Mycosphere 12(1): 89–162 (2021) www.mycosphere.org
ISSN 2077 7019
Article
Doi 10.5943/mycosphere/12/1/12
Indian Pucciniales: taxonomic outline with important descriptive notes
Gautam AK1, Avasthi S2, Verma RK3, Devadatha B4, Jayawardena RS5, Sushma6,
Ranadive KR7, Kashyap PL8, Bhadauria R2, Prasher IB9, Sharma VK3,
Niranjan M4,10, Jeewon R11
1
School of Agriculture, Abhilashi University, Mandi, Himachal Pradesh, 175028, India
School of Studies in Botany, Jiwaji University, Gwalior, Madhya Pradesh, 474011, India
3
Department of Plant Pathology, Punjab Agricultural University, Ludhiana, Punjab, 141004, India
4
Fungal Biotechnology Lab, Department of Biotechnology, School of Life Sciences, Pondicherry University, Kalapet,
Pondicherry, 605014, India
5
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, 57100, Thailand
6
Department of Botany, Dolphin PG College of Science and Agriculture Chunni Kalan, Fatehgarh Sahib, Punjab, India
7
Department of Botany, P.D.E.A.’s Annasaheb Magar Mahavidyalaya, Mahadevnagar, Hadapsar, Pune, Maharashtra,
India
8
ICAR-Indian Institute of Wheat and Barley Research (IIWBR), Karnal, Haryana, India
9
Department of Botany, Mycology and Plant Pathology Laboratory, Panjab University Chandigarh, 160014, India
10
Department of Botany, Rajiv Gandhi University, Rono Hills, Doimukh, Itanagar, Arunachal Pradesh, 791112, India
11
Department of Health Sciences, Faculty of Medicine and Health Sciences, University of Mauritius, Reduit, Mauritius
2
Gautam AK, Avasthi S, Verma RK, Devadatha B, Jayawardena RS, Sushma, Ranadive KR,
Kashyap PL, Bhadauria R, Prasher IB, Sharma VK, Niranjan M, Jeewon R 2021 – Indian
Pucciniales: taxonomic outline with important descriptive notes. Mycosphere 12(1), 89–162,
Doi 10.5943/mycosphere/12/1/2
Abstract
Rusts constitute a major group of the Kingdom Fungi and they are distributed all over the world
on a wide range of wild and cultivated plants. It is the largest natural group of plant pathogens
including 95% of the subphylum Pucciniomycotina and about 8% of all described Fungi. This article
provides an overview and outline of rust fungi of India with important descriptive notes. After
compilation of available literature on Indian rust fungi from various sources, it was observed that
these fungi are distributed in 16 families, 69 genera and 640 species. They belong to
Coleosporiaceae, Crossopsoraceae, Gymnosporangiaceae, Melampsoraceae, Milesinaceae,
Ochropsoraceae,
Phakopsoraceae,
Phragmidiaceae,
Pileolariaceae,
Pucciniaceae,
Pucciniastraceae, Raveneliaceae, Skierkaceae, Sphaerophragmiaceae, Tranzscheliaceae and
Zaghouaniaceae. There are still many rust fungi with uncertain taxonomic position, and they have
been referred to incertae sedis. The placement of all fungal genera is provided at the class, order and
family-level along with number of species in a genus. Notes for each rust family along with total
Indian records and other taxonomic information on transferred genera and species are also presented.
A phylogenetic analysis from a combined LSU and ITS dataset for 25 rust genera is presented to
provide a better understanding of their phylogeny and evolution.
Key words – India – Phylogeny – Pucciniomycotina – rust fungi – Systematics
Introduction
Rust fungi (Basidiomycota, Pucciniales) are a highly diverse group of obligate biotrophic
parasites, distributed in all geographical areas on a wide range of wild and cultivated plants ranging
Submitted 8 December 2020, Accepted 9 February 2021, Published 22 February 2021
Corresponding Author: Ajay K. Gautam – e-mail – a2gautam2006@gmail.com
89
from ferns, Gymnosperms and Angiosperms (Duplessis et al. 2011). They are called rusts as one of
their spore types i.e. the urediniospores and (uredinia) on host surface are often rust coloured. These
fungi are unique and fascinating group of organisms possessing diverse structures (spermogonia, two
anamorphs, teleomorphs and basidiospores) in their life cycle. Besides having up to five or six
morphologically and functionally distinct spore types, many rust fungi exhibit complicated life cycles
with variable host plant specificity. Because of their obligate parasitic nature, actively growing rusts
survive only on living hosts and produce teliospores towards the end of the growing season. Species
of Pucciniomycotina show simple life cycle (simple teliosporic yeasts) to the complex elaborate fivestage life cycles of the biotrophic rust fungi, the latter regarded as the most complex organisms in
Kingdom Fungi (Lutz et al. 2004). Along with various spore types, some rust fungi require alternation
between two exclusive and unrelated host plant taxa to complete their life cycle (heteroecious rust),
although others can complete their life cycle on a single host plant (autoecious rust) (Kolmer et al.
2009). Parasitism and host specialization is highly developed in rust fungi (Savile et al. 1971,
Duplessis et al. 2011). They constitute one of the major groups of plant pathogenic fungi.
Puccininales is the most speciose order in Pucciniomycotina and include 95% of the subphylum and
ca. 8% of all described Fungi (Kirk et al. 2008).
The rust fungi have a long research history, mostly due to their economic importance in
agriculture and forestry, and easily noticeable symptoms (Cummins & Hiratsuka 2003).
Taxonomically the rust fungi belong to Pucciniomycotina, one of the three subphyla of
Basidiomycota. Further, these fungi are placed in class Pucciniomycetes and order Pucciniales. In
some systems of classifications, the rust fungi are placed in the subclass Heterobasidiomycetes based
on basidial morphology while in other systems, they are with smut fungi in Teliomycetes based on
their similar basidial characteristics (Talbot 1971, Webster 1980). Different morphological characters
have been emphasized in earlier system of rust taxonomy and classifications (Ono & Hennen 1983).
Initially, rust fungi were classified into three (or four) families, Melampsoraceae, (Coleosporiaceae),
Pucciniaceae and Zaghouaniaceae based on the characteristics of basidia and teliospores (Sydow &
Sydow 1915, Cunningham 1931). Classification of rust fungi in subfamilies or tribes and morphology
of telia in taxonomy was also considered (Sydow & Sydow 1915, Dietel 1928). Use of morphology
of spermogonial (0) and aecial (I) stages was emphasized by Hiratsuka & Cummins (1963) and
Hiratsuka (1983) in the classification of rust fungi. The morphological characters of telium
(Thirumalachar & Cummins 1949) and teliospores (Dietel 1928, Thirumalachar & Cummins 1948,
Thirumalachar & Mundkur 1949) played an important role in the taxonomic placement of rust fungi
(Alexopoulus 1962). However, with the combination of different character of uredinia, aecia, type of
spermogonia and telia, there has been several conflicting taxonomic hypotheses with time and this
led to the proposal of 13-family system of classification of rust fungi. Based on telial morphology,
three families were accepted: Melampsoraceae (sessile teliospores formed in columns in the telium),
Pucciniaceae (stalked teliospores, produced in a single layer in the telium) and Coleosporiaceae (do
not form a promycelium but karyogamy and meiosis occur directly within the teliospores that become
septate during the germination process). Cummins & Hiratsuka (1983, 2003) proposed 13 families.
This was one of the the most accepted systems of rust classification till the incorporation of molecular
data in systematic studies.
With the use of modern techniques (molecular studies) along with conventional methods
(morphological studies), new taxonomic suggestions have been proposed regarding interrelationships
of different groups of fungi. However, the earlier systems of classifications of rust fungi were mainly
based on shape, size and other morphological characters of different spores and spore producing
structures. Based on recent molecular techniques (DNA sequence data from the large ribosomal
subunit) and ultrastructural investigations, it was observed that rusts are distantly related to some of
the smuts (Aime et al. 2006, Kijpornyongpan et al. 2018). Subsequently Swann & Taylor (1995a, b)
and Swann et al. (2001) proposed the separation of class Urediniomycetes (the rust fungi), including
the rusts (Uredinales), from class Ustilaginomycetes (the smut fungi) and class Hymenomycetes
mushrooms and shelf or bracket fungi) under division Basidiomycota. Within the class
Urediniomycetes, true rusts (Uredinales) account for over 95% of the species and more than 75% of
90
the genera. Several research series on fungal diversity published in recent years provided an updated
information on addition, exclusion, correct taxonomic position and transferred taxa of fungi. Aime
(2006) examined representative species from the 13 families proposed by Cummins & Hiratsuka
(2003) and based on sequence analyses of the 18S and 28S nuclear rDNA regions, three major
suborders were proposed Uredinineae, Melampsorineae and Mikronegeriineae. The Uredinineae
includes species which produce the aecial stage on angiospermous host with pedicellate teliospores.
Similarly, Melampsorineae comprises heteroecious, mostly macrocyclic rust species producing
aecial stage on gymnosperms and producing sessile teliospores. Mikronegeriineae accomodates
species with aecial stage usually on non-pine gymnosperms and in microcyclic types the teliospores
functioned as urediniospores, with short pedicellate or sessile teliospores. He et al. (2019) presented
notes, outline and divergence times of Basidiomycota. They differentiated Urediniomycetes from
Basidiomycetes, as the nuclear membrane partially degrades during mitosis (semiopen pleuromitosis)
in some Urediniomycetes while mitosis in basidiomycetes proceeds with preservation of the nuclear
membrane (intranuclear pleuromitosis). They also differentiated rusts and smuts as non-basidiomata
forming members of Basidiomycota, which comprises Pucciniomycotina and Ustilaginomycotina,
respectively. They included eight families, Coleosporiaceae, Mikronegeriaceae, Phakopsoraceae,
Phragmidiaceae, Pileolariaceae, Pucciniaceae, Raveneliaceae and Sphaerophragmiaceae in order
Pucciniales, class Pucciniomycetes and subphylum Pucciniomycotina in their phylogenetic studies.
However, they also presented the estimated number for taxa in Basidiomycota, of which, order
Pucciniales possesses 15 families, 162 genera and 8105 species. Recently, Wijayawardene et al.
(2020) outlined the kingdom Fungi up to genus level. They also presented 15 families (138 genera)
under the order Pucciniales, class Pucciniomycetes and phylum Basidiomycota. It is interesting to
mention here that the majority of Pucciniomycotina species (ca. 7500 of nearly 8500) belong to a
single order Pucciniales that cause rust diseases in numerous plants (He et al. 2019, Wijayawardene
et al. 2020, Aime & McTaggart 2020). The recent higher-rank classification for rust fungi is provided
by Aime & McTaggart (2020), wherein they proposed the addition of four new suborders and seven
new families, with some amendments in existing families. The classification of Pucciniales now
comprises seven suborders and 18 families.
To understand the status of research on rust fungi in India, we have started this part here with
the beginning of Indian mycological research. Here foreign visiting scientists or emigrant experts in
the 18th and 19th centuries carried out most of the research. This started with the mycological studies
initiated by K.R. Kirtikar in late 19th century who collected and identified numerous fungi. Although
the special credit to initiate mycological research in India goes to A. Barclay and E.J. Butler because
of their contribution to initiate and organize research on mycology and plant pathology in India. After
the establishment of the Imperial Agricultural Research Institute at Pusa (Bihar) during the year 1905
with the generous grant of 30,000 pounds from an American philanthropist, Mr. Henry Phipps,
mycological research in India gained momentum. With the earlier efforts of E.J. Butler, the first
imperial mycologists to the then British Government of India, a firm foundation of mycology and
plant pathology was laid in this country and he is aptly referred to as the Father of Indian Mycology
(Subramanian 1986). This research continued with the passage of time and involvement of many
more mycologists. With the advancement in mycological research, B.B. Mundkur and M.J.
Thirumalachar laid down a pioneer contribution in the studies of rust and smut fungi as Ustilaginales
of India (Mundkar & Thirumalachar 1952). Foundation of the Indian Phytopathological Society was
also laid down by B. B. Mundkur along with S.R. Bose, both sereved as its earliest presidents. K.C.
Mehta (1940) studied the problem of the recurrence of the wheat rust in plains of India. They mainly
focused on cereal rust of India. Balchandra Bhavanishankar Mundkur and Mandayam Jeersannidhi
Thirumalachar in 1952 jointly published a consolidated list of Indian Ustilaginales. Similarly,
Thirumalachar & Mundkar (1950) published a very useful appendix of genera of rust fungi.
Simultaneously, Hans Sydow, a German mycologist, son of Paul Sydow also contributed a lot in
understanding the Himalayan mycoflora including rust fungi. Similarly, George Baker Cummins, A.
Barclay and Joseph Charles Arthur investigated the rusts occurring near North Western Himalayas.
D.P. Mishra along with other mycologists also investigated rust fungi of agricultural crops in India.
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Several other mycologists contributed significantly to the development of research on rust fungi in
India on a regional basis. Pioneering work of C. Mohanan cannot be neglected as modern
mycologists. He worked on biodiversity of plant pathogenic fungi of the Western Ghats and
published a book entitled “Rust Fungi of Kerala, India”. Some names worth mentioning are Ramesh
Chand Sharma, Sanjeev Sharma, R.K. Sharma, S.N. Sachan and Ajay Kumar Gautam, who actively
worked in the past and are still engaged in studies of various aspects of rust fungi of Himachal
Pradesh. Dr. E.J. Butler & G.R. Bisby compiled a monograph “The Fungi of India”, in 1931. The
fungi of India series have been revised from time to time and updated by several workers. However,
the contribution of great Indian mycologists K.D. Bagchee, T.S. Ramakrishnan, J.H. Mitter, K.J.
Narsimhan, S.N. Das Gupta, R.N. Tandon, R. Prasad, T.S. Sadasivan, C.V. Subramaniam and many
more cannot be ignored. The taxonomic research on rust fungi in India has been based primarily on
morphology of certain spore stages. Only a few studies published recently have employed modern
tools and techniques for identification of rust fungi. Several institutes like Indian Type Culture
Collection (ITCC) New Delhi; National Fungal Culture Collection of India (NFCCI) Pune,
Maharashtra; CSIR-IMTECH Chandigarh, NBAIM Mau and many more are actively carrying
research on fungal taxonomy and other related aspects and providing facilities for molecular
characterisation of fungi including rust fungi in India. A major scientific breakthrough of the Indian
Council of Agricultural Research (ICAR) scientists lead to the decoding of genomes of 15 strains of
wheat rust fungus Puccinia triticina. Herbarium Cryptogamae Indiae Orientalis (HCIO) has a rich
collection of rusts, smuts, powdery mildews and meliolales fungi and has more than 3500 type
specimens (Maheswari et al. 2012). HCIO documented a comprehensive checklist of Puccinia
species of India along with herbarium photographs and brief description (Kamil et al. 2013).
Similarly, “Rust fungi of Kerala, India” (Mohanan 2010); First checklist of rust fungi in the genus
Puccinia from Himachal Pradesh, India” (Gautam & Avasthi 2016a); and “A checklist of rust fungi
from Himachal Pradesh, India” (Gautam & Avasthi 2019) are the recent compilations of rust fungi
from India. In subsequent years, a plethora of articles on Pucciniales (Uredinales) has been
published, which has added a lot to understanding this group of fungi, thus it is essential and
pragmmatic to compile it into a single document. Therefore, we are starting a series of publications
on rust fungal diversity from India, and this first paper provides basic information and the latest
trends related to taxonomic outline of rust fungi of India with important descriptive notes.
Materials & Methods
Layout of the paper
A brief description of each genus up to its higher taxonomic rank related with rust fungi of
India is provided. During the listing of genera, their species, and other higher taxonomic ranks into a
single outline, all generic names are listed from previously published literature pertaining to rust
fungi of India. The detailed literature in reference to Indian rust fungi is summarised in the present
study under the heading “literature used during the study of rust fungi”. The names of some species
have been replaced by currently accepted names after consultation of MycoBank
(www.mycobank.org)/ and Species Fungorum (www.speciesfungorum.org) websites and this has
been indicated in such cases. For general outline of Indian rust fungi, we adopted He et al. (2019)
and Wijayawardene et al. (2020). Aime & McTaggart (2020) was followed to provide a higher-rank
classification of rust fungi. To confirm their scientific entity where some generic/species names have
been updated with currently accepted name, more literature on rust fungi was consulted (Cummins
& Hiratsuka 2003, Aime 2006, Aime et al. 2018, Aime & McTaggart 2020. After complete
verification, accepted taxa of rust fungi of Basidiomycota up to species level are presented in detailed.
The existing and currently accepted names of various genera and species of rust fungi are provided
as a separate section in this manuscript.
Phylogenetic analyses
Most of the rust fungi reported from India were characterized mainly based on the
92
morphological characters of uredia and telia or other successive stages observed on collected
samples. However, rust fungi reported from India lack molecular studies.
Based on earlier studies on Indian rust fungi, a checklist was prepared and the DNA sequence
data from the LSU and ITS rDNA regions available for same rust fungi reported from other countries
were downloaded from GenBank and through published literature (Maier et al. 2003, Aime 2006,
Aime et al. 2006, Aime & McTaggart 2020). Individual nucleotide sequences of LSU and ITS were
aligned distinctly using MAFFT 7 (http://mafft. cbrc.jp/alignment/server/) (Katoh & Standley 2013)
and then manual checking and editing where necessary in BioEdit v.7.0.9 (Hall 1999). The ITS
sequences of taxa containing weak aligned portions, incomplete data, missing sequence data and gaps
were removed. The separate aligned gene regions of LSU and ITS were combined in BioEdit. The
combined multigene sequence alignment was converted to PHYLIP format (.phy) using ALTER
(alignment trans-formation environment: (http://sing.eiuvigo.es/ALTER/; 2021) for randomized
accelerated maximum likelihood (RAxML) analysis. The aligned LSU and ITS single gene datasets
and a concatenated dataset of LSU and ITS genes were analyzed with maximum likelihood using the
RAxML-HPC2 on XSEDE (8.2.8) (Stamatakis et al. 2008, Stamatakis 2014) in the CIPRES Science
Gateway platform (Miller et al. 2010) using GTR+I+G model of evolution. Maximum Likelihood
bootstrap values greater than 70% were given above each node. Phylogenetic trees were visualized
with FigTree v1.4.0 program (Rambaut 2012) and reorganized in Microsoft power point (2016).
Table 1 GenBank and voucher/culture collection accession numbers of species included in the
phylogenetic study
Taxon
Voucher/culture
Cerotelium fici
Chrysomyxa pirolae
Coleosporium asterum
Coleosporium bletiae
Coleosporium campanulae
Coleosporium campanulae
Coleosporium inulae
Coleosporium inulae
Coleosporium ipomoeae
LAH20019AM
190CHP_PCG_DU2
MCA3077
BSC1
-LB09265/ZT_Myc_58002
U717
LB09168/ZT_Myc_57996
JRH 485
GenBank accession
Numbers
ITS
LSU
-MK135779
GU049555
-MG907226
-MN108162
-KP017565
KY810467 --MG907223
KY810470 -MF769644 --
Coleosporium ipomoeae
Coleosporium senecionis
Coleosporium senecionis
Coleosporium xanthoxyli
Cronartium quercuum
Cronartium ribicola
R232
LB08877/ZT_Myc_57995
PDD 98309
KUS-F30013
CqvGa-1
--
-KY810472
--L76495
KX963430
Crossopsora ziziphi
Gymnosporangium
clavariiforme
Gymnosporangium
clavariiforme
Gymnosporangium
confusum
Gymnosporangium
confusum
Hyalopsora polypodii
BPI 877877
LD 1019
-HM114220
EU851160
-KJ716348
MK530184
--AF426240
MG744558
--
HMAS:24626
--
KU342766
Zhao et al. (2016)
LD 1021
HM114219
--
Dervis et al. (2011)
20140808B H22
--
KP261043
BPI 893256
KY798367
--
Hyalopsora polypodii
DB 1681
--
AY512852
Fernandez & Alvarado
(2016)
Demers & Castlebury
(unpublished)
Begerow et al.
(unpublished)
References
Ishaq et al. (unpublished)
Feau et al. (2011)
Aime et al. (2018)
Zhou (unpublished)
Tian et al. (unpublished)
Beenken et al. (2017)
Aime et al. (2018)
Beenken et al. (2017)
McTaggart & Aime
(2018)
Zuluaga et al. (2011)
Beenken et al. (2017)
Shin et al. (2019)
Vogler & Bruns (1998)
Kaitera et al. (2017)
Maier et al. (2003)
Souza et al. (2018)
Dervis et al. (2011)
93
Table 1 Continued.
Taxon
Voucher/culture
Kernkampella breyniae
Kweilingia divina
Leucotelium prunipersicae
Maravalia pterocarpi
Melampsora caprearum
Melampsora caprearum
Melampsora epitea
Melampsora epitea
Melampsora euphorbiae
Melampsora euphorbiae
Melampsora
hypericorum
Melampsora
hypericorum
Melampsora populnea
Melampsora populnea
Melampsora salicisalbae
Melampsora salicisalbae
Melampsoridium
betulinum
Melampsoridium
betulinum
Melampsoridium
hiratsukanum
Melampsoridium
hiratsukanum
Milesina exigua
Milesina polypodii
Nyssopsora thwaitesii
Ochropsora ariae
Phakopsora apoda
BRIP:56909
MCA3493
--
GenBank accession
Numbers
ITS
LSU
-KJ862346
-MG907215
AB097450 --
HNCM1
GE14_3_2
NYS-F-003819
-1046MEE-SA-QC.1
-BPI 863501
BPI 893298
KU301795
KY649193
--GQ479218
EF192199
---
--KU550033
DQ354564
--DQ437504
KY798351
PDD 97325
KJ716353
--
892MPO-PTA-FR3.1
-HMAAC4068 HMAAC4068
EU808037
---
-AY444786
MK372199
13125 F
FJ455128
--
KR-M-0048135
--
MK302187
Eslami et al.
(unpublished)
Bubner et al. (2019)
H 6034375
KF031552
--
McKenzie et al. (2013)
421
KC313888
--
Blomquist et al. (2014)
KR-M-0048149
--
MK302188
Bubner et al. (2019)
KR-M-0050247
KR-M-0043190
AMH:9528
KR-M-42604
PDD 72076
MH908478
-KF550283
KX228773
MG461668
MK302211
MK302190
-KX228778
MG461668
Phakopsora cingens
Phakopsora meibomiae
Phakopsora meibomiae
Phakopsora phyllanthi
Phakopsora phyllanthi
BRIP:55628
Brazil 82-1
R188
83
BPI 843632
-AF333501
-KF528025
--
KP729474
-EU851164
-KY764084
Phakopsora ziziphivulgaris
Phragmidium barclayi
Phragmidium
brevipedicellatum
Phragmidium butleri
Phragmidium fragariae
Phragmidium
mucronatum
Phragmidium
potentillae
HMJAU8595
MK296536
MK296506
Bubner et al. (2019)
Bubner et al. (2019)
Baiswar et al. (2014)
Scholler et al. (2019)
Demers & Castlebury
(unpublished)
Maier et al. (2016)
Frederick et al. (2002)
Zuluaga et al. (2011)
Beenken (2014)
Demers et al.
(unpublished)
Ji (unpublished)
HMAS-67281
HMUT100463
---
MG669117
KU059170
Liu et al. (2018)
Xu et al. (unpublished)
HMAS-67841
-TFS01
----
MG669118
AF426217
KJ867552
Liu et al. (2018)
Maier et al. (2003)
El-Deeb (unpublished)
BRIP:60089
--
KT199403
McTaggart et al. (2016b)
References
McTaggart et al. (2015)
Aime et al. (2018)
Osaki (unpublished)
Wang et al. (2016)
Piskur (unpublished)
Zhao (unpublished)
Aime (2006)
Vialle (unpublished)
Aime (unpublished)
Aime et al. (2006)
Demers & Castlebury
(unpublished)
Padamsee & McKenzie
(2014)
Feau et al. (2009)
Pei et al. (2005)
Wang (unpublished)
94
Table 1 Continued.
Taxon
Voucher/culture
Phragmidium
potentillae
Phragmidium rosaemoschatae
Pileolaria pistaciae
Puccinia actaeaeagropyri
Puccinia antirrhini
HMJAU8609
GenBank accession
Numbers
ITS
LSU
MK296538 --
BPI 893257
--
KY798368
-TUB 14959
MG860928
--
KY314266
DQ917746
BPI 910208
--
KY764090
Puccinia arenariae
BPI 893275
Puccinia argentata
IMI 502182
--
KC433402
Puccinia bistortae
Puccinia brachypodii
Puccinia brachypodii
Puccinia bupleuri
TUB 14964
HSZ0975
BRIP 59466
BPI 910217
-GQ457303
---
DQ917697
-KX999868
KY764100
Puccinia canaliculata
Puccinia carthami
U-77
SAF 1
HQ412647
AF064822
Puccinia carthami
Puccinia cenchri
-BPI055581
--
AY787782
KY575080
Puccinia chloridisincompletae
Puccinia chrysanthemi
HSZ1393
KM096427
KM096427
HSZ2154
KX369039
--
Puccinia chrysanthemi
Puccinia circaeae
Puccinia coronata
Puccinia crepidisjaponicae
Puccinia cyperi
Puccinia dactylidina
Puccinia dioicae
Puccinia drabae
Puccinia ferruginosa
R79F
TUB 14969
-BPI 893282
-----
HQ201322
DQ917716
DQ354526
KY798360
BRIP 60997
PUR F15426
DAR 77052
R209-16555
IBA7553
KU296885
JX533546
EF635897
EU014054
--
KU296885
JX533546
-EU014055
AB190901
Puccinia ferruginosa
TSH-R6237
AB188126
--
Puccinia graminis
Puccinia heterospora
72_2
BPI 893307
-KY798362
JQ688991
--
Puccinia heterospora
Puccinia heucherae
Puccinia hieracii
Puccinia hordei
BRIP 60937
RHS5296/05
DAOM 240969
PDD:101656
-----
KU296886
DQ359702
HQ317515
KX985762
Puccinia hydrocotyles
Puccinia iridis
Puccinia iridis
Puccinia kuehnii
-KUS-F23394
KUS-F30180
BPI 881011
--MK446718
--
GU936635
MK446720
KY798385
HQ666891
References
Ji (unpublished)
Demers & Castlebury
(unpublished)
Ishaq et al. (2020)
Maier et al. (2007)
Demers et al.
(unpublished)
Demers et al.
(unpublished)
Tanner et al.
(unpublished)
Maier et al. (2007)
Jin et al. (2010)
Marin-Felix et al. (2017)
Demers et al.
(unpublished)
Deadman et al. (2011)
Berthier-Schaad
(unpublished)
Deadman et al. (2005)
Demers & Castlebury
(unpublished)
Mahadevakumar
(unpublished)
Szabo & Nguyen
(unpublished)
Alaei (unpublished)
Maier et al. (2007)
Aime et al. (2006)
Demers & Castlebury
(unpublished)
McTaggart et al. (2016a)
Liu et al. (2013)
Morin et al. (2009)
Alaei et al. (2009)
Engkhaninun et al.
(unpublished)
Engkhaninun et al.
(2005)
Berlin et al. (2012)
Demers & Castlebury
(unpublished)
McTaggart et al. (2016a)
Henricot et al. (2007)
Liu et al. (2015)
Padamsee & McKenzie
(2017)
Zuluaga et al. (2011)
Choi et al. (2019)
Choi et al. (2019)
Saumtally et al. (2011)
95
Table 1 Continued.
Taxon
Voucher/culture
Puccinia kuehnii
Puccinia lantanae
Puccinia lateritia
090687
R190
BPI 910249
GenBank accession
Numbers
ITS
LSU
GU564421 --EU851144
-KY764134
Puccinia liberta
Puccinia linkii
BRIP 59686
DAOM:242721
-KM851040
KX999881
--
uccinia linkii
DAOM:243230
--
KM851041
Puccinia malvacearum
AFTOL-ID 1629
EF561641
--
Puccinia malvacearum
Puccinia
melanocephala
Puccinia
melanocephala
Puccinia menthae
U1384
PM_NayMex_Xalsco
-MG564638
MG907249
--
PM_ColMex_CE_Tecoman
--
MG564636
BPI 910255
KY764141
--
Puccinia menthae
Puccinia nakanishikii
BPI 871110
BPI 910261
KY764147
DQ354513
--
Puccinia nakanishikii
Puccinia nepalensis
Puccinia nepalensis
Puccinia nitida
-BA66
BA651
BPI 843472
--KX225481
--
GU058002
KX014746
-KY764148
Puccinia oahuensis
BPI087481
--
KY575092
Puccinia obscura
Puccinia obscura
Puccinia ocimi
KR14322
-BPI 910262
-AF468042
KY764149
FJ669234
-KY764149
Puccinia operta
BPI142436
Puccinia oxalidis
Puccinia oxalidis
KUS-F27920
BPI 893310
MH325473
--
-KY798348
Puccinia paspali
BPI841180
--
KY575098
Puccinia peradeniyae
Puccinia pimpinellae
Puccinia polygoniamphibii
Puccinia polysora
Puccinia porri
BPI 089014
-BPI 893289
KX190906
GU058023
--
KX190906
GU058023
KY798381
-22-16
HM467909
KY492366
GU058024
--
Puccinia recondita
BPI 910319
--
KY798399
Puccinia rufipes
Puccinia scirpi
Puccinia striiformis
Puccinia striiformis
Puccinia substriata
-BRIP 61027
HSZ1834
HSZ1828
--
AJ406071
KX999892
GQ457306
-GU058028
-KX999892
-GQ457304
--
KY575095
References
Glynn et al. (2010)
Zuluaga et al. (2011)
Demers et al.
(unpublished)
Marin-Felix et al. (2017)
Mulvey & Hambleton
(2015)
Mulvey & Hambleton
(2015)
Matheny & Hibbett
(unpublished)
Aime et al. (2018)
Bermudez et al.
(unpublished)
Bermudez et al.
(unpublished)
Demers et al.
(unpublished)
Aime et al. (2006)
Demers et al.
(unpublished)
Dixon et al. (2010)
Ali et al. (2017)
Ali et al. 2017)
Demers et al.
(unpublished)
Demers & Castlebury
(unpublished)
Scholler et al. (2011)
Weber et al. (2003)
Demers et al.
(unpublished)
Demers & Castlebury
(unpublished)
Lee et al. (2019)
Demers & Castlebury
(unpublished)
Demers & Castlebury
(unpublished)
Demers et al. (2017)
Dixon et al. (2010)
Demers & Castlebury
(unpublished)
Yu et al. (unpublished)
Ristic et al.
(unpublished)
Demers & Castlebury
(unpublished)
Virtudazo et al. (2001)
Marin-Felix et al. (2017)
Jin et al. (2010)
Jin et al. (2010)
Dixon et al. (2010)
96
Table 1 Continued.
GenBank accession
Numbers
ITS
LSU
KY575101
Taxon
Voucher/culture
Puccinia substriata
BPI106260
Puccinia tanaceti
Puccinia thlaspeos
Puccinia thlaspeos
Puccinia turgida
Puccinia turgida
Puccinia versicolor
Puccinia violae
Puccinia violae
Puccinia xanthii
Puccinia xanthii
Pucciniastrum coryli
Ravenelia acaciaearabicae
Ravenelia acaciaepennatulae
Ravenelia acaciicola
Ravenelia evansii
Ravenelia evansii
Tranzschelia discolor
Tranzschelia discolor
R221
--NA224
NA223
U902
BPI 842321
-BRIP48819
BRIP 56946
TSH-R4237 (IBA8641)
PREM61853
HQ201323
L76177
L76183
MH144383
MH144382
-DQ354509
-EU659694
References
AB221419
--
MG907252
-GU058029
-KX999896
-MN072675
Demers & Castlebury
(unpublished)
Alaei (unpublished)
Roy et al. (1998)
Roy et al. (1998)
Otálora & Berndt (2018)
Otálora & Berndt (2018)
Aime et al. (2018)
Aime (2006)
Dixon et al. (2010)
Seier et al. (2009)
Marin-Felix et al. (2017)
Liang et al. (2006)
Ebinghaus et al. (2020)
U115
--
MG907213
Aime et al. (2018)
PREM61861
PREM61209
PREM61028
U-884
BRIP 57662
-MG945959
-DQ995341
--
MN072683
-MG945996
-KR994891
Tranzschelia prunispinosae
Uromyces aloes
Uromyces aloes
Uromyces cicerisarietini
Uromyces cicerisarietini
Uromyces clignyi
Uromyces commelinae
Uromyces coronatus
Uromyces coronatus
KR-M-0002755
--
KX228774
Ebinghaus et al. (2020)
Ebinghaus et al. (2018)
Ebinghaus et al. (2018)
Deadman et al. (2007)
Doungsa-Ard
(unpublished)
Scholler et al. (2019)
2020-6-28-0005
WM 3290
--
MT136509
-GU058030
-DQ917740
--
Bily et al. (unpublished)
Maier et al. (2007)
Dixon et al. (2010)
--
GQ914998
Stuteville et al. (2010)
DAOM 192217
JW100
DAOM 32991
BPI 910293
HM131364
KF982855
HM131365
---KY764191
Uromyces dactylidis
Uromyces dactylidis
Uromyces dolicholi
Uromyces eragrostidis
Uromyces eragrostidis
PRC:705
TUB 14997
DAOM 116149
DAOM 106767
BPI004474
KM667955
-HQ317563
HQ317561
--
-DQ917745
HQ317563
-KY575113
Uromyces euphorbiae
PUR N11621
KT750329
--
Uromyces euphorbiae
BPI 863673
--
KT750330
Uromyces geranii
Uromyces hedysariobscuri
Uromyces hedysariobscuri
Uromyces lespedezaeprocumbentis
BRIP 60100
DAOM 189699
-HQ317573
KX999898
--
Liu & Hambleton (2013)
Kwon & Kim (2014)
Liu & Hambleton (2013)
Demers et al.
(unpublished)
Hrabetova et al. (2015)
Maier et al. (2007)
Liu et al. (2015)
Liu et al. (2015)
Demers & Castlebury
(unpublished)
Rosskopf et al.
(unpublished)
Rosskopf et al.
(unpublished)
Marin-Felix et al. (2017)
Liu et al. (2015)
DAOM 223015
--
HQ317572
Liu et al. (2015)
BPI 910294
KY764193
KY764193
Demers et al.
(unpublished)
HQ201324
--
97
Table 1 Continued.
Taxon
Voucher/culture
Uromyces minor
Uromyces muscari
Uromyces orientalis
Uromyces peglerae
MVAP50000151
DAOM 75626
BRIP 60934
BPI843311
GenBank accession
Numbers
ITS
LSU
MK045314 MK045314
HQ317552 HQ317552
KX999899 KX999899
KY575116
Uromyces pisi-sativi
--
DQ521591
--
Uromyces pisi-sativi
Uromyces polygoniavicularis
Uromyces rumicis
BRIP 60151
DAOM 181565
--
KX999900
HQ317558
PDD:93529
KX985763
--
Uromyces rumicis
BPI 910298
Uromyces setariaeitalicae
Uromyces setariaeitalicae
Uromyces striatus
Uromyces striatus
Uromyces strobilanthis
BPI863744
KY575068
BPI863750
--
KY575069
U-675
DAOM 240966
BPI 893253
HQ412651
---
-HQ317512
KY798375
Uromyces tenuicutis
Uromyces trifolii
Uromyces viciae-fabae
Uromyces viciae-fabae
Uromyces vignae
Uromyces vignae
Taphrina pruni
BRIP 60012
---
--AB115665
-AB115731
-MH855700
KX999904
GU936634
-AF426199
-KX999906
MH867219
H92019
BRIP 60213
CBS 358.35
KY764197
References
Blomquist (unpublished)
Liu et al. (2015)
Marin-Felix et al. (2017)
Demers & Castlebury
(unpublished)
Sagliocco et al.
(unpublished)
Marin-Felix et al. (2017)
Liu et al. (2015)
Padamsee & McKenzie
(2017)
Demers et al.
(unpublished)
Demers & Castlebury
(unpublished)
Demers & Castlebury
(unpublished)
Deadman et al. (2011)
Liu et al. (2015)
Demers & Castlebury
(unpublished)
Marin-Felix et al. (2017)
Zuluaga et al. (2011)
Chung et al. (2004)
Maier et al. (2003)
Chung et al. (2004)
Marin-Felix et al. (2017)
Vu et al. 2019)
Genus-wise literature of Indian rust fungi
Angiopsora: Bagchee & Singh 1960, Bahadur & Sinha 1967, Ramakrishnan 1950,
Ramakrishnan & Sundaram 1954b, Thirumalachar & Mundkar 1951, Vaheeduddin 1955. Arthuria:
Gokhle & Patel 1953. Bubakia: Butler & Bisby 1931, Mundkar 1938, 1943, Ramakrishnan &
Ramakrishnan 1950a. Calidion: Ramakrishnan 1950. Chaconia: Butler & Bisby 1931, Mishra et al.
1976, Patel et al. 1949, Patil & Thirumalachar 1971, Patil 1966a, Ramakrishnan 1950, Singh 1966.
Ceropsora: Bakshi & Singh 1960. Cerotelium: Ahmad 1981, Arthur 1917, Bakshi et al. 1972, Butler
1914, Chowdhary 1948, Jain et al. 1966, Joshi & Vashiist 1959, Nagraj et al. 1971, Patil &
Thirumalachar 1971, Patil 1966a, Pawar & Kulkarni 1973, Payak 1949, Ramachar et al. 1978,
Ramakrishnan 1952, Rangaswamy et al. 1968, Sathe 1972b, Sydow & Mitter 1935, Thite & Patil
1970, Vasudeva 1962, Venkatakrishaiya 1958, Wakhloo 1962, Yadav 1963a, Yadav &
Thirumalachar 1955. Chrysocelis: Ramakrishnan & Ramakrishnan 1949. Chrysomyxa: Bakshi &
Singh 1972, Barclay 1890a, Butler 1906, 1910, Dietal 1890, Puri 1955, Sydow & Butler 1901,
Ulbitch 1938. Coleosporium: Anonymous 1950, Arthur 1934, Bagchee 1950b, Bakshi et al. 1972,
Barclay 1890b, Berkeley 1856, Butler & Bisby 1931, Cummins 1943, Dewan & Kar 1974, Fleming
1874, Goswami & Singh 1973, Hafeezkhan 1928b, Kamal et al. 1979, Mitter & Tandon 1932a, b,
Patil & Thirumalachar 1971, Patil 1966a, Prasada 1951, Puri 1955, Sanwal 1951b, Sydow 1922,
Sydow & Mitter 1933, Sydow et al. 1937, Sydow & Butler 1901, Sydow et al. 1907, Sydow et al.
1912, Wani & Thirumalachar 1969, Yadav 1964a, Yadav & Thirumalachar 1955. Corbulopsora:
Thirumalachar 1947. Cronartium: Anonymous 1950, Bagchee 1933, 1950a, b, c, Bakshi et al. 1972,
98
Butler & Bisby 1931, Goswami & Singh 1973, Hafeezkhan 1928a, Puri 1955. Crossopsora: Chavan
& Bakare 1974, Malviya & Jain 1981, Mundkar & Thirumalachar 1945, Mundkar & Thirumalachar
1952, Patil & Thirumalachar 1971, Patil 1966a, Ramakrishnan & Soumini 1946a, Sunderam 1961,
Sydow et al. 1912, Yadav 1963b. Cystopsora: Ramakrishnan & Sundaram 1952a, Rangaswamy et
al. 1968, Sharma 1977. Dasturella: Butler & Bisby 1931, Mishra & Nema 1976, Mundkar &
Kheshwala 1943, Nema & Mishra 1965, Patel et al. 1949, 1951b, Patil & Thirumalachar 1971, Patil
1966a, Rangaswamy et al. 1968, Sathe 1965a, Thirumalachar et al. 1956, Thirumalachar &
Gopalkrishan 1947, Yadav 1964a. Didymopsorella: Rangaswamy et al. 1968, Thirumalachar 1950a,
1951, Thirumalachar & Mundkar 1950. Diorchidium: Sydow et al. 1907, Yadav 1953.
Elateraecium: Thirumalachar et al. 1966. Endophyllum: Arthur 1934, Cummins 1943, Gokhle et al.
1955, Singh & Jalan 1965, Nagraj et al. 1971, Patil 1966a, Thirumalachar & Govindu 1954,
Thirumalachar & Narsimhan 1950b. Gambleola: Butler 1906, Sydow & Butler 1901, Sydow et al.
1907. Goplana: Ramakrishnan & Ramakrishnan 1949. Gymnopuccinia: Ramakrishnan 1951c, Thite
& Patil 1970. Gymnosporangium: Arthur 1934, Arthur & Cummins 1936a, Barclay 1890d, Butler
1906, Cummins 1943, Sydow 1938, Sydow & Butler 1901. Hamaspora: Goswami & Singh 1973,
Massee 1892, Ramakrishnan & Sundaram 1954a. Hapalophragmium: Jain et al. 1966, Mishra 1969,
Ramakrishnan & Ramakrishnan 1948a, Thirumalachar 1950c. Hemileia: Ananth & Chokanna 1961,
Cooke 1876b, Krishnamurthy & Rangaswamy 1947, Massee 1906, Nagraj et al. 1971, Parndekar
1964, Patil & Thirumalachar 1971, Patil 1977, 1966a, Ramakrishnan 1957b, Ramakrishnan &
Soumini 1946b, Rangaswamy et al. 1968, Sydow & Mitter 1933, Sydow & Butler 1901,
Thirumalachar 1947, Thirumalachar 1950b, Thirumalachar & Narsimhan 1947, Thite & Patil 1970,
Yadav 1963b. Hyalopsora: Anonymous 1936, Arthur 1934, Chona & Munjal 1955, Munjal &
Kapoor 1961, Sydow 1938. Kernella: Thirumalachar & Mundkar 1949. Kernkampella: Rajendren
1970, Bhagyanarayana & Ramachar 1985. Kuehneola: Chavan 1975, Malviya & Jain 1981, Patil
1966a, Thite & Patil 1970, Yadav 1963b, Bhagyanarayana & Rao 1995, Hosagoudar 1985.
Kweilingia: Bakshi et al. 1972, Gautam & Avasthi 2018. Leucotelium: Sydow 1939, Sydow et al.
1912, Nema & Mishra 1965. Macabuna: Kamal et al. 1979. Maravalia: Ahmad 1981, Arthur &
Cummins 1936b, Butler & Bisby 1931, Patil & Thirumalachar 1971, Patil 1966a, Thirumalachar
1949a, Yadav & Thirumalachar 1955. Masseeëlla: Joshi & Vashiist 1959, Mundkar & Thirumalachar
1952, Patel et al. 1949, Patil 1966a, Patwardhan 1964, Ramakrishnan 1957a, Sharma 1975,
Thirumalachar 1943b, c. Melampsora: Agarwal et al. 1959, Ajrekar 1912, Bagchee 1950b, Bagchee
& Singh 1960, Bakshi & Singh 1961, Barclay 1891, Butler 1905, 1918, Butler & Bisby 1931, 1960,
Chavan & Bakare 1977, Cooke 1876b, Cummins 1943, Jain et al. 1966, Kala & Gaur 1983a, Kaul
1959a, b, Kern & Thurston 1944, Lele 1952, Manocharachary et al. 1976, Mcrae 1917, Mishra 1969,
1963a, Mishra & Prasad 1966, Pandotra 1966, Pandotra & Ganguly 1964b, Parndekar 1964, Patil &
Thirumalachar 1971, Patil 1966a, Payak 1949, Puri 1955, Ramachar & Bhagyanarayana 1977a, b,
Ramakrishnan & Ramakrishnan 1949, Rangaswamy et al. 1968, Ravindra Nath & Narahari Reddy
1964, Srivastava 1982, Sydow & Butler 1901, Sydow et al. 1907, Sydow et al. 1912, Thirumalachar
1941b, Vasudeva 1948, 1949, 1950a, b. Melampsoridium: Bakshi et al. 1972, Narayan & Kamal
1985, Patil 1966a, Sathe 1966a, Singh & Pandey 1972. Milesina: Anonymous 1959a, Bagchee &
Singh 1960. Monosporidium: Barclay 1890c, Parndekar 1964, Patil 1966a, Sydow et al. 1912,
Gokhle & Patel 1951. Neophysopella: Mundkar 1943, Vaheeduddin 1955, Bagchee & Singh 1960.
Nyssopsora: Mundkar & Thirumalachar 1952, Nagachan & Goswami 1985, Rangaswamy et al. 1968,
Roy 1948. Ochropsora: Arthur & Cummins 1936a. Olivea: Bakshi et al. 1972, Butler & Bisby 1931,
Pandotra & Ganguly 1964b, Patil 1966a. Peridermium: Anonymous 1950, Bakshi & Singh 1972,
Barclay 1890b, Butler & Bisby 1931, Champion 1922, Cooke 1878b, Hafeezkhan 1928a, Mitter &
Tandon 1932a, Puri 1955, Sydow & Butler 1901, Berkeley 1856, Cooke 1877. Peridiopsora: Sathe
1969b. Phakopsora: Bahekar 1966, Butler 1912, Butler & Bisby 1931, Dietal 1890, Jhooty et al.
1977, Kala & Gaur 1983a, Mundkar 1943, Nagraj et al. 1971, Patel et al. 1985, Patil & Thirumalachar
1971, Patil 1977, Ramachar et al. 1978, Ramakrishnan & Subramanian 1952, Ramakrishnan 1951c,
Ramakrishnan 1952, 1955a, 1956, Ramakrishnan & Ramakrishnan 1950a, Ramakrishnan et al.1952,
Ramakrishnan & Sundaram 1954b, 1955a, b, Rangaswamy et al. 1968, Sathe 1972a, Sharma & Jain
99
1981, Sydow 1938, Sydow et al. 1907, Sydow et al. 1912, Thirumalachar 1947, Uppal et al. 1935,
Yadav 1963a, 1964a, Yadav & Thirumalachar 1955. Phragmidiella: Patel et al. 1949, Patil 1966a,
Thirumalachar & Mundkar 1949. Phragmidium: Barclay 1890b, Barclay 1891, Bhattacharya &
Baruah 1953, Cooke 1878a, b, Goswami & Singh 1973, Kala & Gaur 1983a, Mitter & Tandon 1932a,
b, Mundkar 1938, Pandotra & Ganguly 1964b, Ramakrishnan & Sundaram 1953a, Sydow 1938,
Sydow & Mitter 1935, Sydow et al. 1907, Sydow et al. 1911a, b. Physopella: Agarwal et al. 1964,
Khanna & Chandra 1975, Malviya & Jain 1981, Patil & Thirumalachar 1971, Ramachar 1966,
Ramachar & Bhagyanarayana 1976, Sathe 1965d, Subramaniam & Ramakrishnan 1956. Pileolaria:
Sydow 1938, Gautam & Avasthi 2017c. Prospodium: Bagyanarayana & Ravinder 1995,
Bagyanarayana et al. 1998. Puccinia: Agarwal et al. 1959, 1981, Ahmad 1977, Ahmad & Singh
1969, Ahmad et al. 1969, Anonymous 1950, Arthur 1934, Arthur & Cummins 1936a, Bagchee &
Singh 1960, Balasubramanian 1973, Barclay 1890a, b, e, f, 1891, Berkeley 1856, Butler 1905, 1918,
Butler & Bisby 1931, 1960, Butler & Hayman 1906, Chavan 1975, Chavan & Bakare 1974, Chavan
& Bakare 1977, Chavan & Patil 1972, Chona et al. 1958, Chona & Munjal 1950, 1955, 1956, Chona
et al. 1956, Chowdhary 1948, Cooke 1876a, b, Cooke 1878a, Cummins 1943, Cummins 1953, Dalela
1956, Dalela & Sinha 1964, Damle 1943, Darr & Shah 1980, Deoraj 1980, Dietal 1890, Dube 1958,
Fleming 1874, Ganguly & Pandotra 1962, 1963, Gautam & Avasthi 2016b, Gautam & Avasthi
2017a, Gopinathnair 1972, Goswami 1974, Goswami & Singh 1973, Goyal et al. 1971, Gupta 1977,
Gupta & Shukla 1955, Jadhav & Somani 1978a, b, Jain et al. 1966, Joshi & Lele 1984, Joshi &
Merchand 1963, Joshi & Payak 1963, Joshi 1958, Joshi & Vashiist 1959, Kala & Gaur 1983a, b,
Kamal et al. 1979, Kanaujia 1978, Kanaujia & Kishore 1981, Kanadswamy & Vijyalakshmi 1959,
Kaul 1959b, Khanna 1961, Khosla et al. 1975, Khulbe & Verma 1978, Kolte & Awasthi 1979,
Konger & Baruah 1958, Kumar et al. 1975, Malviya & Jain 1981, Mcrae 1917, Mehta 1934, Mishra
et al. 1964, 1965, 1968, Mishra 1969, Mishra & Nema 1976, Mishra et al. 1976, Mishra & Mishra
1975, Mishra 1963b, Mishra & Lele 1963, Mishra & Sharma 1963, 1964, Mishra et al. 1975, Mitter
& Tandon 1930, 1932a, b, 1937, Mundkar 1938, Mundkar & Ahmad 1946, Mundkar &
Thirumalachar 1952, Mutkekar et al. 1968, Munshi 1976, Nema & Agarwal 1960, Nema & Mishra
1965, Padwick 1945a, b, Padwick & Khan 1944, Pandotra 1966, Pandotra & Ganguly 1962, Pandotra
& Ganguly 1964a, b, Pandotra & Sastry 1969a, b, Parndekar 1964, Patel et al. 1949, 1950, Patil &
Thirumalachar 1964, Patil & Thirumalachar 1971, Patil 1966a, Patil & Date 1980, Payak 1949, 1965,
Payak & Khanna 1970, Payak & Mishra 1963, Payak & Renfro 1966, Prasad 1948, Prasada 1951,
Rebenhorst 1878, Sharma & Shankar 1979, Rajendran 1966, Ramachar 1965, Ramachar et al. 1978,
Ramachar & Bhagyanarayana 1976, Ramachar & Cummmins 1965, Ramakrishna & Subbayya 1973,
Ramakrishnan 1950, Ramakrishnan 1952, 1955b, 1956, Ramakrishnan & Narasimhalu 1941,
Ramakrishnan & Ramakrishnan 1946, Ramakrishnan & Ramakrishnan 1947, 1948a, b,
Ramakrishnan & Ramakrishnan 1949, 1950b, c, Ramakrishnan et al. 1952a, b, Ramakrishnan &
Sundaram 1952b, 1953b, 1954a, b, 1955a, 1956a, b, Rangaswamy et al. 1968, Roy 1964, 1968, Roy
& Gupta 1959, Sachan et al. 1980, Sahni & Chona 1965, Salam & Ramachar 1955, Sathe 1965b,
1969a, 1971, Shanmugam et al. 1972, Sharma & Mukerji 1972, Sharma 1975, Sharma et al. 1979,
Sharma & Singh 1964, Sharma 1957, Sharma et al. 1973, Shinde & More 1975, Shukla & Singh
1976, Siddiqui 1971, 1972, 1973, Singh & Kamal 1985, Singh & Sharma 1977, Singh 1962, Mishra
et al. 1965, Mishra 1969, Sinha & Kapooria 1966, Sohi et al. 1967, Somani 1979, Soumini 1949,
Srivastava 1979a, c, 1980, 1982, Subramaniam & Ramakrishnan 1956, Sunderam 1956, 1963,
Sunderam et al. 1966, Sydow 1913, 1914, 1922, 1938, 1939, Sydow & Mitter 1935, Sydow et al.
1937, Sydow & Butler 1901, Sydow et al. 1907, Sydow et al. 1911a, b, Sydow et al. 1912,
Thirumalachar 1941a, 1945, Thirumalachar 1947, 1949a, b, Thirumalachar & Mundkar 1951,
Thirumalachar et al. 1943, Tilak & Rao 1968, Ulbitch 1938, Unni & Philip 1974, Uppal et al. 1935,
Vasudeva 1948, 1949, 1950a, b, 1957, 1958, Wakhloo 1962, Yadav 1953, 1963b, 1964a, Yadav &
Yadav 1965, Yadav et al. 1975. Pucciniastrum: Bakshi et al. 1972, Patil 1966a. Pucciniosira: Arthur
1934, Cummins 1943. Pucciniostele: Barclay 1891, Sydow et al. 1912. Ramakrishnania: Ramachar
& Bhagyanarayana 1979. Ravenelia: Berkeley 1839, Butler & Bisby 1931, Chavan & Bakare 1974,
Cooke 1880a, b, Jain et al. 1966, Kapoor & Agarwal 1972, 1974, Mishra 1969, Mishra et al. 1976,
100
Mundkar & Prasad 1938, Narasimhan & Thirumalachar 1961, Patil & Thirumalachar 1971, Patil
1977, Patil & Thite 1978, Patil 1966a, b, Patil & Date 1977, Pavgi & Singh 1969, Petch 1912,
Ramakrishnan 1952, 1957a, Ramakrishnan & Ramakrishnan 1948b, Ramakrishnan & Sundaram
1952b, Rangaswamy et al. 1968, Roy 1964, Salam & Ramachar 1956, Sanwal 1951a, Sathe 1965c,
Siddiqui 1957, Sydow 1913, Sydow & Mitter 1933, Sydow & Butler 1901, Sydow et al. 1907, Sydow
et al. 1912, Tyagi & Prasad 1972, 1978, Yadav 1963b, Yadav & Thirumalachar 1955. Roestelia:
Barclay 1891, Arthur & Cummins 1936a, Mitter & Tandon 1932a, 1937, Sydow et al. 1907. Scopella:
Butler & Bisby 1931, Cummins1950, Mundkar & Thirumalachar 1952, Narayan & Kamal 1985,
Parndekar 1964, Patil & Thirumalachar 1971, Payak 1949, Solanki et al. 1985, Thirumalachar 1950c,
Thirumalachar & Mundkar 1951. Skierka: Chavan 1968, Gautam & Avasthi 2017c.
Sphaerophragmium: Cooke 1880a. Stakmania: Sathe 1966c. Trachyspora: Arthur 1934, Padwick
& Merh 1943. Tranzschelia: Patel et al. 1949, Waraitch & Khatri 1976, 1977. Trochodium: Gharse
1944, Patil & Thirumalachar 1971, Patil 1966a, Thirumalachar 1942a. Uredinopsis: Anonymous
1936, Munjal & Kapoor 1961, Patil 1966a, Puri 1955, Ramakrishnan & Ramakrishnan 1950b.
Uredopeltis: Ramachar et al. 1978, Sathe 1965e. Uromyces: Agarwal et al. 1959, Ahmad 1941,
Ajrekar 1912, Ajrekar & Tonapy 1923, Anonymous 1950, 1954, Arthur 1934, Arthur & Cummins
1936a, Asthana 1952, Bahadur & Sinha 1967, Barclay 1890a, 1891, Basuchaudhary & Singh 1971,
Behera & Mukerji 1974, Butler & Bisby 1931, 1960, Chaudhari 1958, Chavan 1975, Chavan &
Bakare 1973a, b, Chavan & Bakare 1974, Chavan & Bakare 1977, Cooke 1876a, Cummins 1943,
Dietal 1890, Dube 1958, Dube et al. 1979, Dublish & Singh 1977, Gautam & Avasthi 2017b,
Gerdemann & Bakshi 1976, Jain et al. 1966, Joshi & Reddy 1958, Joshi & Reddy 1959, Joshi 1958,
Kapooria & Sinha 1966, Manocharachary 1975, Mathur 1967, Mathur & Singh 1964, Mishra 1969,
Mishra & Khare 1969, Mishra & Nema 1976, Mishra et al. 1976, Mitter & Tandon 1930, 1932b,
More & Moniz 1964, Nagachan & Verma 1984, Narasimhan & Thirumalachar 1964, Nema &
Agarwal 1960, Nema & Mishra 1965, Padwick & Khan 1944, Padwick & Merh 1943, Pandotra 1966,
Pandotra & Sastry 1969a, Parndekar 1964, Patel et al. 1949, Patil & Thirumalachar 1968, Patil &
Thirumalachar 1962, Patil & Thirumalachar 1971, Patil 1966a, Patil & Date 1980, Pavgi & Upadhyay
1966, Payak 1949, 1953, 1962, Ponappa 1969, Prasad et al. 1962, Rebenhorst 1878, Ramachar et al.
1978, Ramakrishnan & Subramanian 1952, Ramakrishnan 1951b, Ramakrishnan 1952,
Ramakrishnan & Ramakrishnan 1948b, Ramakrishnan & Rangaswamy 1948, Ramakrishnan &
Shrinivasan 1950, Ramakrishnan et al. 1952a, b, Ramakrishnan & Sundaram 1955a, Rangaswamy et
al. 1968, Rolla & Addala 1963, Sachan et al. 1980, Saini & Chand 1984, Saksena 1956, 1930, Sokhi
& Sohi 1976, Srivastava 1979b, Sunderam 1964, Sydow 1913, 1922, 1938, Sydow & Mitter 1935,
Sydow et al. 1937, Sydow & Butler 1901, Sydow et al. 1907, Sydow et al. 1912, Thite & Patil 1970,
Ulbitch 1938, Uppal et al. 1935, Vasudeva 1948, 1949, 1950a, b, Yadav 1963b, Yadav &
Thirumalachar 1955. Xenostele: Ramakrishnan 1951a. Zaghouania: Butler & Bisby 1931.
Pucciniales genera incertae sedis
Aecidium: Ajrekar 1936, Arthur & Cummins 1936a, Bagchee & Singh 1960, Barclay 1890a,
b, 1891, Bhagat & Kelkar 1974, Butler & Bisby 1931, 1960, Chavan & Bakare 1973a, b, 1974,
Chavan & Patil 1972, Cooke 1878a, Cummins 1943, Goswami 1972, Goswami & Bhattacharjee
1973, Gupta & Gupta1985, Hennings 1900, Hosagaudar 1984, Kamal & Singh 1981, Kamal et al.
1979, Mishra et al. 1976, Mitter & Tandon 1932a, b, Mundkar 1938, Mundkar & Thirumalachar
1952, Pandotra & Ganguly 1964b, Pandotra & Sastry 1969b, Patel et al. 1949, Patil & Thirumalachar
1971, Prasad et al. 1962, Ramachar 1956, Ramakrishnan 1959, 1960, Ramakrishnan & Ramakrishnan
1946, 1948a, 1949, Ramakrishnan et al. 1952a, b, Ramakrishnan & Sundaram 1953a, b, Rangaswamy
et al. 1968, Salam & Ramachar 1955, 1956, Sathe 1965c, 1966b, Singh & Kamal 1985, Sunderam &
Rao 1950, Sydow 1913, Sydow 1914, Sydow & Mitter 1933, Sydow & Mitter 1935, Sydow & Sydow
1904a, 1911, 1912a, b, Sydow 1921, Sydow & Sydow 1917, Sydow & Butler 1901, Sydow et al.
1907, Sydow et al. 1912, Thirumalachar 1947, Thirumalachar 1950c, Thirumalachar & Narsimhan
1950a, Thite & Patil 1970, Ulbitch 1938, Uppal et al. 1935, Yadav 19563b, Yadav & Saran 1985.
Phragmotelium: Thirumalachar 1942b, Thirumalachar et al. 1943. Tunicopsora Suj. Singh & P.C.
101
Pandey: Bakshi et al. 1972. Uraecium: Ramakrishnan 1965, Ramakrishnan & Ramakrishnan 1948b.
Uredo: Anonymous 1959b, Bagchee & Singh 1960, Barclay 1890b, 1891, Berkeley 1839, Butler &
Bisby 1931, 1960, Chavan 1975, Chavan & Bakare 1973a, b, 1974, Chavan & Bhambure 1975,
Cooke 1876b, 1879, Kamal et al. 1979, Kanaujia & Kishore 1981, Kapoor & Agarwal 1974, Laundon
& Ponappa 1966, Nagraj et al. 1971, Padwick 1945a, b, Padwick & Khan 1944, Pandotra 1966, Patel
et al. 1951a, Patil 1977, Patil & Thite 1978, Rahalkar 1977, Ramachar et al. 1978, Ramakrishnan
1951a, 1952, 1956, 1957a, Ramakrishnan & Ramakrishnan 1948a, Ramakrishnan & Ramakrishnan
1949, Ramakrishnan et al. 1952a, Ramakrishnan & Sundaram 1953b, Rangaswamy et al. 1968, Sathe
& Rahalkar 1976, Sunderam 1961, Sydow & Mitter 1935, Sydow et al. 1937, Sydow & Sydow
1904b, 1924, Sydow & Butler 1901, Sydow et al. 1912, Thirumalachar 1947, Thirumalachar 1950c,
Vaheeduddin 1955, Yadav 1964b.
Results
Accepted taxa of rust fungi of Basidiomycota up to genus are summarized in Table 1 while, the
species of each genus are mentioned in detailed outline of the fungi reported from India. The rust
fungi of India comprised of 640 species and 69 genera belonging to 16 families. Highest numbers of
species were reported in Pucciniaceae (393) followed by Raveneliaceae (61), Phakopsoraceae (50),
Coleosporiaceae (32), Phragmidiaceae (27), Pucciniastraceae (19), Melampsoraceae (18),
Crossopsoraceae (14), Zaghouaniaceae (13), Gymnosporangiaceae (7), Milesinaceae (5),
Skierkaceae (3), Tranzscheliaceae (3), Pileolariaceae (2), Ochropsoraceae (1),
Sphaerophragmiaceae (1). Similarly, when comparing the rust genera, highest number of species of
rust fungi was found Puccinia (279), followed by Uromyces (89), Ravenelia (33), Phakospora (25),
Coleosporium (19), Phragmidium (18), Melampsora (17) and Maravalia (11). Many taxa of rust
fungi with uncertain taxonomic position are placed in incertae sedis. Similarly, the generic names
have been transferred to new genera, but either their types or records from India still need to be
revised. Such genera and their species are discussed in notes of Indian rust genera section of this
manuscript.
Outline of rust fungi reported from India
The information presented in the outline is arranged as phylum followed by subphylum, class,
order, family, genus and species.
Taxonomy
Basidiomycota R.T. Moore
Pucciniomycotina R. Bauer, Begerow, J.P. Samp., M. Weiss & Oberw.
Pucciniomycetes R. Bauer, Begerow, J.P. Samp., M. Weiss & Oberw.
Pucciniales Clem. & Shear
1. Coleosporiaceae Dietel, In: Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 548. 1900.
emend. Aime & McTaggart
Coleosporium Lév. (19)
Coleosporium asterum (Dietel) Syd. & P. Syd.
Coleosporium barclayense Bagchee
Coleosporium bletiae Dietel
Coleosporium campanulae (Pers.) Tul.
Coleosporium clematidis Barclay
Coleosporium datiscae Tranzschel
Coleosporium inulae Rabenh.
Coleosporium ipomoeae (Schwein.) Burrill
Coleosporium himalayense Durrieu
Coleosporium leptodermidis (Barclay) P. Syd. & Syd.
102
Coleosporium mitteri Syd.
Coleosporium myriactidis Syd.
Coleosporium oldenlandiae E.J. Butler
Coleosporium perillae P. Syd.
Coleosporium plectranthi Barclay
Coleosporium satyrii Mundk. & Thirum.
Coleosporium senecionis (Pers.) Fr.
Coleosporium sidae Sanwal
Coleosporium xanthoxyli Dietel & P. Syd. (1898)
Chrysomyxa Unger (7)
Chrysomyxa deformans (Dietel) Jacz.
Chrysomyxa dietelii Syd. & P. Syd.
Chrysomyxa himalensis Barclay
Chrysomyxa piceae Barclay
Chrysomyxa pirolae (DC.) Rostr.
Chrysomyxa vitis E.J. Butler
Chrysomyxa himalayensis Singh, Khan & Mishra
Cronartium Fr. (4)
Cronartium fici T.S. Ramakr. & K. Ramakr.
Cronartium himalayense Bagchee
Cronartium quercuum (Berk.) Miyabe ex Shirai
Cronartium ribicola J.C. Fisch.
Goplana Racib. (1)
Goplana indica T.S. Ramakr. & K. Ramakr.
Stakmania Kamat & Sathe (1)
Stakmania indica Kamat & Sathe
2. Crossopsoraceae Aime & McTaggart, In: Fungal Systematics and Evolution 7: 21–47. 2020
Angiopsora Mains (2)
Angiopsora cyrtococci T.S. Ramakr. & Sundaram
Angiopsora apoda (Har. & Pat.) Aime & McTaggart
Crossopsora Syd. & P. Syd. (3)
Crossopsora premnae (Petch) Syd. & P. Syd.
Crossopsora premnae-tomentosae T.S. Ramakr. & Soumini
Crossopsora symphorematis Sundaram
Crossopsora ziziphi (Syd., P. Syd. & E.J. Butler) Syd. & P. Syd.
Dasturella Mundk. & Khesw. (3)
Dasturella bambusina Mundk. & Khesw.
Dasturella boswelliae Patel, Payak & N.B. Kulk.
Dasturella oxytenantherae Sathe
Kweilingia Teng (2)
Kweilingia bagchii (Suj. Singh & P.C. Pandey) Buriticá
Kweilingia divina (Syd.) Buriticá
Neophysopella Jing X. Ji & Kakish. (3)
Neophysopella ampelopsidis (Dietel & P. Syd.) Jing X. Ji & Kakish.
Neophysopella meliosmae (Kusano) Jing X. Ji & Kakish.
Neophysopella meliosmae-myrianthae (Henn. & Shirai) Jing X. Ji & Kakish.
Physopella (2)
Physopella artocarpi (Berk. & Broome) Arthur
Physopella vernoniae (T.S. Ramakr.) Ramachar & Bhagyan.
103
3. Gymnosporangiaceae P. Zhou & L. Cai, Persoonia 45: 79. 2020. emend. Aime & McTaggart
Peridiopsora Kamat & Sathe (1)
Peridiopsora adelocaryi Kamat & Sathe
Gymnosporangium R. Hedw. ex DC. (3)
Gymnosporangium clavariiforme (Wulfen) DC.
Gymnosporangium confusum Plowr.
Gymnosporangium cunninghamianum Barclay
Roestelia Rebent. (3)
Roestelia distorta (Arthur & Cummins) F. Kern
Roestelia cunninghamianum (Barclay) F. Kern
Roestelia patula (Syd. & P. Syd.) F. Kern
4. Melampsoraceae Dietel, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 38. 1897
Ceropsora B.K. Bakshi & Suj. Singh (1)
Ceropsora piceae (Barclay) B.K. Bakshi & Suj. Singh
Melampsora Castagne (17)
Melampsora caprearum Thüm.
Melampsora ciliata Barclay
Melampsora damnosa (Sacc.) Lind.
Melampsora epitea Thüm.
Melampsora eucalypti Rabenh.
Melampsora euphorbiae (Ficinus & C. Schub.) Castagne
Melampsora euphorbiae-geniculatae F. Kern & Thurst.
Melampsora geniculatae Ramachar & Bhagyan.
Melampsora hypericorum (DC.) J. Schröt.
Melampsora lini (Ehrenb.) Lév.
Melampsora mundkurii Thirum.
Melampsora oblonga Bagchee
Melampsora populnea (Pers.) P. Karst.
Melampsora salicis-albae Kleb.
Melampsora salicis-wallichianae Ulbr.
Melampsora sancti-johannis Barclay
Melampsora stereospermi T.S. Ramakr. & K. Ramakr.
Melampsora yoshinagai Henn.
Melampsora caprearum Thüm.
5. Milesinaceae Aime & McTaggart, In: Fungal Systematics and Evolution 7: 21–47. 2020
Milesina Magnus (3)
Milesina coniogrammes Hirats. f.
Milesina exigua Faull
Milesina polypodii (F.B. White) Aime & Rossman
Uredinopsis Magnus (2)
Uredinopsis macrosperma (Cooke) Magnus
Uredinopsis syngrammes Munjal & J.N. Kapoor
6. Ochropsoraceae (Arthur) Aime & McTaggart, In: Fungal Systematics and Evolution 7: 21–47.
2020
Ochropsora Dietel (1)
Ochropsora ariae (Fuckel) Ramsb.
7. Phakopsoraceae Cummins & Y. Hirats., Illustr. Gen. Rust Fungi, rev. Edn (St. Paul): 13. 1983.
emend. Aime & McTaggart
104
Arthuria Jackson (2)
Arthuria glochidii Gokhale, Patel & Thirum.
Arthuria tylophorae T.S. Ramakr.
Bubakia Arthur (1)
Bubakia indica T.S. Ramakr. & K. Ramakr.
Cerotelium Arthur (7)
Cerotelium bauhiniae Thirum. & Yadav
Cerotelium fici (Castagne) Arthur
Cerotelium kirganeliae Thirum. & Yadav
Cerotelium peregrinum (P. Syd. & Syd. & E.J. Butler) Arthur
Cerotelium terminaliae-paniculatae Nag Raj, Govindu & Thirum.
Cerotelium trichosanthis (Petch) Nag Raj, Govindu & Thirum.
Cerotelium wagateae Thirum. & Gopalkr.
Macabuna Buriticá & J.F. Hennen (1)
Macabuna ziziphi (Pat.) Buriticá & J.F. Hennen
Monosporidium Barclay (3)
Monosporidium andrachnes Barclay ex Sacc.
Monosporidium euphorbiae Barclay ex Sacc.
Monosporidium pavettae (Gokhale & Patel) Buriticá
Masseeella Dietel (6)
Masseeella breyniae Thirum.
Masseeella capparis (Hobson bis ex Cooke) Dietel
Masseeella flueggeae Syd.
Masseeella narasimhanii Thirum.
Masseeella putranjivae T.S. Ramakr.
Masseeella terminaliae Patw.
Phakopsora Dietel (25)
Phakopsora apludae M.S. Patil
Phakopsora artemisiae Hirats. f.
Phakopsora caseariae Yadav
Phakopsora chorisandrae T.S. Ramakr. & G.S. Reddy
Phakopsora cingens (Syd. & P. Syd.) Hirats.
Phakopsora cronartiiformis Dietel
Phakopsora desmium (Berk. & Broome) Cummins
Phakopsora elephantopodis Hirats
Phakopsora elettariae (Racib.) Cummins
Phakopsora erythrinae Gäum.
Phakopsora fici-elasticae T.S. Ramakr.
Phakopsora formosana Syd. & P. Syd.
Phakopsora incompleta (Syd. & P. Syd.) Cummins
Phakopsora kirganeliae T.S. Ramakr. & K. Ramakr.
Phakopsora mangalorica T.S. Ramakr. & Sundaram
Phakopsora meibomiae (Arthur) Arthur
Phakopsora pachyrhizi Syd. & P. Syd.
Phakopsora odinae Mundk.
Phakopsora pachyrhizi Syd. & P. Syd.
Phakopsora parasnathii Yadav & Thirum.
Phakopsora phyllanthi Dietel
Phakopsora punctiformis (Barclay & Dietel) Dietel
Phakopsora sterculiae Nag Raj, Govindu & Thirum.
Phakopsora zingiberis T.S. Ramakr
Phakopsora ziziphi-vulgaris Dietel
105
Phragmidiella Henn. (3)
Phragmidiella aliena (Syd., P. Syd. & E.J. Butler) Buriticá & J.F. Hennen
Phragmidiella heterophragmatis (Mundk. & Thirum.) Thirum. & Mundk.
Phragmidiella holwayi (H.S. Jacks.) Buriticá
Pucciniostele Tranzschel & K.L. Kom. (1)
Pucciniostele clarkiana (Barclay) Tranzschel & K.L. Kom.
Uredopeltis Henn. (1)
Uredopeltis chevalieri J. Walker & R.G. Shivas
8. Phragmidiaceae Corda Icon. fung. (Prague) 1: 6. 1837
Hamaspora Körn. (2)
Hamaspora longissima (Thüm.) Körn.
Hamaspora rubi-sieboldii (Kawagoe) Dietel
Kuehneola Magnus (6)
Kuehneola grewiae (Mundk. & Thirum.) Thirum.
Kuehneola loeseneriana (Henn.) H.S. Jacks. & Holw.
Kuehneola flacourtiae (Mundk. & Thirum.) Thirum.
Kuehneola ramacharii Bagyan. & K.N. Rao
Kuehneola spondiadis Hosag.
Kuehneola ziziphi (T.S. Ramakr. & Subram.) Thirum.
Phragmidium Link (18)
Phragmidium assamense Syd. & P. Syd.
Phragmidium barclayi Dietel
Phragmidium brevipedicellatum Hirats. f.
Phragmidium bulbosum (Fr.) Schltdl.
Phragmidium butleri Syd. & P. Syd.
Phragmidium egenulum Syd., P. Syd. & E.J. Butler
Phragmidium fragariae (Rabenh.) Ces.
Phragmidium kamtschatkae (H.W. Anderson) Arthur & Cummins
Phragmidium mucronatum (Pers.) Schltdl.
Phragmidium orientale Syd. & P. Syd.
Phragmidium potentillae (Pers.) P. Karst.
Phragmidium rosae-moschatae Dietel
Phragmidium incompletum Barclay
Phragmidium laceianum Barclay
Phragmidium malvacearum Bert.
Phragmidium nepalense Barclay
Phragmidium octoloculare Barclay
Phragmidium quinqueloculare Barclay
Trachyspora Fuckel (1)
Trachyspora alchemillae (Pers.) Fuckel
9. Pileolariaceae (Arthur) Cummins & Y. Hirats., llustr. Gen. Rust Fungi, rev. Edn (St. Paul): 14.
1983. emend. Aime & McTaggart
Pileolaria Castagne (2)
Pileolaria indica Syd.
Pileolaria pistaciae F.L. Tai & C.T. Wei
10. Pucciniastraceae Gäum. ex Leppik, Ann. bot. fenn. 9: 139. 1972. emend. Aime & McTaggart
Hyalopsora Magnus (2)
Hyalopsora orientalis Chona & Munjal
Hyalopsora polypodii (Pers.) Magnus
106
Melampsoridium Kleb. (4)
Melampsoridium betulinum (Pers.) Kleb.
Melampsoridium hiratsukanum S. Ito ex Hirats. f.
Melampsoridium indicum Sathe
Melampsoridium inerme Suj. Singh & P.C. Pandey
Pucciniastrum G.H. Otth (6)
Pucciniastrum aceris Syd.
Pucciniastrum agrimoniae (Dietel) Tranzschel
Pucciniastrum celastri Syd. & P. Syd.
Pucciniastrum coriariae Dietel
Pucciniastrum coryli Kom.
Pucciniastrum gaultheriae Syd. & P. Syd.
Peridermium (Link) J.C. Schmidt & Kunze (7)
Peridermium brevius Barclay
Peridermium cedri Barclay
Peridermium ephedrae Cooke
Peridermium himalayense Bagchee
Peridermium orientale Cooke
Peridermium piceae Barclay
Peridermium thomsonii Berk.
11. Pucciniaceae Chevall., Fl. gén. env. Paris (Paris) 1: 413. 1826. emend. Aime & McTaggart
Caeoma Link (4)
Caeoma himalayense Suj. Singh, S.N. Khan & B.M. Misra
Caeoma scopariae K.N. Rao
Caeoma euphorbiae-geniculatae Ramachar & Bhagyan.
Caeoma indicum Rajendren
Chrysocelis Lagerh. & Dietel (1)
Chrysocelis butleri (Dietel, Syd. & P. Syd.) G.F. Laundon
Corbulopsora Cummins (1)
Corbulopsora cumminsii Thirum.
Endophyllum Lév. (8)
Endophyllum cassiae Nag Raj, Govindu & Thirum.
Endophyllum cassiae (Bres.) F. Stevens & Mendiola
Endophyllum elaeagni-latifoliae (Petch) Gokhale, Thirum. & Patel
Endophyllum emiliae-sonchifoliae Nag Raj, Govindu & Thirum.
Endophyllum heliotropii Thirum. & Naras.
Endophyllum kaernbachii (Henn.) F. Stevens & Mendiola
Endophyllum maheshwarii Hard. Singh & Jalan
Endophyllum macowanianum (Thüm.) Pole-Evans
Endophyllum spilanthis Thirum. & Govindu
Gambleola Massee (1)
Gambleola cornuta Massee
Hapalophragmium Syd. & P. Syd. (4)
Hapalophragmium anamalaiense T.S. Ramakr. & K. Ramakr.
Hapalophragmium mysorense Thirum.
Hapalophragmium ponderosum Syd., P. Syd. & E.J. Butler
Hapalophragmium tandonii Mitter
Kernella Thirum. (1)
Kernella lauricola (Thirum.) Thirum.
Puccinia Pers. (279)
Puccinia acanthospermi Henn.
107
Puccinia acrophila Peck
Puccinia actaeae-agropyri E. Fisch.
Puccinia adjuncta Mitter
Puccinia aggregata Syd. & P. Syd.
Puccinia agrostidis Plowr.
Puccinia ahmadiana Syd.
Puccinia ainsliaeae P. Syd. & Syd.
Puccinia altii Rud.
Puccinia amphilophidis Doidge
Puccinia angelicae (Schumach.) Fuckel
Puccinia anodae P. Syd. & Syd.
Puccinia antirrhini Dietel & Holw.
Puccinia apii Desm.
Puccinia apludae Syd. & P. Syd.
Puccinia arachidis Speg.
Puccinia arenariae (Schumach.) J. Schröt.
Puccinia argentata (Schultz) G. Winter
Puccinia aristidae Tracy
Puccinia aristidicola Henn.
Puccinia arthraxonis-ciliaris Cummins
Puccinia arundinellae Barclay
Puccinia asterum (Schwein.) F. Kern
Puccinia atropuncta Peck & Clinton
Puccinia azanzae Yadav
Puccinia baradensis P.B. Chavan & U.V. Kulk.
Puccinia barbeyi (Roum.) Magnus
Puccinia behenis G.H. Otth
Puccinia belamcandae Dietel
Puccinia bellurensis Thirum.
Puccinia betae-bengalensis Mundk. & Thirum.
Puccinia bistortae (F. Strauss) DC.
Puccinia blepharidis Henn.
Puccinia bottomleyae Doidge
Puccinia brachypodii G.H. Otth
Puccinia bulbostylidis Doidge
Puccinia bupleuri F. Rudolphi
Puccinia butleri Syd. & P. Syd.
Puccinia cacao McAlpine
Puccinia caheunsis Ell. & Ev.
Puccinia calcitrapae var. filicinae Barclay
Puccinia calcitrapae DC.
Puccinia calosperma Syd., P. Syd. & E.J. Butler
Puccinia calthae Link
Puccinia caricis-filicinae Barclay
Puccinia caricis-nubigenae Padwick & A. Khan
Puccinia cenchri Dietel & Holw.
Puccinia cephalandrae-indicae Syd. & P. Syd.
Puccinia chaerophylli Purton
Puccinia chloridis-incompletae T.S. Ramakr., Sriniv. & Sundaram
Puccinia chrysanthemi Roze
Puccinia circaeae Pers.
Puccinia citrina P. Syd. & Syd.
108
Puccinia citrulli Syd., P. Syd. & E.J. Butler
Puccinia citrullina Raghun. & K. Ramakr. ex Hosag. & Raghun.
Puccinia collettiana Barclay
Puccinia conclusa Thüm.
Puccinia congesta Berk. & Broome
Puccinia coronata Corda
Puccinia courtoisiae (Syd. & P. Syd.) Syd.
Puccinia crepidis-japonicae (Lindr.) Dietel
Puccinia crepidis-sibiricae Lindr.
Puccinia cressae Lagerh.
Puccinia ctenolepidis Ramachar & Bagyan.
Puccinia curculiginis Racib.
Puccinia curcumae T.S. Ramakr. & Sundaram
Puccinia cynodontis Lacroix ex Desm.
Puccinia cyperi Arthur
Puccinia cyperi-tagetiformis (Henn.) F. Kern
Puccinia dactylidina Bubák
Puccinia deodikarii K.R.G. Nair
Puccinia desertorum Syd. & P. Syd.
Puccinia digitariae Ramachar & George
Puccinia digitariae-biformis P.B. Chavan & Hosag.
Puccinia digitariae-vestitae Ramachar & Cummins
Puccinia dioicae Magnus
Puccinia dioscoreae Kom.
Puccinia dovrensis A. Blytt
Puccinia drabae F. Rudolphi
Puccinia droogensis E.J. Butler
Puccinia abutilonis Berk. & Broome
Puccinia aristidae var. chaetariae Cummins & S.M. Husain
Puccinia bulbocastani (A. Cumino) Fuckel
Puccinia canaliculata (Schwein.) Lagerh.
Puccinia centaureae H. Mart.
Puccinia chrysopogoni Barclay
Puccinia coronata f.sp. avenae P. Syd. & Syd.
Puccinia dissiliens Cooke
Puccinia duthiei Ellis & Tracy
Puccinia ellisii De Toni
Puccinia elytrariae Henn.
Puccinia engleriana Henn.
Puccinia enteropogonis P. Syd. & Syd.
Puccinia eragrostidis Petch
Puccinia eremuri Kom.
Puccinia erianthi Padwick & A. Khan
Puccinia eulaliae Barclay
Puccinia eutela Syd.
Puccinia exhauriens Thüm.
Puccinia expallens Syd. & P. Syd.
Puccinia echinopis DC.
Puccinia excelsa Barclay
Puccinia fagopyri Barclay
Puccinia fagopyricola Jørst.
Puccinia ferruginosa P. Syd. & Syd.
109
Puccinia festucae Plowr.
Puccinia fimbristylidis Arthur
Puccinia fimbristylidis-ferrugineae Ramachar, Bhagyan. & A. Kumar
Puccinia flaccida Berk. & Broome
Puccinia flavipes Syd. & P. Syd.
Puccinia fuirenicola Arthur
Puccinia fusca G. Winter
Puccinia garnotiae T.S. Ramakr. & Sundaram
Puccinia gentianae (F. Strauss) Link
Puccinia geranii-silvatici P. Karst.
Puccinia gerberae Pole-Evans
Puccinia gouaniae Holw.
Puccinia gracilenta Syd., P. Syd. & E.J. Butler
Puccinia graminis f. avenae Erikss. & Henning
Puccinia graminis Pers.
Puccinia graminis f. agropyri P.R. Mehta & R. Prasad
Puccinia graminis f. poae Erikss. & Henning
Puccinia graminis f. tritici Erikss. & Henning
Puccinia gymnopetali-wightii T.S. Ramakr., Sriniv. & Sundaram
Puccinia helianthi Schwein.
Puccinia heraclei Grev.
Puccinia heracleicola Cummins
Puccinia herqueri
Puccinia heterospora Berk. & M.A. Curtis
Puccinia heucherae (Schwein.) Dietel
Puccinia hieracii (Röhl.) H. Mart.
Puccinia himachalensis A.K. Gautam & S. Avasthi
Puccinia holboelliae-latifoliae Cummins
Puccinia hookeri P. Syd. & Syd.
Puccinia hordei G.H. Otth
Puccinia hyderabadensis Bagyan. & Ravinder
Puccinia hydrocotyles (Mont.) Cooke
Puccinia hypoxidis McAlpine
Puccinia inayatii Syd. & P. Syd.
Puccinia insidiosa Berk.
Puccinia intermixta Peck
Puccinia invenusta Syd. & P. Syd.
Puccinia investita Schwein.
Puccinia iridis Wallr.
Puccinia isachnes Petch
Puccinia jagopyri Barclay
Puccinia jasminicola T.S. Ramakr. & K. Ramakr.
Puccinia joerstadii S. Ahmad
Puccinia kalchbrenneri De Toni
Puccinia kenmorensis Cummins
Puccinia kraussiana Cooke
Puccinia kuehnii (W. Krüger) E.J. Butler
Puccinia kunthiana T.S. Ramakr., Sriniv. & Sundaram
Puccinia lantanae Farl.
Puccinia lateripes Berk. & Ravenel
Puccinia lateritia Berk. & M.A. Curtis
Puccinia launaeae Maire
110
Puccinia leiocarpa Thirum.
Puccinia leonotidicola Henn.
Puccinia leucadis P. Syd. & Syd.
Puccinia leucophaea Syd., P. Syd. & E.J. Butler
Puccinia leveillei Mont.
Puccinia levis (Sacc. & Bizz.) Magnus
Puccinia libani Magnus
Puccinia liberta F. Kern
Puccinia ligustici Ellis & Everh.
Puccinia linkii Klotzsch
Puccinia lithospermi Ellis & Kellerm.
Puccinia longirostris Kom
Puccinia luculenta (Syd. & P. Syd.) T.S. Ramakr. & K. Ramakr.
Puccinia macrorhynchi Rabenh.
Puccinia malvacearum Bertero ex Mont.
Puccinia melanocephala Syd. & P. Syd.
Puccinia melasmioides Tranzschel
Puccinia menthae Pers.
Puccinia merrillii Henn.
Puccinia microspora Dietel
Puccinia minutissima Arthur
Puccinia monticola Kom.
Puccinia mysorensis Syd.
Puccinia melothriicola Syd. & P. Syd.
Puccinia nakanishikii Dietel
Puccinia nepalensis Barclay & Dietel
Puccinia neyraudiae Syd. & P. Syd.
Puccinia nitida Barclay
Puccinia oahuensis Ellis & Everh.
Puccinia obscura J. Schröt.
Puccinia ocimi Doidge
Puccinia oligocarpa Syd., P. Syd. & E.J. Butler
Puccinia operculinae T.S. Ramakr. & Sundaram
Puccinia operta Mundk. & Thirum.
Puccinia opizii Bubák
Puccinia oplismeni Syd. & P. Syd.
Puccinia oreogeta Syd.
Puccinia oryzopsidis Syd., P. Syd. & E.J. Butler
Puccinia ottochloae T.S. Ramakr.
Puccinia oxalidis Dietel & Ellis
Puccinia pachypes Syd. & P. Syd.
Puccinia pacifica Blasdale ex Arthur
Puccinia padwickii Cummins
Puccinia panici-montani Fujik. ex Ramachar & Cummins
Puccinia paspali Tracy & Earle
Puccinia pectiniformis T.S. Ramakr., Sriniv. & Sundaram
Puccinia peradeniyae Demers & Castl.
Puccinia peraffinis Syd. & P. Syd.
Puccinia phragmitis (Schumach.) Tul.
Puccinia phyllocladiae Cooke
Puccinia phyllostachydis Kusano
Puccinia pieridiz Hazel.
111
Puccinia pimpinellae (F. Strauss) Link
Puccinia plicata Kom.
Puccinia pogonatheri Petch
Puccinia polliniae Barclay
Puccinia polliniae-quadrinervis Dietel
Puccinia polygoni-amphibii Pers.
Puccinia polygoni-weyrichii Miyabe
Puccinia polysora Underw.
Puccinia porri (Sowerby) G. Winter
Puccinia praecox Bubák
Puccinia prainiana Barclay
Puccinia prenanthis-purpureae (DC.) Lindr.
Puccinia princeps Syd. & P. Syd.
Puccinia propinqua Syd., P. Syd. & E.J. Butler
Puccinia prostii Moug.
Puccinia pseudocesatii Cummins
Puccinia pulverulenta Grev.
Puccinia pulvinata Rabenh.
Puccinia punctata Link
Puccinia purpurea Cooke
Puccinia pusilla Syd. & P. Syd.
Puccinia recondita Roberge ex Desm.
Puccinia ribis DC.
Puccinia recondita var. simlensis A.P. Misra, S.T. Ahmad & Sheodh. Singh
Puccinia ribis-caricis Kleb.
Puccinia rhynchosporae Syd. & P. Syd.
Puccinia romagnoliana Maire & Sacc.
Puccinia roscoeae Barclay
Puccinia rostrata Cooke
Puccinia rottboelliae P. Syd. & Syd.
Puccinia ruelliae Lagerh.
Puccinia rufipes Dietel
Puccinia sacchari Patel, Kamat & Y.A. Padhye
Puccinia sonchi Roberge ex Desm.
Puccinia satarensis P.B. Chavan & Bakare
Puccinia saussureae Thüm.
Puccinia saviculae Grev.
Puccinia saxifragae-ciliatae Barclay
Puccinia schedonnardi Kellerm. & Swingle
Puccinia schirajewskii Tranzschel
Puccinia scirpi DC.
Puccinia senecionis-scandentis Lindr.
Puccinia shiraiana P. Syd.
Puccinia silvaticella Arthur & Cummins
Puccinia solanacearum Sacc. & P. Syd.
Puccinia solani-giganteae T.S. Ramakr. & K. Ramakr.
Puccinia solmsii (Kuntze) Sacc. & P. Syd.
Puccinia sorghi Schwein.
Puccinia spongiosa Berk. & Broome
Puccinia stenotaphricola J. Walker
Puccinia striiformis Westend.
Puccinia striiformis f. muehlenbergii
112
Puccinia suaveolens (Pers.) Rostr.
Puccinia substriata Ellis & Barthol.
Puccinia swertiae G. Winter
Puccinia tanaceti DC.
Puccinia terminaliae T.S. Ramakr. & Sundaram
Puccinia thlaspeos Ficinus & C. Schub.
Puccinia thomasiana T.S. Ramakr. & K. Ramakr.
Puccinia thunbergiae Cooke
Puccinia thwaitesii Berk.
Puccinia tiliaefolia T.S. Ramakr. & Sundaram
Puccinia tragiae Cooke
Puccinia tricholepidis Syd.
Puccinia trollii P. Karst.
Puccinia turgida P. Syd. & Syd.
Puccinia tweediana T.S. Ramakr. & K. Ramakr.
Puccinia unica var. chica Cummins & S.M. Husain
Puccinia urticae Barclay
Puccinia ustalis Berk.
Puccinia verruca Thüm.
Puccinia vernoniae-monosis T.S. Ramakr. & K. Ramakr.
Puccinia versicolor Dietel & Holw.
Puccinia violae (Schumach.) DC.
Puccinia volutarellae Thirum.
Puccinia wangikarii Somani
Puccinia wattiana Barclay
Puccinia weyrehii Miyabe
Puccinia woodii (Kalchbr. & Cooke) P. Syd. & Syd.
Puccinia xanthii Schwein.
Puccinia xanthocarpi R.Y. Roy & P.C. Gupta
Puccinia xanthopoda Syd. & P. Syd.
Puccinia xanthosperma Syd. & P. Syd.
Puccinia zingiberis T.S. Ramakr.
Pucciniosira Lagerh (1)
Pucciniosira tuberculata (Ellis & Kellerm.) Buriticá & J.F. Hennen
Ramakrishnania Ramachar & Bhagyan. (1)
Ramakrishnania ixorae Ramachar & Bhagyan.
Trochodium Syd. & P. Syd. (2)
Trochodium ajrekarii Gharse
Trochodium sampathense Thirum.
Uromyces (Link) Unger (89)
Uromyces achrous Syd. & P. Syd.
Uromyces aconiti Fuckel
Uromyces acori T.S. Ramakr. & Rangaswami
Uromyces agropyri Barclay
Uromyces aloes (Cooke) Magnus
Uromyces ambiens Cooke
Uromyces amphilophis-insculptae T.S. Ramakr., Sriniv. & Sundaram
Uromyces andropogonis Tracy
Uromyces andropogonis-annulati Syd., P. Syd. & E.J. Butler
Uromyces anotidis-monospermatis T.S. Ramakr. & Sundaram
Uromyces anthyllidis (Grev.) J. Schröt.
Uromyces apludae Syd., P. Syd. & E.J. Butler
113
Uromyces appendiculatus (Pers.) Link
Uromyces behenis (DC.) Unger
Uromyces bidentis Lagerh.
Uromyces blainvilleae Berk.
Uromyces callicarpae (Petch) Fujik. ex S. Ito
Uromyces capitatus Syd. & P. Syd.
Uromyces ciceris-arietini (Grognot) Jacz. & G. Boyer
Uromyces clignyi Pat. & Har.
Uromyces clivalis Mitter
Uromyces commelinae Cooke
Uromyces coronatus Yoshin.
Uromyces dactylidis G.H. Otth
Uromyces decoratus Syd. & P. Syd.
Uromyces dolicholi Arthur
Uromyces eragrostidis Tracy
Uromyces eriochloae (Syd. & P. Syd.) Syd., P. Syd. & E.J. Butler
Uromyces euphorbiae Cooke & Peck
Uromyces fritillariae Thüm.
Uromyces geranii (DC.) G.H. Otth & Wartm.
Uromyces haussknechtii Tranzschel
Uromyces hedysari-obscuri (DC.) Carestia & Picc.
Uromyces heterogeneus Cooke
Uromyces hobsonii Vize
Uromyces hyderabadensis Ramachar, K.N. Rao & Bagyan.
Uromyces indigoferae f.sp. tinctoriae L.M. Joshi & A.R. Reddy
Uromyces ignobilis (Syd. & P. Syd.) Arthur
Uromyces inayatii Syd. & P. Syd.
Uromyces indicus Pat.
Uromyces indigoferae Dietel & Holw.
Uromyces lapponicus Lagerh.
Uromyces lespedezae-procumbentis (Schwein.) Lagerh.
Uromyces lespedezae-sericeae S. Ahmad
Uromyces lineolatus (Desm.) J. Schröt.
Uromyces loculiformis T.S. Ramakr. & K. Ramakr.
Uromyces macintirianus Barclay
Uromyces minor J. Schröt.
Uromyces mucunae Rabenh.
Uromyces muscari (Duby) Niessl
Uromyces mussooriensis Syd. & P. Syd.
Uromyces nilagiricus T.S. Ramakr. & K. Ramakr.
Uromyces orientalis Syd. & P. Syd.
Uromyces orthosiphonis T.S. Ramakr. & Sriniv.
Uromyces ottochloae Ramakr. T.S.
Uromyces panici-sanguinalis Rangel
Uromyces pavgii R.N. Goswami & Ngachan
Uromyces peglerae Pole-Evans
Uromyces pianhyensis Henn.
Uromyces pisi-sativi (Pers.) Liro
Uromyces polygoni-avicularis (Pers.) G.H. Otth
Uromyces pontederiicola Speg.
Uromyces poonensis W.D. More & Moniz
Uromyces proeminens (DC.) Lév.
114
Uromyces pseudarthriae Cooke
Uromyces ramacharii Ravinder & Bagyan.
Uromyces rottboelliae Arthur
Uromyces rugulosus Pat.
Uromyces rumicis (Schumach.) G. Winter
Uromyces satarensis P.B. Chavan & Bakare
Uromyces schoenanthi Syd. & P. Syd.
Uromyces setariae-italicae Yoshino
Uromyces sommerfeltii Hyl., Jørst. & Nannf.
Uromyces spegazzinii (De Toni) Arthur
Uromyces sphaeropleus Cooke
Uromyces sporgoni Clint & Peck. subsp. asiaticus
Uromyces striatus J. Schröt.
Uromyces strobilanthis Barclay
Uromyces superfluus P. Syd. & Syd.
Uromyces tenuicutis McAlpine
Uromyces triandrae T.S. Ramakr. & Sriniv.
Uromyces trichoneurae Doidge
Uromyces trifolii (R. Hedw.) Lév.
Uromyces trigonellae Pass.
Uromyces tripogonicola Payak & Thirum.
Uromyces trollii-caroli Ulbr.
Uromyces valerianae (Schumach.) Fuckel
Uromyces valerianae-wallichii (Dietel) Arthur & Cummins
Uromyces vestergrenii P. Syd. & Syd.
Uromyces viciae-fabae (Pers.) J. Schröt.
Uromyces vignae Barclay
Uromyces vossiae Barclay
Uromyces wedeliae-biflorae Boedijn
Uromyces wellingtonicus T.S. Ramakr. & K. Ramakr.
Xenostele Syd. & P. Syd. (1)
Xenostele litseae (Pat.) Syd. & P. Syd.
12. Raveneliaceae Leppik, Ann. bot. fenn. 9: 139. 1972. emend. Aime & McTaggart
Chaconia Juel (1)
Chaconia butleri (Syd. & P. Syd.) Mains
Didymopsorella Thirum. (1)
Didymopsorella macrospora (Mundk. & Thirum.) Thirum.
Diorchidium Kalchbr. (3)
Diorchidium levigatum Syd., P. Syd. & E.J. Butler
Diorchidium orientale Syd., P. Syd. & E.J. Butler
Diorchidium tricholaenae Syd. & P. Syd.
Gymnopuccinia K. Ramakr. (1)
Gymnopuccinia pulneyensis K. Ramakr.
Kernkampella Rajendren (6)
Kernkampella breyniae (Syd. & P. Syd.) Rajendren
Kernkampella breyniae-patentis (Mundk. & Thirum.) Rajendren
Kernkampella coimbatorica (T.S. Ramakr. & Sundaram) G.F. Laundon
Kernkampella emblicae (Syd. & P. Syd.) G.F. Laundon
Kernkampella kirganeliae (Mundk. & Thirum.) G.F. Laundon
Kernkampella phyllanthi (Mundk. & Thirum.) G.F. Laundon
115
Maravalia Arthur (11)
Maravalia achroa (Syd.) Arthur & Cummins
Maravalia aulica (Syd.) Y. Ono
Maravalia ascotela (Syd.) Mains
Maravalia echinulata (Niessl ex Rabenh.) Y. Ono
Maravalia fici (Mundk. & Thirum.) Y. Ono
Maravalia gentilis (Syd.) Y. Ono
Maravalia ichnocarpi (Barclay) Sathe
Maravalia millettiae Yadav & Thirum.
Maravalia milletticola Y. Ono & J.F. Hennen
Maravalia mimusops (Cooke) Y. Ono
Maravalia pterocarpi (Thirum.) Thirum.
Scopella Mains (1)
Scopella dalbergiae (T.S. Ramakr. & K. Ramakr.) Ragunathan & K. Ramakr.
Olivea Arthur (3)
Olivea colebrookiana Thirum. & Yadav
Olivea isonandrae Hosag.
Olivea tectonae (Racib.) Thirum.
Ravenelia Berk. (33)
Ravenelia acaciae-arabicae Mundk. & Thirum.
Ravenelia acaciae-caesiae Tyagi & S.S. Prasad
Ravenelia acaciae-concinnae Mundk. & Thirum.
Ravenelia acaciae-intsiae B.V. Patil & Thirum.
Ravenelia acaciae-pennatulae Dietel
Ravenelia acaciae-pennatulae Dietel
Ravenelia acaciae-sumae Thirum. & Mundk.
Ravenelia acaciae-senegalae Sanwal
Ravenelia acaciicola Sanwal
Ravenelia aculeifera Berk.
Ravenelia ajmerensis Sanwal
Ravenelia albiziae-amarae Bacc.
Ravenelia burmanica Thaung
Ravenelia cassiicola G.F. Atk.
Ravenelia clemensae Syd.
Ravenelia deformis Tyagi & S.S. Prasad
Ravenelia tephrosiicola (Henn.) Hirats. f.
Ravenelia esculenta Naras. & Thirum.
Ravenelia evansii Syd. & P. Syd.
Ravenelia fragrans Long
Ravenelia hansfordii Cummins
Ravenelia hobsonii Cooke
Ravenelia indica Berk.
Ravenelia japonica Dietel & P. Syd.
Ravenelia karadensis P.B. Chavan & U.V. Kulk.
Ravenelia mitis Syd. & P. Syd.
Ravenelia mitteri Syd.
Ravenelia odoratissimae Tyagi & S.S. Prasad
Ravenelia ornata Syd. & P. Syd.
Ravenelia parasnathii Yadav
Ravenelia radhanagarensis Patil
Ravenelia satarensis P.B. Chavan & U.V. Kulk.
Ravenelia sayeedii M.A. Salam & Ramachar
116
Ravenelia sessilis Berk.
Ravenelia spicigerae B.V. Patil & Thirum.
Ravenelia stictica Berk. & Broome
Ravenelia sumatii S.D. Patil & Date
Ravenelia tandonii Syd.
Ravenelia taslimii Mundk.
Ravenelia versatilis (Peck) Dietel
Prospodium Arthur (2)
Prospodium erebia (Syd. & P. Syd.) Bagyan. & Ravinder
Prospodium tirumalense Bagyan., Ravinder & P. Ramesh
13. Skierkaceae (Arthur) Aime & McTaggart, Fungal Systematics and Evolution 7: 21–47, 2020.
Skierka Racib. (3)
Skierka agallochae Racib.
Skierka himalayensis A.K. Gautam & S. Avasthi
Skierka toddaliae (Petch) Hirats.
14. Sphaerophragmiaceae Cummins & Y. Hirats., Illustr. Gen. Rust Fungi, rev. Edn (St. Paul): 15.
1983. emend. Aime & McTaggart
Sphaerophragmium Magnus (1)
Sphaerophragmium acaciae (Cooke) Magnus
15. Tranzscheliaceae (Arthur) Aime & McTaggart, Fungal Systematics and Evolution 7: 21–47,
2020
Leucotelium Tranzschel (1)
Leucotelium pruni-persicae (Hori) Tranzschel
Tranzschelia Arthur (2)
Tranzschelia discolor (Fuckel) Tranzschel & M.A. Litv.
Tranzschelia pruni-spinosae (Pers.) Dietel
16. Zaghouaniaceae P. Syd. & Syd., Monogr. Uredin. (Lipsiae) 3(3): 586. 1915. emend. Aime &
McTaggart
Cystopsora E.J. Butler (1)
Cystopsora antidesmatis T.S. Ramakr. & Sundaram
Elateraecium Thirum., F. Kern & B.V. Patil (1)
Elateraecium salaciicola Thirum., F. Kern & B.V. Patil
Hemileia Berk. & Broome (10)
Hemileia canthii Berk. & Broome
Hemileia gardeniae-thunbergiae (Henn.) Maubl. & Roger
Hemileia holarrhenae Syd. & P. Syd.
Hemileia indica Massee
Hemileia jasmini C.S. Krishnam. & Rangaswami
Hemileia mysorensis Thirum. & Gopalakrishn.
Hemileia pavetticola Maubl. & Roger
Hemileia thomasii Thirum. & Naras.
Hemileia vastatrix Berk. & Broome
Hemileia wrightiae (Racib.) Racib.
Zaghouania Pat. (1)
Zaghouania oleae (E.J. Butler) Cummins
Pucciniales genera incertae sedis
Aecidium Vuill.
117
Aecidium adhatodae Syd. & P. Syd.
Aecidium aechmantherae Syd. & P. Syd.
Aecidium ajugae Syd. & P. Syd.
Aecidium anaphalidis-leptophyllae T.S. Ramakr., Sriniv. & Sundaram
Aecidium argyreiae Berk. & Broome
Aecidium brasiliense Dietel
Aecidium callianthum Syd.
Aecidium campanulae Pandotra & K.S.M. Sastry
Aecidium carviae Sathe
Aecidium cassiae-torae P.B. Chavan & Bakare
Aecidium cinnamomi Racib.
Aecidium cleomes Ellis & H.W. Anderson
Aecidium clerodendri Henn.
Aecidium colchici-aurei Ulbr.
Aecidium crassocephali Wakef. & Hansf.
Aecidium crini Kalchbr.
Aecidium crypticum Kalchbr. & Cooke
Aecidium cuspidatum T.S. Ramakr., Sriniv. & Sundaram
Aecidium delphinii Barthol.
Aecidium deutziae Dietel
Aecidium dichrocephalae Henn.
Aecidium diospyri A.L. Sm.
Aecidium distinctum Arthur & Cummins
Aecidium elaeocarpi-tuberculati Hosag.
Aecidium esculentum Barclay
Aecidium flavescens Barclay
Aecidium garciniae Sundaram & A.V. Rao
Aecidium girardiniae Syd. & P. Syd.
Aecidium gymnematis T.S. Ramakr. & Sundaram
Aecidium hartwegiae Thüm.
Aecidium hedyotidis Syd.
Aecidium hemidesmi Syd. & P. Syd.
Aecidium hemigraphidis B.V. Patil & Thirum.
Aecidium infrequens Barclay
Aecidium innatum Syd., P. Syd. & E.J. Butler
Aecidium inquitosense P. Henn.
Aecidium kamatii Sathe
Aecidium latifolium Massee
Aecidium leeae M.A. Salam & Ramachar
Aecidium lepidagathis Syd. & P. Syd.
Aecidium leucadinum Mitter
Aecidium lophanthi Henn.
Aecidium lophopetali Wakef.
Aecidium marsdeniae T.S. Ramakr. & K. Ramakr.
Aecidium melaleucum Syd. & P. Syd.
Aecidium meliosmae-wightii T.S. Ramakr., Sriniv. & Sundaram
Aecidium memecyli Thirum.
Aecidium microrhynchi Henn.
Aecidium miliare Berk. & Broome
Aecidium montanum E.J. Butler
Aecidium mori Barclay
Aecidium morobeanum Cummins
118
Aecidium myriactidis (Barclay) P. Syd. & Syd.
Aecidium nummulare Berk.
Aecidium ocimi Henn.
Aecidium orbiculare Barclay
Aecidium osmanthi Syd., P. Syd. & E.J. Butler
Aecidium painavuense Hosag.
Aecidium paramignyae Racib.
Aecidium pavoniae-odoratae T.S. Ramakr., Sriniv. & Sundaram
Aecidium peristrophes Syd. & P. Syd.
Aecidium petchii Sacc. & Trotter
Aecidium plectranthicola Cummins
Aecidium plectroniae Cooke
Aecidium ponderosum Syd. & P. Syd.
Aecidium poonense Sathe
Aecidium pulneyense T.S. Ramakr. & Sriniv.
Aecidium pupaliae Prasad, L.C. Sharma & R.D. Singh
Aecidium pygei Syd. & P. Syd.
Aecidium quintum Syd. & P. Syd.
Aecidium randiae Henn.
Aecidium rhododendri Barclay
Aecidium rhynchosiae Bagyan. & Ramachar
Aecidium rhytismoideum Berk. & Broome
Aecidium salamii G.F. Laundon
Aecidium satarense P.B. Chavan & S.K. Patil
Aecidium saussureae Johanson
Aecidium scutellariae Syd. & P. Syd.
Aecidium sinhagadense Sathe
Aecidium solani Mont.
Aecidium spilanthis T.S. Ramakr. & Sundaram
Aecidium stewartianum Cummins
Aecidium stewartii Arthur & Cummin
Aecidium stranvaesiae Syd. & P. Syd.
Aecidium strobilanthis Barclay
Aecidium tandonii Mitter
Aecidium terminaliae T.S. Ramakr. & K. Ramakr.
Aecidium travancoricum T.S. Ramakr.
Aecidium tricholepidis P.B. Chavan & Bakare
Aecidium tubulosum Pat. & Gaillard
Aecidium urceolatum Cooke
Aecidium urgineae Henn. & Pole-Evans
Aecidium vangueriae Cooke
Aecidium verbenae Speg.
Aecidium vernoniae-cinereae Petch
Aecidium walayarense T.S. Ramakr. & Sundaram
Aecidium withaniae Thüm.
Nyssopsora Arthur
Nyssopsora cedrelae (Hori) Tranzschel
Nyssopsora thirumalacharii R.N. Goswami & Ngachan
Nyssopsora thwaitesii (Berk. & Broome) Syd.
Phragmotelium Syd.
Phragmotelium burmanicum (Syd. & P. Syd.) Syd.
Phragmidium mysorense (Thirum. & Mundk.) B. Ali & Berndt
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Tunicopsora Suj. Singh & P.C. Pandey
Tunicopsora bagchii Suj. Singh & P.C. Pandey
Uraecium Arthur
Uraecium nothopegiae T.S. Ramakr. & K. Ramakr.
Uredo Pers.
Uredo acaciae-concinnae Kapoor bis & D.K. Agarwal
Uredo acalyphae-fruticosae T.S. Ramakr., Sriniv. & Sundaram
Uredo allmaniae P.B. Chavan & U.V. Kulk.
Uredo alpestris J. Schröt.
Uredo amomi Petch
Uredo apludae Barclay
Uredo arachidis Lagerh.
Uredo brachylepidis T.S. Ramakr. & Sundaram
Uredo cajani Syd. & P. Syd.
Uredo carissae Thirum.
Uredo carissae–occidentalis Chavan & Kulkurni
Uredo cassiae K.N. Rao
Uredo cassiae-occidentalis T.S. Ramakr.
Uredo celastri Arthur & Cummins
Uredo celastri-paniculatae T.S. Ramakr. & Sundaram
Uredo chasaliae Petch
Uredo citri Vaheed.
Uredo davaoensis Syd. & P. Syd.
Uredo dalbergiae-latifoliae Hosag. & N.C. Nair
Uredo deutziae Barclay
Uredo dioscoreae Henn.
Uredo dioscoreae-sativae Syd. & P. Syd.
Uredo echinulata (Niessl.) Syd.
Uredo ehretiae Barclay
Uredo elephantopodis Petch
Uredo elettariae Thirum.
Uredo emiliae-scabrae T.S. Ramakr. & Sundaram
Uredo exasperata (Cooke) Sacc.
Uredo garugae Sundaram
Uredo gayanae J.C. Lindq.
Uredo gharsei Sathe & Rahalkar
Uredo gomphrenae Barclay
Uredo hygrophilicola G.F. Laundon & Ponnappa
Uredo hyperici-mysorensis Petch
Uredo khandalensis Sathe & Rahalkar
Uredo launeae–coromandelicae Chavan & Bakare
Uredo lipocarphae Syd. & P. Syd.
Uredo malabarica T.S. Ramakr. & K. Ramakr.
Uredo mannanurensis K.N. Rao
Uredo microspora (Vize) Sacc.
Uredo mundkurii P.B. Chavan
Uredo myriactidis Sundaram
Uredo neilgherriensis T.S. Ramakr.
Uredo niterogensis Rangel
Uredo ochnae K.N. Rao
Uredo ophiorrhizae Petch
Uredo ophiuri Syd., P. Syd. & E.J. Butler
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Uredo paederiae Syd. & P. Syd.
Uredo pallidula Cooke & Massee
Uredo panacis Syd. & P. Syd.
Uredo paspali Pandotra
Uredo paspali-scrobiculati Syd. & P. Syd.
Uredo phyllanthi-niruris M.S. Patil
Uredo pileae Barclay
Uredo pouzolziae Syd. & P. Syd.
Uredo pterocarpi T.S. Ramakr.
Uredo punctoidea Cooke
Uredo ravennae Maire
Uredo rhinacanthi T.S. Ramakr. & Sundaram
Uredo rottboelliae Dietel
Uredo shuteriae T.S. Ramakr.
Uredo sesbaniae Henn.
Uredo setariae-tomentosae Ramachar, Bhagyan. & A. Kumar
Uredo sissoo Syd., P. Syd. & E.J. Butler
Uredo spinulosa (Cooke) Sacc.
Uredo tephrosiae Rabenh.
Uredo terminaliae Henn.
Uredo terminaliae-paniculatae T.S. Ramakr. & K. Ramakr.
Uredo thelypteridis Yadav
Uredo thelypteridis var. thelypteridis Yadav
Uredo verecunda Syd.
Uredo victoriae Cummins
Phylogenetic analysis
The phylogenetic analyses based on the combined LSU and ITS rDNA sequence dataset
comprised 189 taxa including various genera from 13 families belonging to Pucciniales were assessed
with Taphrina pruni CBS 358.35 as an outgroup taxon. RAxML analysis of the combined dataset
produced the best tree with a final ML optimization likelihood value of -34581.032655. The genera
from different families included in the phylogenetic analyses are Coleosporiaceae, Cronartiaceae,
Crossopsoraceae,
Melampsoraceae,
Milesinaceae,
Ochropsoraceae,
Phakopsoraceae,
Phragmidiaceae, Pileolariaceae, Pucciniastraceae, Pucciniaceae, Raveneliaceae and
Tranzscheliaceae. The genera Coleosporium, Chrysomyxa clustered together in Coleosporiaceae
clade with significant support from ML 93% and Cronartia formed a distinct clade sharing a sister
group relationship with Coleosporium and Chrysomyxa. The Melampsoraceae clade is supported by
taxa from Melampsora with significant support from ML 100%. The Pucciniastraceae clade consists
of taxa from Hyalospora, Melampsoridium and Pucciniastrum and Milesinaceae includes Milesina.
The genera Ravenelia, Kernkampella clustered in a Raveneliaceae clade and Maravalia formed a
distinct lineage. While Nyssopsora belonging to Uredinineae incertae sedis formed a different
lineage. The Tranzschelia includes Tranzschelia and Leucotelium and Ochropsoraceae comprises
Ochropsora. The Phakopsoraceae includes Cerotelium and Phakopsora in a monophyletic clade.
The Crossopsoraceae clade comprises Angiopsora, Crossopsora and Kweilingia. The
Phragmidiaceae includes taxa from Phragmidium in monophyletic clade with significant support
from ML 94%. The Gymnosporangiaceae comprised Gymnosporangium in a monophyletic clade
with significant support from ML 96%. The Pucciniaceae includes polyphyletic taxa from Puccinia
and Uromyces.
Notes of Indian rust genera
After going through the available literature on Indian rust fungi, it is now clear that
identification of these fungi has primarily been based on morphological characters. Few studies are
121
reported to use modern tools and molecular based techniques (specifically DNA-based) in their
taxonomy. But wth the use of all modern molecular methdologies, a number of alterations in existing
system of classification of rust fungi has been proposed by Cummins & Hiratsuka (2003). Number
of changes as proposed recently in classification of rust fungi (Aime 2006, Aime & McTaggart 2020),
has led to the introcution of many new families as well as transfer of many genera and species.
Keeping in view all the proposed changes, this outline of Indian Pucciniales incorporates updated
changes. Of the 18 families of rust fungi, 16 are reported from India, consists 69 genera and 640
species. Most of the genera and species of Indian collections still required DNA based identification.
Therefore, a brief description of each rust family along with total number of genera as well as species
reported from India and their host families are provided in this section. In addition, a brief note, where
further studies on Indian collections are urgently required to resolve their taxonomic uncertainty is
also provided.
Phylum – Basidiomycota R. T. Moore, Bot. Mar.23: 371 (1980).
Synonyms – Basidiomycota Bold, Morph. Pl.: 7, 198 (1958), nomen invalidum
Basidiomycota is the second largest phyla of kingdom Fungi which shares 97% of all fungal
species along with phylum Ascomycota (Wijayawardene et al. 2017, 2018, 2020, Niskanen et al.
2018). The fungi included in Basidiomycota possess basidia as meiosporocysts in the sexual life
stage. The hyphae appeared to have single-layered wall (which actually is multi-layered) are divided
by septa into mononucleate, binucleate, or multinucleate segments. The septal pore is generally
closed, however, in some cases barrel-like thickening is present on both sides. Chemotaxonomy,
formation of urease, siderochromes, and the type of ubiquinone system also differentiate
basidiomycetous fungi from ascomycetes. In addition, the guanine-cytosine content of the total DNA
exceeds 50% in basidiomycetous species. The updated outline of Basidiomycota includes four
subphyla, 18 classes, 68 orders, 241 families, 1928 genera and 41270 species, of which rust fungi are
included in subphylum Pucciniomycotina (He et al. 2019, Wijayawardene et al. 2020).
Subphylum: Pucciniomycotina R. Bauer, Begerow, J.P. Samp., M. Weiß & Oberw., Mycol. Progr.5:
45 (2006).
Equivalent to Urediniomycetes (Swann & Taylor 1995b, Kirk et al. 2001, Swann et al. 2001).
Pucciniomycotina is a diverse group of fungi, including rusts, yeasts, smut-like and jelly-like
fungi. It is the sister to the Ustilaginomycotina and Agaricomycotina, forming the basal lineage of
Basidiomycota. Species of Pucciniomycotina studied so far lack dolipores (septal pore swellings) and
septal pore caps. Absence of the predominant cell wall sugar, mannose (Prillinger et al. 1993) and
disc like spindle pole bodies (McLaughlin et al. 1995, Wells 1994), distinguishes them from most
other Basidiomycota. These fungi show very simple to complex life cycle, considered as most
complex organisms (Lutz et al. 2004). Most described species are predominantly phytopathogens but
also include asymptomatic members. Subphylum Pucciniomycotina is estimated to have 10 classes,
22 orders, 49 families, 270 genera, and 8653 species with rust fungi placed in class Pucciniomycetes
(He et al. 2019, Wijayawardene et al. 2020).
Class: Pucciniomycetes R. Bauer, Begerow, J.P. Samp., M. Weiss & Oberw., Mycol. Progr. 5: 48
(2006).
Equivalent to Urediniomycetidae (Swann et al. 2001).
Pucciniomycetes is a diverse class of subphylum Pucciniomycotina. It is one of the major
classes of basidiomycete fungi containing about 8000 species (Kirk et al. 2008). Based on rDNA
phylogenetic studies, rust fungi and their closest relatives in Pucciniomycetes are reported to have
ambiguous phylogenetic positions within the Pucciniomycotina (Aime et al. 2006). All species of
Pucciniomycetes are dikaryotic (containing two haploid nuclei per cell) except for Septobasidiales,
which are monokaryotic (containing a single haploid nucleus per cell). Lack of clamp connections in
their hyphae (Bauer et al. 2006) is another characteristic feature of these fungi. Production of asexual
spores, especially among rusts, is often well developed (Bruckart et al. 2010). Sexual reproduction
takes place via the formation of basidiospores. This class contains 5 orders, 20 families, 180 genera,
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and 8016 species. Majority of the fungal species in Pucciniomycetes are parasitic in nature.
Pucciniales is the most species-rich group of the Pucciniomycetes with over 95% of the species and
75% of the genera placed in this order, the plant parasitic rust fungi.
Order: Pucciniales Clem. & Shear, Gen. Fungi (2nd edn): 147 (1931).
Equivalent to Uredinales.
Exemplar genera: Puccinia Pers. 1801, Uromyces (Link) Unger 1832.
The Pucciniales is one of the largest and major orders in Basidiomycota (class
Pucciniomycetes). This order mainly contains many important plant pathogens popularly known as
rusts. These fungi are obligate plant parasites occurring on ferns, gymnosperms and angiosperms.
They have been studied in detailed as many of the most devastating plant diseases in agricultural
crops are caused by the members of Pucciniales. Morphologically, the species of Pucciniales are
characterized by their rusty appearance on infected host parts such as leaves, petioles, tender shoots,
stems, fruits, etc. and named for the typically rusty coloration of their urediniospores. These obligate
parasites have highly complex life cycles with up to five spore stages and two unrelated hosts
(Cummins & Hiratsuka 2003). Of 7800 described species, Pucciniales constitutes 25% of all known
species in Basidiomycota and ca. 8% of all described Fungi (Kirk et al. 2008). The Pucciniales is
estimated to have 15 families, ca. 150 genera and ca.7, 800 species, and is considered as the most
speciose order of fungi (Kirk et al. 2008, Wijayawardene et al. (2020). Recently, a higher-rank
classification for rust fungi, with notes on genera was provided by Aime & McTaggart (2020). They
have proposed four new suborders and seven new families based on the evaluation of 80 % of
accepted genera including type species wherever possible, and three DNA loci. As per this
classification, Pucciniales now comprises seven suborders as Araucariomycetineae,
Melampsorineae, Mikronegeriineae, Raveneliineae, Rogerpetersoniineae, Skierkineae, and
Uredinineae. There are now 18 families Araucariomycetaceae, Coleosporiaceae, Crossopsoraceae,
Gymnosporangiaceae, Melampsoraceae, Milesinaceae, Ochropsoraceae, Phakopsoraceae,
Phragmidiaceae,
Pileolariaceae,
Pucciniaceae,
Pucciniastraceae,
Raveneliaceae,
Rogerpetersoniaceae, Skierkaceae, Sphaerophragmiaceae, Tranzscheliaceae, and Zaghouaniaceae
(Aime & McTaggart 2020). Therefore, we followed Aime & McTaggart (2020) to present updated
information on genera and species in each family. The numbers of genera and species of rust fungi
were presented as per Wijayawardene et al. (2020). The distinct characteristics of various spore stages
(telia, teliospores, uredinia, uredinospores, aecia, aeciospores, spermogonia and basidia) for each
family are summarized in this section. The notes on rust families reported from India are described
here in this section of manuscript.
Coleosporiaceae Dietel, In: Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 548. 1900.
emend. Aime & McTaggart
Type genus – Coleosporium Lév., Ann. Sci. Nat. Bot. III, Ser. 8: 373. 1847
The species of family Coleosporiaceae bear large, bladder-like aecia covered with welldeveloped peridium and have catenulate, verrucose aeciospores. Uredinia with rudimentary peridium
or none and urediniospores are formed in chain with verrucose echinulate ornamentation on surface.
The germ pores are mostly obscure and scattered. The teliospores are formed one by one (catenulate,
pseudocatenulate or in a single layer) in erumpent, hard, waxy or gelatinous, pulvinate or columnar
telia under the epidermis of the host plant. The teliospores are generally unicellular, thin walled and
sessile with non-differentiated germ pores. The spermogonia are of Group I (type 2 or 3) (but Group
II, type 9 in Cronartium). Most of the species are heteroecious and macrocyclic, with aecial stage on
needle, buds and cones of conifers (Cummins & Hiratsuka 2003, Aime & McTaggart 2020).
Genera reported in India – Coleosporium (19), Chrysomyxa (7), Cronartium (4), Goplana (1),
Stakmania (1) total 32 species.
Host families – Asteraceae, Campanulaceae, Ericaceae, Gentianaceae, Grossulariaceae,
Lamiaceae, Lauraceae, Moraceae, Phyllanthaceae, Pinaceae, Ranunculaceae.
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Figure 1 – Maximum likelihood phylogeny of Indian rust fungi based on two concatenated loci (LSU
and ITS) of taxa from Pucciniales. Bootstrap support values for maximum likelihood equal to or
124
greater than 70% are given above each branch respectively. Outgroup taxon is Taphrina pruni CBS
358.35.
Figure 1 – Continued.
Notes – Two species of rust genus Stakmania (S. formosana and S. indica) were reported from
India. Among the two, S. formosana Syd. & P. Syd. has been transferred to Phakopsora formosana
while, S. indica Kamat & Sathe remains unchanged (Index Fungorum 2020). Hence, this species has
been retained in this family.
Crossopsoraceae Aime & McTaggart, In: Fungal Systematics and Evolution 7: 21–47. 2020
Type genus – Crossopsora Syd. & P. Syd., Annls mycol. 16(3/6): 243. 1919.
Fig. 9
125
Crossopsoraceae is characterized by producing Group VI (type 7) spermogonia and aecidiumtype aecia wherever known. Uredinia are of malupa-type and usually paraphysate. The teliospores
produced by these fungi are 1-celled, compact and often produced in catenulate chains of a few to
many cells. These spores germinate externally, with or without dormancy. The fungi in this family
are mostly identified from the sporothallus (uredinial, and telial) stages. The genus Neophysopella in
this family is macrocyclic and heteroecious in nature (Aime & McTaggart 2020) and considered the
same for other genus/species also. A total of 7 genera have been included in this family, of which,
six genera being reported from India.
Genera reported in India – Angiopsora (2), Crossopsora (3), Dasturella (3), Kweilingia (2),
Neophysopella (2), Physopella (2); total 14 species.
Host families – Poaceae, Rhamnaceae Verbenaceae.
Notes – Two Indian records of the genus Physopella i.e. P. artocarpi (Berk. & Broome) Arthur
and P. vernoniae (T.S. Ramakr.) Ramachar & Bhagyan are solely based on morphs. Since the genus
Physopella has now been changed to Neophysopella, these two Indian records are still unchanged
(Index Fungorum 2020). Hence, we retained this species in this family and proposed DNA based
studies to resolve its taxonomic position.
Gymnosporangiaceae P. Zhou & L. Cai, Persoonia 45: 79. 2020. emend. Aime & McTaggart
Type genus – Gymnosporangium R. Hedw. ex DC., In: Lamarck & de Candolle, Fl. franç.,
Edn 3 (Paris) 2: 216. 1805.
The family Gymnosporangiaceae consists of Group V (type 4) Spermogonia, bounded with
well developed peripheral flexuous hyphae. Aecia Roestelia-type, subepidermal, with welldeveloped peridia (Gymnosporangium) or less frequently aecidium-type (Gymnotelium).
Aeciospores catenulate, with intercalary cells. Urediniospores borne singly on pedicels in Uredotype, subepidermal Uredinia. Telia subepidermal, erumpent consists mostly 2-celled teliospores,
borne singly on gelatinising pedicels without dormancy via external basidia. Life cycle mostly
demicyclic and heteroecious (Zhao et al. 2020, Aime & McTaggart 2020).
Genera reported in India – Peridiopsora (1) Gymnosporangium (3) Roestelia (3)
Host families – Boraginaceae, Cupressaceae, Rosaceae.
Notes – Based on puccinioid character of 2-celled, pedicellate teliospores, the genus name
Gymnosporangium has been conserved against the older name Roestelia Rebent. (Aime et al. 2018b).
Hence, the genus Roestelia has now been transferred to Gymnosporangium. Because of lack of
molecular data, Indian records of this genus reported as Roestelia distorta (Arthur & Cummins) F.
Kern, Roestelia cunninghamianum (Barclay) F. Kern and Roestelia patula (Syd. & P. Syd.) F. Kern
are still unchanged (Index Fungorum 2020).
Melampsoraceae Dietel, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 38. 1897.
Figs 2, 7, 9
Type genus – Melampsora Castagne, Obs. Plantes Acotylédonées Fam. Urédinié 2: 18. 1843.
Melampsoraceae is characterized by aecia without peridium or rudimentary if present with
catenulate and verrucose aeciospores. Uredinia contain abundant paraphyses (sometimes
rudimentary peridium also) and echinulate urediniospores with scattered or bizonate germ pores and
borne singly. Telia are embedded (subepidermal or rarely subcuticular) containing unicellular,
sessile, pigmented teliospores with one germ pore. Germination external or semi-external
(Ceropsora). The basidium is external and spermogonia are of Group I (Type 2 or 3). These fungi
are mostly macrocyclic and inhabit two different unrelated hosts or same host to produce all spore
stages. Most of the species are heteroecious and macrocyclic; however, the species of Ceropsora are
microcyclic. Total 100 species of Melampsora have been reported globally (He et al. 2019,
Wijayawardene et al. 2020), of which only 17 species have been reported so far in India.
Genera reported in India – Ceropsora (1), Melampsora (17); total 18 species.
Host families – Euphorbiaceae, Hypericaceae, Linaceae, Pinaceae, Salicaceae.
126
Notes – The type species of the genus Ceropsora (C. picea) infecting Picea sp. was reported
from India (Bakshi & Singh 1960). It was placed in family Coleosporiaceae (Cummins & Hiratsuka
2003, Wijayawardene et al. 2020). The telia of two species of Ceropsora viz., C. picea and C. weirii
contain some thin-walled sterile cells on the sides (been interpreted as remnants of a peridermium).
Teliospores of the genus are subtended by adherent crusts of sterile basal cells in the beginning that
separate at dispersal stage (Bakshi & Singh 1960, Crane et al. 2000). Aime & McTaggart (2020)
proposed the inclusion of the genus Ceropsora in to this family.
Figure 2 – Rust fungi Melampsora sp. A-B Euphorbia helioscopia. C-D Populus sp.
Milesinaceae Aime & McTaggart, In: Fungal Systematics and Evolution 7: 21–47. 2020
Type genus – Milesina Magnus, Ber. Deutsch. Bot. Ges. 27: 325. 1909.
Two important characters, production of colourless urediniospores in species that infect ferns
and production of aecia (milesia-type) in species that infect Ericaceae differentiate this family from
Melampsorineae. The important characters of this family include production of Group I (mostly type
1, also type 2 and 3) Spermogonia and colourless sori. Although urediniospores of Naohidemyces are
orange in colour. The presence of peridermium-type aecia (milesia-type in Naohidemyces); milesiatype uredinia; 1- to many-celled, barely differentiated, sometimes laterally adherent teliospores with
dormant germination are found. These fungi are mostly macrocyclic and heteroecious in nature. They
produce sporothalli on ferns (except Naohidemyces on Ericaceae), and gametothalli on Pinaceae.
Genera reported in India – Milesina (3), Uredinopsis (2); total 5 species
Host families – Dryopteridaceae, Pteridaceae.
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Note – This is newly proposed family by Aime & McTaggart (2020). Out of 4 genera included
in this family (Aime & McTaggart 2020), two genera namely, Milesina and Uredinopsis are reported
from India. Aime et al. (2018) recommended the protection of the name Milesina Magnus over
Milesia F.B. White, but they do not found the type species of both the genera congeneric and
recommended retaining both the genera. In India, the molecular studes on Milesina and Uredinopsis
are also required to demonstrate their correct taxonomic position.
Ochropsoraceae (Arthur) Aime & McTaggart, In: Fungal Systematics and Evolution 7: 21–47. 2020
Type genus – Ochropsora Dietel, Ber. Dtsch. Bot. Ges. 13: 401. 1895.
The family Ochropsoraceae contains Group VI (type 7) spermogonia. Aecia are of aecidiumtype, these states systemic in nature and overwintering as mycelium. Uredinia are of malupa-type.
The genera of this family produce 1-cell deep telia forming crusts, which are at first subepidermal,
then erumpent. Teliospores germinate without dormancy, either internally (Ochropsora) or
externally (Aplopsora). These fungi are macrocyclic and heteroecious in nature.
Genera reported in India – Ochropsora (1); total 1 species
Host families – Ranunculaceae.
Note – Aime & McTaggart (2020) proposed Ochropsoraceae as new family. The rust genera
Aplopsora and Ochropsora included in this family were previously treated within the Chaconiaceae
(Cummins & Hiratsuka 2003). Of these, Ochropsora (Ochropsora ariae (Fuckel) Ramsb.) was also
reported from India. However, only morphotaxonomic studies are available for this record, and
molecular studies can be helpful to establish the correct taxonomic position.
Phakopsoraceae Cummins & Y. Hirats., Illustr. Gen. Rust Fungi, rev. Edn (St. Paul): 13. 1983.
emend. Aime & McTaggart
Type genus – Phakopsora Dietel, Ber. Deutsch. Bot. Ges. 13: 333. 1895.
The genera of Phakopsoraceae show both heteroecious and autoecious (species of Bubakia,
Masseeëlla and Nothoravenelia) mode of life cycle on various hosts but not host restricted.
Spermogonia are of Group IV (type 7). The aecia are caeoma-type (some Masseeëlla with aecidiumtype) with aeciospores either verrucose or echinulate, borne singly or in chains. Uredinia lecythea or
uredo-type consisting echinulate urediniospores, mostly borne singly (catenulate in Arthuria) with
scattered germ pores or obscure. Both erumpent and embedded types of telia are found which contain
1-celled, sessile, catenulate or irregularly arranged teliospores with mostly 1 germ pore in each cell
The majority of Phakopsora and Uredopeltis species are only known from the sporothallus (uredinial,
and telial) stages (Cummins & Hiratsuka 2003, Aime & McTaggart 2020). This family comprises 15
genera and 209 species (Wijayawardene et al. 2020). Aime & McTaggart (2020) in recent treatment
included 12 genera in this family, 10 being reported from India.
Genera reported in India – Arthuria (2), Bubakia (1), Cerotelium (7), Macabuna (1),
Monosporidium (3), Masseeëlla (6), Phakopsora (25), Phragmidiella (3), Pucciniostele (1),
Uredopeltis (1); total 50 species.
Host families – Bignoniaceae, Burseraceae, Combretaceae, Euphorbiaceae, Fabaceae,
Phyllanthaceae, Putranjivaceae, Rhamnaceae, Rubiaceae, Salicaceae, Saxifragaceae and Vitaceae.
Notes – The rust genus Bubakia has now been transferred to Phakopsora (Index Fungorum
2021). It is often treated as a synonym of Phakopsora (Cummins & Hiratsuka 2003). But, Aime &
McTaggart (2020) found a distinct lineage of the Bubakia argentinensis within Phakopsoraceae and
similar characteristics and hosts with type, B. crotonis. Hence, they proposed Bubakia for these
species. However, the only India record of Bubakia indica T.S. Ramakr. & K. Ramakr. still requires
molecular studes, as it was identified based on morphology only. The genus Masseeëlla is also placed
within this family (Aime & McTaggart 2020), although previously treated as incertae sedis within
Pucciniales (Cummins & Hiratsuka 2003).
Phragmidiaceae Corda Icon. fung. (Prague) 1: 6. 1837
Type genus – Phragmidium Link, Mag. Ges. Naturfr. Freunde Berlin 7: 30. 1816.
Figs 3, 9
128
Phragmidiaceae is characterized by forming spermogonia Group IV (type 6, 8, 10 or 11). The
aecia variable, caeoma-, petersonia- or uredo-type and are formed with or without peridium.
Aeciospores are verrucose or echninulate aeciospores borne in chains or singly on pedicels with few
or no paraphyses. Uredinia lecythea or uredo-type, thin-walled, with incurved paraphyses consisting
of echninulate urediniospores borne singly and have scattered germ pores. Telia erumpent, with or
without paraphyses, with teliospores born on short pedicels, 3- to several cells (by transverse septa)
with 1 or more apical germ pore in each cell. Basidium is external. Life cycle is mainly of autoecious
type. Wijayawardene et al. (2020) documented 13 genera and 178 species in this family; however,
Aime & McTaggart (2020) included only 9 genera.
Genera reported in India – Hamaspora (2), Kuehneola (6), Phragmidium (18), Trachyspora
(1); total 27 species.
Host families – Poaceae, Rosaceae.
Notes – Of the 9 genera included in this family, only 4 genera and 27 species have been reported
in India. Phragmidiaceae species are confined almost exclusively to the Rosoideae subfamily of
Rosaceae (Aime 2006).
Figure 3 – A-B Phragmidium sp. on Rosa sp.
Pileolariaceae (Arthur) Cummins & Y. Hirats., llustr. Gen. Rust Fungi, rev. Edn (St. Paul): 14. 1983.
emend. Aime & McTaggart
Figs 4A, 9
Type genus – Pileolaria Castagne, Obs. Plantes Acotylédonées Fam. Urédinées 1: 22. 1842.
The genera of the family are characterized by forming spermogonia Group VI (type 7). Aecia
(uredo-type) with or without peridium contains reticulate, ridged, verrucose, spirally marked
aeciospores borne singly and germ pores zonate. Uredia (uredo-type) are similar to aecia except not
accompanied by spermogonia. Telia erumpent and having unicellular teliospores 1-celled, borne one
or few on each pedicel with one germ pore per spore. Basidium is external. Species are mostly
macrocyclic and autoecious. The genera of this family mostly inhabit members of Anacardiaceae
(Cummins & Hiratsuka 2003, Aime & McTaggart 2020).
Genera reported in India – Pileolaria (2); total 2 species.
Host family – Anacardiaceae.
Notes – Four genera and 43 species have been documented in this family (Wijayawardene et
al. 2020), of which two genera, Pileolaria and Skierka were reported from India. However, Aime &
McTaggart (2020) included only the genus i.e. Pileolaria and proposed the placement of Skierka
within another family. A total 25 species of Pileolaria are recorded worldwide (Index Fungorum
2021). The two Indian records (P. indica Syd. and P. pistaciae F.L. Tai & C.T. Wei) require further
molecular studies.
129
Figure 4 – Rust fungi. A Pileolaria sp. on Pistacia sp. B Skierka sp. on Pistacia sp.
Pucciniastraceae Gäum. ex Leppik, Ann. bot. fenn. 9: 139. 1972. emend. Aime & McTaggart
Type genus – Pucciniastrum G.H. Otth, Mitt. Naturforsch. Ges. Bern 1861: 71. 1861.
Pucciniastraceae members are heteroecious and mostly macrocyclic. Spermogonia and aecia
produced on conifers; uredinia and telia often on ferns. Spermogonia are of Group I (type 2 or 3).
Aecia with well developed peridium and contain catenulate aeciospores. Uredinia are formed with
cellular peridium and ostiolar opening and contain pedicellate or non pedicellate, echinulate
urediniospores, borne singly and possess obscure, scattered or bizonate germ pores. Telia are not well
differentiated i.e. either subepidermal or intradermal, not erumpent, composed of unicellular or
multicellular, sessile teliospores having obscure or 1 germ pore per cell. Basidia are external
(Cummins & Hiratsuka 2003, Aime & McTaggart 2020). The family was reported to contain 10
genera and 210 species (Wijayawardene et al. 2020). The recent higher rank classification for rust
fungi proposed the inclusion of only six genera in this family (Aime & McTaggart 2020), of which
four were recorded from India.
Genera reported in India – Hyalopsora (2), Melampsoridium (4), Peridermium (7),
Pucciniastrum (6); total 19 species.
Host families – Betulaceae, Celastraceae, Coriariaceae Magnoliaceae, Phyllanthaceae,
Pinaceae, Pteridaceae, Rosaceae, Sapindaceae.
Notes – As per the proposal to conserve the name of rust fungi (Aime et al. 2018), the sexual
name Melampsorella has been conserved over Peridermium. However, the Indian records of
Peridermium are devoid of any molecular studies and are still unchanged (Index Fungorum 2021).
Pucciniaceae Chevall., Fl. gén. env. Paris (Paris) 1: 413. 1826. emend. Aime & McTaggart
Figs 5, 6, 7, 8
Type genus – Puccinia Pers., Synopsis methodica fungorum: 225. 1801.
Pucciniaceae is a largest family that infects nearly all the major angiospermous orders. They
are destructive phytopathogens infecting mainly cereals such as wheat. Spermogonia belong to Group
V (type 4). Aecidia are with or without a peridium and have verrucose aeciospores borne singly, or
in chain (catenulate). Uredinia with palisade like peridium contain echinulate urediniospores borne
singly and have many germ pores. The most important distinguishing character is that these fungi
having stalked teliospores borne singly or united in telia with or without paraphyses. In some cases,
palisade like peridium or separation of telia into locules by stromatoid paraphyses is also observed.
130
Teliospores are mostly 1 or 2 (rarely more) celled with one germ pore in each cell and germination
mostly by external basidium (sometimes internal). The genera are mostly heteroecious exhibiting
four spore stages usually upon two or more distinct hosts followed by an independent promycelial
stage upon germination of the teliospores. The largest two genera are Puccinia, with two-celled
teliospores, and Uromyces where they are one-celled. This is the most speciose family of the
Pucciniales, and contains 21 genera and over 4961 species (Wijayawardene et al. 2020). Aime &
McTaggart (2020) proposed the inclusion of 23 genera and 9 more genera likely to include in this
family. Total 12 genera with 393 species have been recorded in India.
Genera reported in India – Caeoma (4), Chrysocelis (1), Corbulopsora (1), Endophyllum (8),
Gambleola (1), Hapalophragmium (4), Kernella (1), Puccinia (279), Pucciniosira (1),
Ramakrishnania (1), Trochodium (2), Uromyces (89); total 393 species.
Host families – Acanthaceae, Apiaceae, Asteraceae, Berberidaceae, Boraginaceae,
Celastraceae, Convolvulaceae, Cyperaceae, Elaeagnaceae, Euphorbiaceae, Fabaceae, Lamiaceae,
Lauraceae, Liliaceae, Loranthaceae, Malvaceae, Orchidaceae, Pinaceae, Plantaginaceae, Poaceae,
Polygonaceae, Ranunculaceae, Rubiaceae, Saxifragaceae, Schisandraceae, Solanaceae.
Notes – The genus Caeoma, as typified by C. berberidis, is a synonym of Puccinia (Aime et
al. 2018). One of its species C. torreyae is presented as basionym of Rogerpetersonia torreyae
(Bonar) Aime & McTaggart (Aime & McTaggart 2020) based on broad molecular assessment.
Similarly, the genus Trochodium has now been transferred to Uromyces. The types of two genera,
Gambleola and Ramakrishnania are of Indian origin and identified mainly based on
mophotaxonomic characters. The Indian records for all three genera are still unchanged (Index
fungorum 2021) and required investigation at molecular level.
Raveneliaceae Leppik, Ann. bot. fenn. 9: 139. 1972. emend. Aime & McTaggart
Fig. 9
Type genus – Ravenelia Berk., Gard. Chron. 13:132. 1853.
Raveneliaceae consists Group VI (type 5, 7) spermogonia. The genera of this family produce
typically uredo- (rarely aecidium-, caeoma-, or lecythea-) type aecia with or without peridium and
paraphyses. Aeciospores are pedicellate and echinulate or verrucose borne singly or in chain.
Uredinia are of subepidermal or erumpent, with or without paraphyses. These are similar to aecia
while unknown in some cases. Urediniospores borne singly, mostly echinulate with several germ
pores. Telia are erumpent with or without paraphyses contain pedicellate, vertically septate or
vertically or radially arranged 1- to many-celled teliospores (2 or more) on the top of pedicel, often
subtended by hygroscopic crystals or with pedicel having apical cells. Each cell or spore contains
one or two germ pores. Basidium is of external type. Members of this family have autoecious and
macrocyclic type of life cycle mostly on Fabaceae or Rosaceae. Although 24 genera and 384 species
have been documented within this family (Wijayawardene et al. 2020), Aime & McTaggart (2020)
proposed the inclusion of 16 genera and 16 more genera likely to include in this family. Total 10
genera with 61 species have been recorded in India.
Genera reported in India – Chaconia (1), Didymopsorella (1), Diorchidium (3), Gymnopuccinia
(1), Kernkampella (5), Maravalia (11), Scopella (1), Olivea (3), Prospodium (2) Ravenelia (33); total
61 species.
Host families – Fabaceae, Oleaceae, Phyllanthaceae, Poaeae, Rubiaceae, Rutaceae.
Notes – The identification of Indian records in this family, Didymopsorella macrospora
(Mundk. & Thirum.) Thirum., Gymnopuccinia pulneyensis K. Ramakr. and Scopella dalbergiae (T.S.
Ramakr. & K. Ramakr.) Ragunathan & K. Ramakr. is mainly based on their morphological
caharcters. Similarly, the genus Scopella has now been transferred to Maravalia. However, its indian
record Scopella dalbergiae (T.S. Ramakr. & K. Ramakr.) Ragunathan & K. Ramakr. is still
unchanged. DNA sequence studies can be helpful to establish their correct taxonomic placement.
Skierkaceae (Arthur) Aime & McTaggart, Fungal Systematics and Evolution 7: 21–47, 2020.
Figs 4B, 9
Type genus – Skierka Racib., Parasit. Alg. Pilze Javas (Jakarta) 2: 30. 1900.
131
Skierkaceae is a newly introduced family by Aime & McTaggart (2020), mainly characterized
by subepidermal, periphysate, deep-seated spermogonia with convex hymenium. Aecia and uredinia
uredo-type. The Uredia and telia (sporothalli sori) are deep-seated and subepidermal which
differentailed these from all other rust fungi. Urediniospores and teliospores are single-celled
produced on sporogenous cells through a narrow sorus opening. Before emenrgence, these spores
leaving behind new spores on sporogenous cells from which they are detached. Teliospores strongly
adherent, extruded in hair-like columns, germination external, without dormancy. These fungi
possess autoecious and macrocyclic type of life cycle.
Genera reported in India – Skierka (3); total 3 species.
Host families – Anacardiaceae.
Notes – A total of 14 species of the genus Skierka have been reported so far (Index Fungorum
2021), three species reported from India too. Skierka himalayensis A.K. Gautam & S. Avasthi was
reported as new from India (Gautam & Avasthi 2017c). But all three records are identified on based
on morphological characters only and require molecular identification.
Sphaerophragmiaceae Cummins & Y. Hirats., Illustr. Gen. Rust Fungi, rev. Edn (St. Paul): 15. 1983.
emend. Aime & McTaggart
Type genus – Sphaerophragmium Magnus Ber. dt. bot. Ges 9: 121. 1891.
Spermogonia are mostly lacking and unknown, Group V (type 4) in Sphenorchidium, if present.
Aecia are aecidium-type and uredinia resembling aecia (lecythea-type in Sphenorchidium).
Teliospores 2- to multicelled, pedicellate, globose to subglobose, with furcated or simple blunt wall
projections, with one germ pore per cell borne in compact telia (Beenken & Berndt 2010). In
Austropuccinia, urediniospores with a smooth patch (tonsure) as comparison to echinulate or
verrucose in other cases. Telia subepidermal to erumpent, cylindrical to ellipsoidal, with a rounded
apex, 2-celled teliospores, constricted at the septum. Basidia are mostly external. Species are
autoecious with variable life cycles. Wijayawardene et al. (2020) documented 2 genera and 25 species
in this family, however, Aime & McTaggart (2020) proposed the inclusion of 5 genera.
Genera reported in India – Sphaerophragmium (1); total 1 species.
Host families – Fabaceae.
Notes – Only single genus Sphaerophragmium with one species (S. acacia (Cooke) Magnus)
was reported from India. The identification of this species is solely based on moprhological
characters, and DNA based molecular studies are necessary.
Tranzscheliaceae (Arthur) Aime & McTaggart, Fungal Systematics and Evolution 7: 21–47, 2020
Type genus – Tranzschelia Arthur, Rés. Sci. Congr. Int. Vienne: 340. 1906.
The rust fungi of family Tranzscheliaceae are mostly macrocyclic and heteroecious in nature.
However, some microcyclic species may be found. The species consists of Group VI (type 7)
spermogonia. Aecia are of aecidium type while, uredinia uredo-type. Teliospores produced cby
theses fungi are 2-celled, pedicellate, produced from sterile basal cells.
Genera reported in India – Leucotelium (1), Tranzschelia (2); total 3 species
Host families – Rosaceae.
Notes – Tranzscheliaceae is proposed as new family by Aime & McTaggart (2020), included
two genera, Leucotelium and Tranzschelia. Both the genera were previously treated within
Uropyxidaceae (Cummins & Hiratsuka 2003). The Indian records of these genera still required DNA
sequence based studies to establish their correct taxonomic position.
Zaghouaniaceae P. Syd. & Syd., Monogr. Uredin. (Lipsiae) 3(3): 586. 1915. emend. Aime &
McTaggart
Type genus – Zaghouania Pat., Bull. Soc. mycol. Fr. 17: 187. 1901
The species of this family most often consists of deep seated and non-periphysate Group III
(type 12) spermogonia. Aecia are mostly petersonia-type (without peridium or intercalary cells).
Aeciospores are echinulate or verrucose; borne singly or in chain. The uredinia are generally
132
produced without peridium (most often uredo-type) or weakly developed peridium in some cases and
produce echinulate, singly borne ureniniospores. Telia are with or without paraphyses, produce
teliospores germinating externally (internally in some cases) by apical growth without dormancy.
These rust fungi have both autoecious and heteroecious mode of life cycle.
Genera reported in India – Cystopsora (1), Elateraceium (1), Hemileia (10), Zaghouania (1);
total 13 species
Host families – Apocynaceae, Celastraceae, Oleaceae, Penaeaceae, Phyllanthaceae,
Rubiaceae.
Notes – Aime & McTaggart (2020) included 8 genera in this family, of which, four genera
namely Cystopsora, Elateraceium, Hemileia, Zaghouania have been reported from India. The genus
Cystopsora is now considered as synonym of Zaghouania (Aime & McTaggart 2020). The identity
of Indian record Cystopsora antidesmatis T.S. Ramakr. & Sundaram is mainly based on
morphological observations. DNA sequence based studies are required to provide its exact taxonomic
position.
Figure 5 – Rust fungi Puccinia spp. A Rubia cordifolia. B Clematis sp. C Berberis sp. D Clematis
sp. E Oxalis sp. F Mentha sp.
Figure 6 – Rust fungi Uromyces spp. A Trifolium sp. B Geranium sp. C Rumex sp.
133
Figure 7 – Uredinospores. A Puccinia himachalensis on Clematis sp. B P. tiliaefolia on Grewia sp.
C P. fagopyri on Fagopyrum sp. D P. menthae on Mentha sp. E P. Oxalidis on Oxalis sp. F P. flavipes
on Duchesnea sp. G P. abrupta on Parthenium sp. H P. colletiana on Rubia sp.
I Melampsora caprearum on Salix sp. J P. Nepalensis on Rumex sp. K Uredo sp. on Ehretia sp.
L Melampsora populnea on Populus sp. Scale Bar = 10µm.
134
Figure 8 – Teliospores of Puccinia. A P. himachalensis on Clematis sp. B P. fagopyri on Fagopyrum
sp. C P. tiliaefolia on Grewia sp. D P. agrostdis on Aquilegia sp. E P. Cynodontis on Cyanodon sp.
F P. cynodontis on Cyanodon sp. G P. colletiana on Rubia sp. H P. colletiana on Rubia sp. I P.
gouriana on Clematis sp. Scale Bar = 10µm.
Excluded and replaced names of rust fungi in India
With the use of DNA sequence based techniques, the economically important rusts are
relatively well explored, but not much attention has been paid to species infecting wild plants in
general. The identification of most of the Indian rust fungi is largely based on morphological
characters especially morphology of certain spore stages. Use of DNA sequence based studies along
with morphotaxonomic characters has made identification and characterization of rust fungi more
efficient and accurate. The names of many rust genera/species as reported in the cited publications
have been replaced with new accepted names. Numbers of genera and species of Indian rust fungi
have also been replaced with currently accepted name according to MycoBank
(www.mycobank.org)/ and Species Fungorum (www.speciesfungorum.org) websites and this is
indicated in Table 3.
135
Figure 9 – Teliospores of rust fungi. A Uromyces on Trifolium sp. B Uromyces on Rumex sp.
C Uromyces on Geranium sp. D Ravenelia on Pongamia sp. E Skierka on Pistacia sp. F Kweilingia
on Bamusa sp. G Phragmidium on Rosa sp. H Pileolaria on Pistacia sp. I Melampsora on Euphorbia
sp. Scale Bar = 10µm.
Table 3 Excluded and replaced names of rust fungi in India. (Index Fungorum 2020, Mycobank
2020)
Old Name
Acervulopsora ichnocarpi (Barclay) Thirum.
Aecidium acanthospermi P.B. Chavan & Bakare
Aecidium asterum Schwein.
Aecidium barleriae M.A. Salam & Ramachar
Aecidium berberidis Pers. ex J.F. Gmel.
Aecidium cassiae Bres.
Aecidium cunninghamianum Barclay
New Name
Maravalia ichnocarpi (Barclay) Sathe
Puccinia acanthospermi Henn.
Puccinia asterum (Schwein.) F. Kern
Aecidium salamii G.F. Laundon
Puccinia graminis Pers.
Endophyllum cassiae (Bres.) F. Stevens & Mendiola
Roestelia cunninghamianum (Barclay) F. Kern
136
Table 3 Continued.
Old Name
Aecidium elaeagni-latifoliae Petch
Aecidium kaernbachii Henn.
Aecidium leucospermum DC.
Aecidium macowanianum Thüm.
Aecidium meliosmae-myrianthae Henn. & Shirai
Aecidium patulum Syd. & P. Syd.
Aecidium plectranthi Barclay
Aecidium ranunculacearum DC.
Angiopsora ampelopsidis (Dietel & P. Syd.) Thirum.
& F. Kern
Angiopsora elephantopodis (Hirats.) Mundk. &
Thirum.
Angiopsora meliosmae (Kusano) Thirum. & F. Kern
Angiopsora vernoniae T.S. Ramakr.
Bubakia cingens (Syd. & P. Syd.) S. Ito
Catenulopsora flacourtiae Mundk. & Thirum.
Catenulopsora grewiae Mundk. & Thirum.
Catenulopsora ziziphi T.S. Ramakr. & Subram.
Cerotelium fici (Castagne) Arthur
Chnoopsora butleri Dietel, Syd. & P. Syd.
Chnoopsora sancti-johannis (Barclay) Dietel
Cystopsora oleae E.J. Butler
Dasturella divina (Syd.) Mundk. & Khesw.
Dasturella grewiae (Pat. & Har.) Thirum.
Didymopsorella toddaliae (Petch) Thirum.
Elateraecium divinum (Syd.) Thirum., F. Kern &
B.V. Patil
Endophyllum tuberculatum (Ellis & Kellerm.) Arthur
& Fromme
Frommea obtusa (F. Strauss) Arthur
Gymnosporangium distortum Arthur & Cummins
Hamaspora benguetensis Syd.
Hemileia woodii Kalchbr. & Cooke
Jacksoniella holwayi (H.S. Jacks.) Kamat & Sathe
Kamatomyces narasimhanii (Thirum.) Sathe
Kuehneola aliena (Syd., P. Syd. & E.J. Butler) P.
Syd. & Syd. & E.J. Butler
Kuehneola trichosanthis (Petch) T.S. Ramakr. &
Sundaram
Kuehneola vitis (E.J. Butler) P. Syd. & Syd.
Melampsora laricis-epitea Kleb.
Nyssopsora schefflerae Ramachar, Bagyan., Subbal.
& Hosag.
Ochropsora sorbi Dietel
Oplophora cedrelae (Hori) Syd.
Phakopsora apoda (Har. & Pat.) Mains
Phakopsora gossypii (Lagerh.) Hirats. f.
Phakopsora grewiae (Pat. & Har.) Cummins
New Name
Endophyllum elaeagni-latifoliae (Petch) Gokhale,
Thirum. & Patel
Endophyllum kaernbachii (Henn.) F. Stevens &
Mendiola
Ochropsora ariae (Fuckel) Ramsb.
Endophyllum macowanianum (Thüm.) Pole-Evans
Neophysopella meliosmae-myrianthae (Henn. & Shirai)
Jing X. Ji & Kakish.
Roestelia patula (Syd. & P. Syd.) F. Kern
Coleosporium plectranthi (Barclay) Sacc.
Uromyces dactylidis G.H. Otth
Neophysopella ampelopsidis (Dietel & P. Syd.) Jing X.
Ji & Kakish.
Phakopsora elephantopodis Hirats.
Neophysopella meliosmae (Kusano) Jing X. Ji &
Kakish.
Physopella vernoniae (T.S. Ramakr.) Ramachar &
Bhagyan.
Phakopsora cingens (Syd. & P. Syd.) Hirats.
Kuehneola flacourtiae (Mundk. & Thirum.) Thirum.
Kuehneola grewiae (Mundk. & Thirum.) Thirum.
Kuehneola ziziphi (T.S. Ramakr. & Subram.) Thirum.
Puccinia fuirenicola Arthur
Chrysocelis butleri (Dietel, Syd. & P. Syd.) G.F.
Laundon
Melampsora sancti-johannis Barclay
Zaghouania oleae (E.J. Butler) Cummins
Kweilingia divina (Syd.) Buriticá
Uredopeltis chevalieri J. Walker & R.G. Shivas
Skierka toddaliae (Petch) Hirats.,
Kweilingia divina (Syd.) Buriticá
Pucciniosira tuberculata (Ellis & Kellerm.) Buriticá &
J.F. Hennen
Phragmidium potentillae (Pers.) P. Karst.
Roestelia distorta (Arthur & Cummins) F. Kern
Hamaspora rubi-sieboldii (Kawagoe) Dietel
Puccinia woodii (Kalchbr. & Cooke) P. Syd. & Syd.
Phragmidiella holwayi (H.S. Jacks.) Buriticá
Masseeella narasimhanii Thirum.
Phragmidiella aliena (Syd., P. Syd. & E.J. Butler)
Buriticá & J.F. Hennen
Cerotelium trichosanthis (Petch) Nag Raj, Govindu &
Thirum.
Chrysomyxa vitis E.J. Butler
Melampsora epitea Thüm.
Nyssopsora thwaitesii (Berk. & Broome) Syd.
Ochropsora ariae (Fuckel) Ramsb.
Nyssopsora cedrelae (Hori) Tranzschel
Angiopsora apoda (Har. & Pat.) Aime & McTaggart.
Phakopsora desmium (Berk. & Broome) Cummins
Uredopeltis chevalieri J. Walker & R.G. Shivas
137
Table 3 Continued.
Old Name
Phakopsora tecta H.S. Jacks. & Holw.
Phakopsora vignae (Bres.) Arthur
Phakopsora ampelopsidis Dietel & P. Syd.
Phragmidium disciflorum (Tode) J. James
Phragmidium rubi (Pers.) G. Winter
Puccinia absinthii DC.
Puccinia abutili Berk. & Broome
Puccinia allii (DC.) F. Rudolphi
Puccinia artemisiella P. Syd. & Syd.
Puccinia arthraxonis (Henn.) Syd., P. Syd. & E.J.
Butler
Puccinia bullata (Pers.) J. Schröt.
Puccinia calcitrapae var. centaureae (DC.) Cummins
Puccinia carduorum Jacky
Puccinia caricis Rebent.
Puccinia coronata f. agrostidis Erikss.
Puccinia coronata var. himalensis Barclay
Puccinia dispersa Erikss. & Henning
Puccinia epilobii-tetragoni G. Winter
Puccinia extensicola Plowr.
Puccinia glumarum (J.C. Schmidt) Erikss. & Henning
Puccinia graminis var. tritici Erikss. & Henning
Puccinia himalensis (Barclay) Dietel
Puccinia leptodermidis (Barclay) Sacc.
Puccinia levis var. panici-sanguinalis (Rangel)
Ramachar & Cummins
Puccinia lolii E. Nielsen
Puccinia lycoctoni Fuckel
Puccinia maydis Bérenger
Puccinia obtegens (Link) Tul.
Puccinia orientalis (Syd., P. Syd. & E.J. Butler)
Arthur & Cummins
Puccinia orientalis (Syd., P. Syd. & E.J. Butler)
Arthur & Cummins
Puccinia penniseti Zimm.
Puccinia persistens Plowr.
Puccinia poae-nemoralis G.H. Otth
Puccinia polygoni Alb. & Schwein.
Puccinia prunicolor Syd., P. Syd. & E.J. Butler
Puccinia pruni-persicae Hori
Puccinia pruni-spinosae Pers.
Puccinia pulsatillae Kalchbr.
Puccinia punctiformis (F. Strauss) Röhl.
Puccinia rubigo-vera var. tritici (Erikss.) Carleton
Puccinia rubigo-vera (DC.) G. Winter
Puccinia saxifragae-micranthae Barclay
Puccinia substriata var. indica Ramachar & Cummins
Puccinia substriata var. penicillariae (Speg.)
Ramachar & Cummins
Puccinia taraxaci Plowr.
Puccinia tricholaenae (Syd. & P. Syd.) T.S. Ramakr.
& K. Ramakr.
Puccinia triticina Erikss.
New Name
Uromyces spegazzinii (De Toni) Arthur
Phakopsora pachyrhizi Syd. & P. Syd.
Neophysopella ampelopsidis (Dietel & P. Syd.) Jing X. Ji
& Kakish.
Phragmidium mucronatum (Pers.) Schltdl.
Phragmidium bulbosum (Fr.) Schltdl.
Puccinia chrysanthemi Roze
Puccinia abutilonis Berk. & Broome
Puccinia porri (Sowerby) G. Winter
Puccinia tanaceti DC.
Kuehneola loeseneriana (Henn.) H.S. Jacks. & Holw.
Puccinia angelicae (Schumach.) Fuckel
Puccinia carthami Corda
Puccinia calcitrapae DC.
Puccinia dioicae Magnus
Puccinia coronata Corda
Puccinia coronata Corda
Puccinia recondita Roberge ex Desm.
Puccinia pulverulenta Grev.
Puccinia dioicae Magnus
Puccinia striiformis Westend.
Puccinia graminis Pers.
Chrysomyxa himalensis Barclay
Coleosporium leptodermidis (Barclay) P. Syd. & Syd.
Uromyces panici-sanguinalis Rangel
Puccinia coronata Corda
Uromyces dactylidis G.H. Otth
Puccinia sorghi Schwein.
Puccinia suaveolens (Pers.) Rostr.
Diorchidium orientale Syd., P. Syd. & E.J. Butler
Diorchidium orientale Syd., P. Syd. & E.J. Butler
Puccinia substriata Ellis & Barthol.
Puccinia recondita Roberge ex Desm.
Puccinia brachypodii G.H. Otth
Puccinia polygoni-amphibii Pers.
Puccinia purpurea Cooke
Leucotelium pruni-persicae (Hori) Tranzschel
Tranzschelia pruni-spinosae (Pers.) Dietel
Puccinia ustalis Berk.
Puccinia suaveolens (Pers.) Rostr.
Puccinia striiformis Westend.
Puccinia recondita Roberge ex Desm.
Puccinia heucherae (Schwein.) Dietel
Puccinia substriata Ellis & Barthol.
Puccinia substriata Ellis & Barthol.
Puccinia hieracii (Röhl.) H. Mart.
Diorchidium tricholaenae Syd. & P. Syd.
Puccinia recondita Roberge ex Desm.
138
Table 3 Continued.
Old Name
Puccinia anomala Rostr.
Puccinia anthistiriae Barclay
Puccinia baryi (Berk. & Broome) G. Winter
Puccinia brizae-maximae T.S. Ramakr.
Puccinia bupleuri-falcati (DC.) G. Winter
Puccinia cichorii Bellynck ex J. Kickx f.
Ravenelia berkeleyi Mundk. & Thirum.
Ravenelia breyniae Syd. & P. Syd.
Ravenelia breyniae-patentis Mundk. & Thirum.
Ravenelia emblicae Syd. & P. Syd.
Ravenelia evernia Syd.
Ravenelia kirganeliae Mundk. & Thirum.
Ravenelia phyllanthi Mundk. & Thirum.
Scopella aulica (Syd.) Mundk. & Thirum.
Scopella echinulata (Niessl) Mains
Scopella fici Mundk. & Thirum.
Scopella gentilis (Syd.) Mundk. & Thirum.
Scopella mimusops (Cooke) Cummins
Stakmania formosana (Syd. & P. Syd.) Sathe
Teloconia rosae (Kuntze) Syd.
Thekopsora gaultheriae (Syd. & P. Syd.) P. Syd. &
Syd.
Trachyspora intrusa (Grev.) Arthur
Tranzschelia punctata Arthur
Tunicopsora bagchii Suj. Singh & P.C. Pandey
Uredo artocarpi Berk. & Broome
Uredo bombacis Petch
Uredo callicarpae Petch
Uredo colebrookeae Barclay
Uredo fici Castagne
Uredo gaultheriae (Syd. & P. Syd.) Hirats. f.
Uredo ipomoeae Yadav
Uredo plumeriiae Pravenna, Nasheema & Balakrishna
Uredo tephrosiicola Henn.
Uredo ziziphi Pat.
Uredopeltis boswelliae (Patel, Payak & N.B. Kulk.)
Sathe
Uredopeltis boswelliae (Patel, Payak & N.B. Kulk.)
Sathe
Uromyces fabae (Pers.) de Bary
Uromyces leptodermus Syd. & P. Syd.
Uromyces linearis Berk. & Broome
Uromyces lycoctoni (Kalchbr.) Trotter
Uromyces phaseoli G. Winter
Uromyces pisi (DC.) G.H. Otth
Uromyces scillarum (Grev.) G. Winter
Uromyces scirpi Burrill
Uromyces sojae (Henn.) Syd. & P. Syd.
Xenostele indica Thirum.
New Name
Puccinia hordei G.H. Otth
Puccinia graminis Pers.
Puccinia brachypodii G.H. Otth
Puccinia graminis Pers.
Puccinia bupleuri F. Rudolphi
Puccinia hieracii (Röhl.) H. Mart.
Ravenelia cassiicola G.F. Atk.
Kernkampella breyniae (Syd. & P. Syd.) Rajendren
Kernkampella breyniae-patentis (Mundk. & Thirum.)
Rajendren
Kernkampella emblicae (Syd. & P. Syd.) G.F. Laundon
Ravenelia fragrans Long
Kernkampella kirganeliae (Mundk. & Thirum.) G.F.
Laundon
Kernkampella phyllanthi (Mundk. & Thirum.) G.F.
Laundon
Maravalia aulica (Syd.) Y. Ono
Maravalia echinulata (Niessl ex Rabenh.) Y. Ono
Maravalia fici (Mundk. & Thirum.) Y. Ono
Maravalia gentilis (Syd.) Y. Ono
Maravalia mimusops (Cooke) Y. Ono
Phakopsora formosana Syd. & P. Syd.
Phragmidium kamtschatkae (H.W. Anderson) Arthur &
Cummins
Pucciniastrum gaultheriae Syd. & P. Syd.
Trachyspora alchemillae (Pers.) Fuckel
Tranzschelia pruni-spinosae (Pers.) Dietel
Kweilingia bagchii (Suj. Singh & P.C. Pandey) Buriticá
Physopella artocarpi (Berk. & Broome) Arthur
Calidion bombacis (Petch) D.J. Soares & R.W. Barreto
Uromyces callicarpae (Petch) Fujik. ex S. Ito
Olivea colebrookeae (Barclay) Thirum. & Yadav
Cerotelium fici (Castagne) Arthur
Pucciniastrum gaultheriae Syd. & P. Syd.
Coleosporium ipomoeae (Schwein.) Burrill
Coleosporium plumeriae Pat.
Ravenelia tephrosiicola (Henn.) Hirats. f.
Macabuna ziziphi (Pat.) Buriticá & J.F. Hennen
Dasturella boswelliae Patel, Payak & N.B. Kulk.
Dasturella boswelliae Patel, Payak & N.B. Kulk.
Uromyces viciae-fabae (Pers.) J. Schröt.
Uromyces setariae-italicae Yoshino
Puccinia peradeniyae Demers & Castl.
Uromyces dactylidis G.H. Otth
Uromyces appendiculatus (Pers.) Link
Uromyces pisi-sativi (Pers.) Liro
Uromyces muscari (Duby) Niessl
Uromyces lineolatus (Desm.) J. Schröt.
Phakopsora pachyrhizi Syd. & P. Syd.
Xenostele litseae (Pat.) Syd. & P. Syd.
139
Discussion
This study provides an outline for rust fungi of India based on the literature. It provides
complete information of Indian Pucciniales in one compilation as 69 genera and 640 species
belonging to 16 families. Rust fungi are one of the extensively studied fungal groups of India, as
evident from the number of researchers who have investigated these fungi since pre-independance.
The outline presented in this study helps to better understand the taxonomy of Indian rust fungi. In
addition to broadly studied fungal group of India, rust fungi possessed a broad host range and
distribution too. As per earlier reports, rust fungi cause diseases on various plant hosts (Misra et al.
1975, Bisht & Srivastava 1990, Cummins & Hiratsuka 2003, Jiao et al. 2016). High relative humidity
and dense forest cover might be the possible reasons to promote these rust fungi in these regions to
cause diseases. The occurrence of 167 species of rust fungi belonging to 23 genera and 11 families
on 170 plant species belonging to 52 families from Himachal Pradesh justified their diversity in this
hilly state. Similarly, the broader host range from Poaceae with highest number of records followed
by Ranunculaceae, Rosaceae, Asteraceae, Polygonaceae, Fabaceae, Salicaceae, Acanthaceae,
Lamicaeae, Pinnaceae, Apiaceae, Rubiaceae, Saxifragaceae, Cyperaceae, Euphorbiaceae,
Berberidiaceae, Geraniaceae, Linaceae and Zinziberaceae support a wide distribution of these fungi
(Gautam & Avasthi 2019). The occurence of 12 rust genera with 35 species belonging to 7 families
on large number of herbaceous, shrubby plants including climbers, grass and trees also support the
diversity and distribution of rust fungi in Himalayan regions (Singh & Palni 2011). However, these
fungi are not only limited to hilly regions of India; Mohanan (2010) documented a total of 95 rust
fungi belonging to 25 genera associated with 117 forest plant species belonging to 80 host genera
under 43 host families from the Western Ghats regions of Kerala. Similarly, the checklist of the rust
genus Uromyces was documented by Gautam & Avasthi (2017b), also support broad diversity of
these fungi on wide-ranging host range.
Besides the diversity and distribution, rust fungi cause very devastating diseases on various
agricultural crops in India. Wheat rusts caused by three species of Puccinia namely, stripe rust (by
P. striiformis f. sp. tritici Westend.), stem rust (by P. graminis Pers. f. sp. tritici Eriks. & Henn.) and
leaf rust (caused by P. triticina Eriks.) pose a threat to global wheat production. The detection of
Ug99 led the global community to work together to combat this disease, re-emerged as a threat and
the establishment of the Borlaug Global Rust Initiative (BGRI, earlier Global Rust Initiative) in
September 9, 2005. Pathotyping of rust pathogens and their identification are some important steps
being deployed by researchers to find out management strategies of rust diseases. Emphasis is laid
on evaluation of germplasm for rust resistance and development of various rust resistance varieties
through various breeding programmes. In comparison to other crops, wheat has achieved a record
high production in India during the current century. The development of rust resistance varieties has
also progressed which played a vital role in protecting wheat from any epidemic threat (Tomar et al.
2014). The ICAR scientists cracked the whole genome of Puccinia triticina which is now proving
very helpful in understanding the nature of this wheat rust pathogen (Kiran et al. 2016). Similarly,
about 22 varieties of wheat which are are resistant to the deadly Ug99 fungal disease caused by
Puccinia graminis- tritici are developed, of which, some of the varieties such as DBW 17, PBW 550,
and Lok 1 are being cultivated in wheat-growing states in India (Bhardwaj et al. 2019).
Both morphological and molecular characterization of rust fungi is required to understand the
natural classification and evolutionary relationship of rust fungi. Because with the difficulty of
culturing of these fungi artificially, the success rate of culturing is also not so encouraging. The direct
sequencing of rust fungi is also not so easy because of the isolation of DNA of other microbes
associated with main rust pathogens. This may affects the quality and purity of DNA and ultimately
the sequencing process and final identification. Due to all the above reasons, the information on
molecular identification of Indian rust fungi is not adequate. Although a phylogeny of Indian rust
fungi based on the sequence data of LSU and ITS available for corresponding rust fungi in GenBank
(NCBI) is presented, the lack of molecular data for most of the rust fungi included requires the
application of molecular techniques. Our phylogenetic studies based on LSU and ITS sequence data
showed that few taxa of family Pucciniaceae include polyphetic taxa i.e. from Puccinia and
140
Uromyces. More studies are still required for the better understanding of their taxonomic placement
at different levels. Hence, fresh collections are required to generate molecular data to understand
their phylogenetic relationships. This study has set the foundation for the systematics and taxonomic
studies of rust fungi in India at generic and species level.
Although 69 genera of Indian rust fungi were included in this study, very few of these are
known to have DNA sequence data. Molecular studies of these fungi are still scanty and there is much
scope for exploratory work on this fungal group. Due to the lack of molecular studies, many genera
or species need to be recollected and epitypifed, in order to place them in their correct taxonomic
position. Some Indian rust fungi require much attention as there is confusion in their correct
taxonomic placement. Therefore, future works are likely to focus on reassessing the samples of rust
fungi reported from India on both morphological and molecular characterization. Emphasis should
also be given on understanding the relationship between rust pathogens and host preference to
elaborate this fungal group more precisely. In addition, a digital web based platform should be
developed which help the researchers to identify Indian Pucciniales and to provide all information
on their diversity, distribution and host association.
Acknowledgements
Authors are grateful to their respective organizations for providing the necessary laboratory
facilities and valuable support during the study. We also thank the Anonymous reviewers for their
helpful comments and suggestions.
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