyzed, do not occur in Brazilo Ua corrugatus„ U0 .johowii» U„ nordensk.joldii„ and U8 trifolii-megalanthi reported by Watson (1971), probably do not occur in Brazil since there was no reaffirmation of these records from the source or from Brazilian literaturee U. neurocarpi, IAN No. 88^ = PUR FI6909, reported as occurring at Belem, appears to be a new species on Clitoria racemosa Benth., found only in the uredinial stage. Uo verus, described as having teliospores verrucose, is identical to Uo floralis that has teliospores minutely and uniformly reticulate. U. dietelianus appears to be a morphologically variable specieso When many speci mens were examined, it was difficult to classify them either as U. pannosus or U. dietelianus even when based on study of urediniospore germ pores® I concluded that U0 pannosus and U_. dietelianus represent one species. TABLE 1. Summary of the results of a taxonomic study of the species of Uromyces reported on legumes in Brazil

Species of Uromyces Reported Species Recognized

1, Uo anthemophilus Vest. Uo anthemophilus Vest, (type not studied) 2. U. anthyllidis (Grev.) Schroet. He anthyllidis (Grev.) Schroet,

3« U« bauhiniicola Arth.# IACM No# 3317* Uo floralis Vest0 if. Uo bauhiniicola Arth.„ IACM No. 309^** U® dietelianus Paz. var. and Noe U-222 nervicola var0 nov. 5. U. belemensis Albuq. & Figue. Uo belemensis Albuq. & Figue. 6. U. bradbur.yae Jacks. & Holw. Uo bradbur.yae Jacks. & Holw.

7. Uo castaneus Syd. Uo castaneus Syd0 8o Uo clavatus Diet. Uo lathyrinus Speg. 9o Uo corrugatus Speg. probably does not occur in Brazil 10. Us crotalariae (Arth.) J. Baxter Uo crotalariae (Arth.) J. Baxter 11. Uo desmodiicola Jorst. Uo desmodiicola Jorst. var. desmodiicola

12. U. dietelianus Paz. Ua dietelianus Paz. var. dietelianus TABLE 1. (Continued) Summary of a taxonomic study of species of Uromyces

Species of Uromyces Reported Species Recognized

13. U8 fabae (Pers.) dBy. Uo viciae-fabae (Pers.) Schroet.

14. U. floralis Vest. UB floralis Vests 15. U. foveolatus Juel Uo foveolatus Juel

16. ? Uo foveolatus Juel, Holw. No. 12?0 (S)* U0 foveolatus Juel

17. ? Uo foveolatus Juel, Holw. No. 1998 (S)* Uo bauhiniae P0 Henn.

18. ? Uo foveolatus Juel, Holw. Noe 1120 (S)* Uo bauhiniae P0 Henn.

19. U. goyazensis P. Henn0 Uo goyazensis P0 Henn. 20. U« hedysari-paniculati (Schw.) Farl. Uo hedysari-paniculati TSchWo) Farl0

21. Uo hedysari-paniculati (Schw0) Farl., Uo desmodiicola Jorst. var. Muller No. 5^1 (CUP)* desmodii var. nov0

220 U. hemmendorffii Vest. Uo hemmendorffii Vesto (type not studied)

23. U. hemmendorffii Vest., IACM No0 1361* Uo viegasii spB nov8 24. U. hemmendorffii Vest., Holw. No, 1025 Uo dietelianus Paz. var. (BPI)* dietelianus TABLE 1. (Continued) Summary of a taxonomic study of species of Uromyces

Species of Uromyces Reported Species Recognized

25. ? U« hemmendorffii Vest*, Holw0 Nos i883 Uo dietelianus Paz. var. (BPI)* dietelianus

26. ? U. imperfectus Arth., Muller No0 1077 U0 foveolatus Juel (CUP)*

27• ? Us imperfectus Arth.g Muller No. 296 Uo foveolatus Juel (CUP)*

289 ? Uo imperfectus Arth., Waller No. 7^1 Uo vicosensis sp8 nov0 (CUP) 29. Uo .jo'now 11 Dieto & Neger probably does not occur in Brazil

30. Uo nerviphilus (Grog.) Hotson Uo nerviphilus (Grog0) Hotson

31. Uo neurocarpi Dieto Uo neurocarpi Diet, 32e. ? Uo neurocarpi Dieta, IAN No. 884=PUR Uredo goeldii ined. F1^9 09"" 33® nordensk.joldii Diet. probably does not occur in Brazil

3^0 Uo occidentalis Dieto» IACM No0 ^035* Uo anthyllidis (Grev0) Schroet. 35c Uo orbicularis Diet, Uo orbicularis Diet#

# 36c Uo pannosus Vest., Vest. No0 1517 (BPI) Uo dietelianus Paz. var. dietelianus TABLE 1. (Continued) Summary of a taxonomic study of species of Uromyces

Species of Uromyces Reported Species Recognized

37. U. peraff ir.is Diet. = Uo peraffinis Diet. (type not studied)

38. Uo perlebiae Vest0 = Uo perlebiae Vest, (type not studied)

39. U. -perlebiae Vesto, IACM No. 3^59* - Us dietelianus Paz. var. dietelianus if-O. ? Uo perlebiae Vest0| Holw0 No0 1652 (BPI)* = U0 dietelianus Paz0 var0 dietelianus bio ? U0 perlebiae Vest., Holw. No, 1702 (BPI)* = Uo dietelianus Paz. var0 nervicola var8 nov0

^2. Ue phaseoli t.ypica Arth. = Uo appendiculatus (Pers0) Ung. var° appendiculatus

43. U. phaseoli longepedunculati Viegas nom0 = Uo appendiculatus (Pers0) Ungo nudum var0 brasiliensis var» nov,

U. pisi (Perso) V/int0 - U. pisi (DC.) Otth i}-5« Uo praetextus Vest. = Uo bauhiniae P0 Henn0

46. U. regius Vest. = U8 regius Vest, (type not studied)

47o U. striatus Schroet. Uo striatus Schroet. TABLE 1. (Continued) Summary of a taxonomic study of species of Uromyces

Species of Uromyces Reported Species Recognized

^8. U. superfixus Vest0, 1ACM No0 3829* Uo bauhiniae P. Henn.

^9» Ue trifolii (Hedw. f.) Lev. Uo trif olii (Hedw. f. ex DC.) Wint0 var. trifolii 50. Ue trifolii-megalanthi (Diet, & Neger) probably does not occur in Jackso & Holw. Brazil 51o U. unionensis Viegas Uo unionensis Viegas

; 52. U. verus Jacks. & Holw.f Holw. No. 1319 Uo floralis Vest. TBPI)*

* The citations of collection numbers for some specimens implies that these specimens were either doubtfully or incorrectly identified,, Herbarium citations follow Holmgren and Keuken (197^)«. 77

I have not been able to study the type specimens of

U. anthemophilus, U0 hemmendorffii„ Uo peraffinis„ U0

perlebiae and U0 regiuso However, based on the literature

(Vestergren5 1905s Dietel, 1922) the first three appear to

be very distinct species? the later two9 although recognized as distinct^ probably could be considered as varieties of U«

dietelianuse Also (Table 1)„ the author recognized two new spe

cies f Uo vicosensis and U„ viegasii on Bauhinia and two new

varieties9 Uo dietelianus var. nervicola, and U. desmodiicola var. desmodiia The life cycles and spore stages of the species found on legumes in Brazil are given (Table 2). Only two species

(U, nerviphilus and U0 orbicularis) are autoecious and

demicyclico Three species (Uo anthyllidis,, U0 pisin and U. striatus) are heteroecious and macrocyclic„ with spermogonia and aecia on Euphorbiaceaej and four species (all on Bauhinia) are microcyclic,. Six species and U® appendiculatus

var. appendiculatus„ U„ desmodiicola var0 desmodiicola B and

two varieties of U0 dietelianus are autoecious and macrocyclic. Also 11 species and two varieties (U„ appendiculatus var, brasiliensis and Uo desmodiicola var. desmodii) have life cycles considered uncertain because only uredinia and

telia have been foundo Uredinoid aecia have been found in U. neurocarpi, U. dietelianus var. dietelianus» U. TABLE 2. Life cycles and spore stages of the species of Uromyces on legumes in Brazil

UROMYCES species life cycle* spore stages* single or most 0 I II III important host

1. Uo anthemophilus (A) +B Bauhinia longifolia Do Dietr0 2. Uo anthyllidis (H) + + +B +B Lupinus albus L. 3. Uo appendiculatus var. appendiculatus (Am) + + +B Phaseolus vulgaris L,

k, u. appendiculatus var. brasiliensis (?) - - +B +B Phaseolus longepedunculatus Mart. 5. Uo bauhiniae (?) +B +B Bauhinia sp. 6o Uo belemensis (?) +B +B Ormosia nobilis Tul. 7. Uo bradburyae (?) +B +B Centrosema pubescens Bentho

8. Uo castaneus (?) +B +B Desmodium incanum DC0 9. Uo crotalariae (?) +B + Crotalaria vitellina Ker o 10. Uo desmodiicola var. desmodiicola (Am) +B +B +B +B Desmodium affine Schlechto (Do albiflorum Salzm.) TABLE 2. (Continued) Life cycles and spore stages of species of Uromyces

UROMYCES species life cycle* spore stages* single or most 0 I II III important host 11. U. desmodiicola var« desmodii (?) +B +B Desmodium incanum DCo (D« supinum DCo)

12. u- dietelianus var. dietelianus (Am) +B +B +B +B Bauhinia candicans Bentho

13. Uo dietelianus var. nervicola (Am) +B +B +B +B Bauhinia forficata Link

14. u. floralis (A) +B - - +B Bauhinia hiemalis Malme

15. u. foveolatus (?) - - 4-B +B Bauhinia hirsuta Vog.

16. u„ so.vazensis (A) - - - +B Bauhinia sp.

17. Uo hedvsari-paniculati (Am) + +B +B +B Desmodium uncinatum DC.

• - 00 Uo hemmendorffii (?) - +B +B Bauhinia forficata Link

19. UP lathvrinus (Am) - +B +B +B Lathyrus ma^ellanicus Lam«

20. nerviphilus (Ad) +B +B - +B Trifolium repens Lo ^3 VO TABLE 2. (Continued) Life cycles and spore stages of species of Uromyces

UROMYCES species life cycle* spore stages* single or most 0 I II III important host

21. U' neurocarpi (Am) +B +B +B +B Clitoria ca.ianifolia Bentho

22. U. orbicularis (Ad) - +B - +B Desmodium sp.

23. u. peraffinis (?) - - +B +B Bauhinia sp„

24. II* perlebiae (?) - - +B +B Bauhinia pentandra Vos. 25. pisi (H) + + +B +B Pisum sativum L.

26c u. regius (?) - - +B +B Bauhinia candicans Bentho

27. u. striatus (H) + + +B +B Medicago sativa L. 28. u. trifolii var. trifolii (Am) + + +B +B Trifolium sp.

29® u. unionensis (?) - - +B +B Desmodium sp. 30. u. viciae-fabae (Am) + + +B +B Vicia faba L. TABLE 2. (Continued) Life cycles and spore stages of species of Urom.yces

UROMYCES species life cycle* spore stages* single or most 0 I II III important host

31b U. vicosensis (Am) +B +B +B +B Bauhinia std.

32. UB viegasii (A) +B — +B Bauhinia forficata Link

* Symbols used are: 0 = Spermogonium + = Stage known 1 = Aecium = Stage unknown II = Uredinium +B = Stage found in Brazil III = Telium (A) = Autoecious microcyclic (Ad) = Autoecious demicyclic (Am) = Autoecious macrocyclic (H) = Heteroecious with 0 and I on Euphorbiaceae (?) = Life cycle uncertain because only II and III are known

oo f—1 82

dietelianus var« nervicola8 and U. vicosensis, but with more field work the discovery of other species with this type of aecium is probable0 Paraphyses in uredinia have been found in only two species9Uo hedysari-paniculati and U9 unionensis,, and teliospores with wall bilaminate have been found only in U9 bauhiniaeB U0 crotalariae8 and U0 foveolatusB

Geographicallyp Brazil is divided in five regions:

North (Norte)p Northeast (Nordeste)D Center-West (Centro-

Oeste)g East (Leste)„ and South (Sul)9 and species of Uromyces on legumes have been reported in all these regions

(Pig. 40)o Uo appendiculatus var0 ap-pendiculatus is the most widely distributeds followed by U0 hedysari-paniculati, Ua bauhiniae„ U„ dietelianus var0 dietelianusB U0 floralis„ U. viciae-fabae fl and Ue neurocarpi,, respectively® A few species of Uromyces on legumes have been trans ferred to other genera or are considered invalid0 U« arachidis, was described as belonging to this genus (Hennings,

1896)t probably because of the affinity of Uromyces for le gumes,, This species had been correctly described 12 years

before by Spegazzini (188^) as Puccinia arachidisn The only taxonomic distinction between Puccinia and Uromyces is in the number of cells of teliospores0 Geographical distribution of the species of Uromyces on legumes in Brazil.—Species are numbered with same numbers as found in key to species based on uredinia and telia. 84

U, albescens Syda v;as transferred "by Dietel (1924) to the genus Maravalia„ which differs from Uromyces in hav ing teliospores which germinate without dormancy by apical

prolongation of the wall without a germ pore0 Mains (1938) showed that the one-celled teliospores of this species devel

op in groups from basal cells and are sessile0 The basal

cells are not pedicels but sporogenous cellss He considered Maravalia albescens Sydo to be a synonym of Chaconia

alutacea Juel0

U0 ingicola P0 Henna (the name is based on the uredinial stage) was treated as a synonym of Bitzea ingae by Mains (1939)a Thirumalachar and Cummins (19^9) reduced Bitzea to synonymy with Chaconia and Cummins (1956) trans

ferred Bitzea ingae (Syde) Mains to Chaconiae Baxter (1962) transferred Haplopyxis crotalariae

(Arth«) Syd0 to the genus Uromyces because he found that

this rust was identical to U0 harmsianus Doidge„ The genus Haplopyxis was established by Sydow and Sydow (1919) to ac commodate one species originally described by Arthur (1918)

as Uropyxis crotalariae Artheo Since the first published description of the telial stage is that by Arthur (1918), the valid name of this rust is Uromyces crotalariae (Arth.) J. Baxter,, Consequently, he reduced Haplopyxis to synonymy with Uromyces. 85

The results of the study of the species of Uromyces reported on legumes in Brazil suggest the following conclusions: 1» Twenty-nine species of Uromyces occur on legumes in Brazilo Among them two varieties of U«

appendiculatus„ two varieties of Ua desmodiicola»

two varieties of U0 dietelianus, and one variety of Uo trifoliie

2. U® bauhiniicola, U„ corrugatus„ Uo .johowii9 U8

imperfectus8 U0 nordensk.ioldiie Ua occidentalis,

U, superfixus9 and U0 trifolii-megalanthi, prob ably do not occur in Brazilo 3* U* verus is identical to U. floralis, and U.

pannosus and U0 dietelianus represent a single specieso Among the taxa recognized„ U® vicosensis, U.

viegasiic U0 desmodiicola var» desmodii, and U«

dietelianus var0 nervicola are considered to be new,

5. U, phaseoli longepedunculati Viegas nom0 nudum was recognized as a distinct variety and named U.

appendiculatus var» brasiliensis var8 nov0 LITERATURE CITED

Albuquerque, F. C. 1971# Relacao das especies de Uredina- les coletadas na Amazoniao Pesq® Agropec. Braso Ser® Agron0 6s 1^7-150o j and Mo M„ FigueiredOo 1971° Descricao de uma nova especie de Uredinales da Amazonia Uromyces belemensis0 Pesqa Agropecs Bras0 Sero Agron0 6s 1^-5-1^6 0

Arthur,, J0 Co 1905o Leguminous rusts from Mexico0 Bot® Gazo 29s 385-396o

1907~1927o Uredinaleso No Amer0 Flora 7(2-12)s 85-8^80 I9I80 Uredinales of Guatemala based on col lections by E0 Wo Do Holway I0 Amer«, Jo Bot0 5« 325-356c 1920o New species of Uredineae-XIIo Bui. Torrey Bota Club ^-7s ^72 0 1962o Manual of the rusts in United States and Canada0 Hafner Publishing Co09 New Yorkp ^38 p0 Supplement by Go B0 Cummins0 p0 1A-2^AS

Augustin, Eos, and J0 Go C0 da Costa0 1971» Levantamento de racas fisiologicas de Uromyces phaseoli typica no Rio Grande do Sul e Santa Catarina em 1968 e 19690 Pesq0 Agropeco Bras« Ser9 Agron0 6s 109-113*

Baxter^ Jo Wa 19620 The status of the genus Haplopyxis (Uredinales) 0 Mycologia 5^°> ^37-^39°

Carvalhoff B® Ce L0 de , and C. Vieira0 1972e Comportamento de variedades de feijao (Phaseolus vulgaris L0)9 nas regioes de Irece e Tucanos Estado da Bahiao Rev0 Ceres 19(105)* 367-383®

Cummins, Go B0 1956® Nomenclatural changes for some North American Uredinales0 Mycologia **8: 6oi-6o80 1959• Illustrated genera of rust fungio Burgess Publishing Co®, Minneapolis, Minnesota, I3I P• 86 8?

Cummins, G. B. 1971o The rust fungi of cereals„ grasses and bamboose Springer-Verlag New York Inc00 New York Incop New Yorkp Heidelberg5 Berlin,, 570 p0

Dias, I, R0j and J0 Co da Costaa 19680 Identificacao de racas fisiologicas da ferrugem (Uromyces •phaseoli typica Artho) do fei.ioeiro '(Phase olus vulgaris L0) em duas regioes fisiograficas do Rio Grande do Sul0 Brasilo Pesq0 Agropeco Bras0 Ser0 Agrons 3s l65-l?0o

Dietel, P8 1895° Einige neue exotische Pilze0 Hedwigia 3^g 291-292o 1897o Uredineae brasilienses a clo Ule lectae® Hedwigia 36 s 26-37o

1899o Uredineae brasilienses a cl0 Ea Ule lectae IIo Hedwigia 38s 2^8-259» i903o Uber die Uromyces - Arten auf Lupinen. Hedwigia h2% 95-990 19220 Einige neue oder bemerkenswerte Uredinee0 Ann0 Mycol® 20s 293-295° 192^0 Kleine Beitrage zur Systematik der ~Uredineen IV0 Anna Mycolo 22s 269-273°

Fontoura, Og Sob and M0 Jo Nowackio 196?-70O Contribuicao ao levantamento fitossanitario do ParanaB Arq« Biolo Tecnolo 13(1*0* 109-183® Galli , Fo et al. 1968„ Manual de Fitopatologia.Doencas das plantas e seu controles Editora Agronomica Ceres LTDA8 Sao Paulo9 Brasil9 6^0 p0

Guyot, Ao Lo I9380 Les Uredinees0 Genre Uromyceso Encycl. Mycol®9 Paul Lechevalier9 Paris9 Vole 89 8s 4*38 p8 1957° Les Uredinees® Genre Uromyceso Encycl» Mycolot Paul Lechevalier8 Paris8 VolQ 29» 647 p®

Hariot9 P0 1892a Les Uromyces des legumineuseso Rev0 Mycolo 14 s ll-220

Hennings, P0 1895® Fungi goyazenses0 Hedwigia 34s 88-116. i8960 Beitrage zur Pilzflora Sudamerikas I. Hedwigia 35s 202-2o2» 88

Hennings, P. 1902a. Fungi S, Paulenses I. a cl. Puttemans collecti. Hedwigia 41: 104-118. 1902b. Fungi S. Paulenses II. a cl. Puttemans collecti. Hedwigia 4l: 295-3H< 1909. Fungi S. Paulenses IV, a cl. Puttemans collecti. Hedwigia 48: 1-20, Holmgren, P. K., and W. Keuken. 1974. Index herbariorum. Part I the herbaria of the world. 6th. ed. Inter national Bureau for Plant Taxonomy and Nomencla ture, Utrecht, Netherlands, 397 p«

Jackson, H. S0 I93I0 The Rusts of South America based on the Holway collections IV. Mycologia 23s 332-364. Jorstad, I. 1959° Uredinales from South America and Trop ical North America, chiefly collected by Swedish botanists II. Ark. Bot„ Ser 2, 4(5); 59-103<' 0 Junqueira Netto, A., K. L. Athow, and C. Vieira. 1969. Identificacao de racas fisiologicas de Uromyces phaseoli var. phaseoli, no Estado de Minas Gerais. Rev. Ceres 16(87)s 1-9« Krutman, S. 1968. Primeiros resultados sobre feijoeiro (Phaseolus vulgaris L.) na zona da mata de Pernambucoo Pesq. Agropec. Bras. Ser. Agron. 3: 111-125.

Lindquist, J. Co, and J. P. da Costa Neto. 1963. Uredi nales de Rio Grande do Sul0 Rev, Fac. Agron. 39: 111-159. Luz, N. K. 1970. Fungos de plantas olericolas no Rio Grande do Sul. Pesq. Agropec. Bras. Ser. Agron. 5: 53-59o Mains, E. B. 1938. Studies in the Uredinales, the genus Chaconia. Bull. Torrey Bot. Club 65: 625-629. 1939« Bitzea, a new genus in the Pucciniceae. Mycologia 31s33-42. Menezes, 0. B. de. 1952. Identificacao de 4 racas fisio logicas de ferrugem do feijoeiro. Dusenia, Curi- tiba 4: 309-312. 89

Pazschke^ 0. i89i. Rabenhorst-Winter0 Fungi europaei et extraeuropaei® Hedwigia 30s 199B ______1892o Erstes Verzeichnis der von Eo Ule in den jahren i883-87 in Brasilien gesammelten Pilze0 Hedwigia 31s 93-1140

Ponte9 Jo J0 da0 1974 0 Convenio de fitossanidade DNOCS/ UFC-relatorio anual/l9740 Departamento Nacional de Obras Contra as Secas e Universidade Federal do Cearas Fortalezap 43 p0 (mimeografado)«

Silveira® V0 Do 1951<> Elementos de fitopatologia0 Agrono- mia 10s 7-44®

Spegazzini0 Co 18840 Fungi guaraniticio Anala Soc0 Cients Argento 17s 90o

Sydow, He5 and P0 Sydow0 1907° Verzeichnis der von Herrn Fo Noack in Brasilien gesammelten Pilzee Ann8 %-colo 5s 3^8-363o 1910o Fungi paraenseso Hedwigia 49s 78-848 1919e Uber einige Uredineen mit quellbaren Membranen und erhohter Keimporenzahl9 Ann0 Mycolo 17s 101-107o

- Sydow# Pc, and Ho Sydow® I9IO0 Monographia Uredinearum0 Fre Borntraeger, Leipsigj volo 29 396 p<>

Thirumalachar, M» J®, and G* B8 Cummins0 19^9° The Taxo- nomic significance of sporogenous basal cells in the Uredinaleso Mycologia 4ls 523-526e

Thurston, H„ W0, Jr„ 194os The rusts of Minas Gerais„ Brazil based on collections by Ae So Muller* Mycol ogia 32s 290-3096

USDA. 1970o Index of plant diseases in the United States0 U0So Depto of Agriculture9 Agric0 Handbook Nos 165» Washington^ 531 P«>

Vasconcelos, I. 1971o Terceira contribuicao ao inventario de fungos de plantas do Ceara. Cien. Agron. 1(2)j 121-128. 90

Vestergren, T. 1905. Monographie der auf der Leguminosen- Gattung Bauhinia vorkommenden Uromyces - Arten. Ark* Bote 4(15)s 1-3^0

Viegas, A. P. 1945a0 Alguns fungos do Brasil IYS Uredina- les» Bragantia 5s 1-144. 1945bo Uns poucos fungos do Brasilo Bragantia 5s 561-582< 196lo Indice de fungos da America do Sul. Instituto Agronomico, Campinas, Sao Paulo, Brasil, 921 p. Watson, A. J. 1971o Foreign bacterial and fungus diseases of food, forage, and fiber cropss an annotated list. U.S. Dept. of Agriculture, Agric. Handbook No. 418. Washington, 111 p. Wilson, Me, and D. M. Henderson. 1966. British rust fungi. Cambridge University Press., London, 384 p.