A Global Overview of Diversity and Phylogeny of the Rust Genus Uromyces

Abstract

Uromyces is the second-largest plant pathogenic rust genus, is responsible for numerous diseases, and has major effects on both agricultural and non-agricultural plants. The genus is generally characterized by its unicellular teliospores that help to characterize it and distinguish it from another important rust genus, Puccinia. In this study, a global overview of the diversity and distribution of Uromyces is presented based on both online and offline resources. The information obtained was analyzed for numerical and graphical summaries to provide the diversity and distribution of the genus by country and continent. Besides this, broad taxonomical aspects, a brief life cycle, and other comparative aspects on diversity and distribution were also provided. In addition, a phy-logenetic analysis based on the ITS and nLSU DNA sequence data available in GenBank and published literature was performed to examine the intergeneric relationships of Uromyces. The results obtained revealed that the rust genus is found distributed over 150 countries, territories, and occu-pancies of the world on around 647 plant genera belonging to 95 plant families. Phylogenetic studies based on LSU and ITS sequence data revealed that Uromyces species are polyphyletic and require more DNA-based analyses for a better understanding of their taxonomic placement.

Full text

Citation: Gautam, A.K.; Avasthi, S.;
Verma, R.K.; Sushma; Niranjan, M.;
Devadatha, B.; Jayawardena, R.S.;
Suwannarach, N.; Karunarathna, S.C.
A Global Overview of Diversity and
Phylogeny of the Rust Genus
Uromyces.J. Fungi 2022,8, 633.
https://doi.org/10.3390/jof8060633
Academic Editor: Liangdong Guo
Received: 26 April 2022
Accepted: 6 June 2022
Published: 14 June 2022
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4.0/).
Fungi
Journal of
Review
A Global Overview of Diversity and Phylogeny of the Rust
Genus Uromyces
Ajay Kumar Gautam 1, * , Shubhi Avasthi 2,* , Rajnish Kumar Verma 3, Sushma 4, Mekala Niranjan 5,6 ,
Bandarupalli Devadatha 6, Ruvishika S. Jayawardena 7, Nakarin Suwannarach 8, *
and Samantha C. Karunarathna 9
1School of Agriculture, Abhilashi University, Mandi 175028, Himachal Pradesh, India
2School of Studies in Botany, Jiwaji University, Gwalior 474011, Madhya Pradesh, India
3Department of Plant Pathology, Punjab Agricultural University, Ludhiana 141004, Punjab, India;
vermarajnish1985@gmail.com
4Department of Biosciences, Chandigarh University, Gharuan 140413, Punjab, India; smunihal@gmail.com
5Department of Botany, Rajiv Gandhi University, Rono Hills, Doimukh,
Itanagar 791112, Arunachal Pradesh, India; neeru436@gmail.com
6Fungal Biotechnology Lab, Department of Biotechnology, School of Life Sciences, Pondicherry University,
Kalapet 605014, Pondicherry, India; devadatha796@gmail.com
7Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand;
ruvishika.jay@mfu.ac.th
8Research Center of Microbial Diversity and Sustainable Utilization, Chiang Mai University,
Chiang Mai 50200, Thailand
9
Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and
Food Engineering, Qujing Normal University, Qujing 655011, China; samanthakarunarathna@gmail.com
*Correspondence: a2gautam2006@gmail.com (A.K.G.); shubh.avasth@gmail.com (S.A.);
suwan.462@gmail.com (N.S.); Tel.: +91-7018321256 (A.K.G.); +6686-5127518 (N.S.)
Abstract:
Uromyces is the second-largest plant pathogenic rust genus, is responsible for numerous
diseases, and has major effects on both agricultural and non-agricultural plants. The genus is generally
characterized by its unicellular teliospores that help to characterize it and distinguish it from another
important rust genus, Puccinia. In this study, a global overview of the diversity and distribution of
Uromyces is presented based on both online and offline resources. The information obtained was
analyzed for numerical and graphical summaries to provide the diversity and distribution of the
genus by country and continent. Besides this, broad taxonomical aspects, a brief life cycle, and other
comparative aspects on diversity and distribution were also provided. In addition, a phylogenetic
analysis based on the ITS and nLSU DNA sequence data available in GenBank and published
literature was performed to examine the intergeneric relationships of Uromyces. The results obtained
revealed that the rust genus is found distributed over 150 countries, territories, and occupancies of
the world on around 647 plant genera belonging to 95 plant families. Phylogenetic studies based
on LSU and ITS sequence data revealed that Uromyces species are polyphyletic and require more
DNA-based analyses for a better understanding of their taxonomic placement.
Keywords: phylogeny; Pucciniomycotina; rust fungi; taxonomy; Uromyces
1. Introduction
Uromyces (Link) Unger, a genus of rust fungi, was proposed by Unger (1833). The
genus contains several important plant pathogens, parasitizes in both monocots and dicots
throughout the world, and affects a range of crops, causing a varying amount of yield
loss annually, with losses being very severe in many cases. This genus shares synonymy
with numerous genera of fungi such as Alveomyces Bubák., Capitularia Rabenh., Coeomurus
Gray., Dichlamys Syd. & P. Syd., Groveola Syd., Haplopyxis Syd. & P. Syd., Haplotelium Syd.,
Hypodermium subgen. Uromyces Link., Klebahnia Arthur., Nielsenia Syd., Poliotelium Syd.,
J. Fungi 2022,8, 633. https://doi.org/10.3390/jof8060633 https://www.mdpi.com/journal/jof

J. Fungi 2022,8, 633 2 of 99
Poliotelium Syd., Pucciniola L. Marchand., Teleutospora Arthur & Bisby., Telospora Arthur.,
Trochodium Syd. & P. Syd., and Uromycopsis Arthur. Hypodermium subgen. Uromyces Link
1816 has been considered the Basionym of this genus [
1
]. Uromyces is the second largest
genus of rust fungus after Puccinia in the family Pucciniaceae, which contains more than
1568 epithets and more than 800 reported species [
1
–
5
]. This genus produces unicellular
teliospores that help to characterize it and distinguish it from another important rust
genus, Puccinia.
Species of Uromyces occur on a wide variety of plant hosts around the world. The most
important ones are caused by Uromyces spp. that cause infections and damage to various
agricultural crops such as alfalfa or lucerne rust (Uromyces straitus J. Schröt.), bean rust
[Uromyces appendiculatus (Pers.) Unger], beet rust [Uromyces betae (Pers.) Tul.], carnation
rust [Uromyces dianthi (Pers.) Niessl], chickpea rust (Uromyces ciceris-aerietinus Jacz.), clover
rust (Uromyces trofolii-repentis Liro), and pea rust [Uromyces pisi (DC.) G.H. Otth.] [
2
]. The
Uromyces species was recorded on the host plants belonging to the families Asteraceae,
Euphorbiaceae,Fabaceae,Liliaceae,Loranthaceae, and Poaceae, and among these, Loranthaceae is
known to be the most affected. However, the known distribution of most Uromyces species
on Loranthaceae is restricted to Mexico and Central and South America [6].
The present manuscript is focused on providing a global overview of the diversity and
distribution of the genus Uromyces. Based on the combined ITS and LSU DNA sequence
data available in GenBank and published literature, phylogenetic analyses were performed
to examine the intergeneric relationships of Uromyces. In addition, broad taxonomical
aspects, a brief life cycle, and other comparative aspects of its diversity and distribution are
also provided.
2. Taxonomy of the Genus Uromyces
The genus Uromyces belongs to the phylum Basidiomycota, the second-largest [
7
] phy-
lum after Ascomycota in the kingdom Fungi; both phyla are considered higher fungi and
share 97% of all fungal species [
3
,
8
–
11
]. Taxonomically, Uromyces contains rusts under the
class Pucciniomycetes and order Pucciniales. Based on telial morphology, the spermogonial
and aecial type, basidiospores, and uredinial characteristics, 13 rust families were proposed
by Cummins and Hiratsuka [
2
]. However, Aime [
12
] and Aime and McTaggart [
5
] proposed
several changes in basal systematics of rust fungi based on molecular studies along with
morphological characteristics. This resulted in the introduction of many new families and
the transmission of many genera and species. The Wijayawardene et al. [
3
] included 21 gen-
era with Uromyces and 4961 species in the rust family Pucciniaceae. The recent higher-level
classification for rust fungi provided recently by Aime and McTaggart [
5
] proposed the
inclusion of 23 genera (including Uromyces) and 9 likely belonging to the family Pucciniaceae.
The genus Uromyces was described by Unger in 1833 with Uromyces appendiculatus Pers. as
the type species of the genus. Uredo appendiculatus Pers. was considered as a basionym for
the type species, now regarded as Uromyces appendiculatus (Pers.) Link 1816. The detailed
taxonomy of the genus Uromyces is described as follows:
Uromyces (Link) Unger, Exanth. Pflanzen (Wien): 277 (1833):
=Alveomyces Bubák, Annln K.K. naturh. Hofmus. Wien 28 (1–2): 190 (1914);
=Capitularia Rabenh., Bot. Ztg. 9 (25): 449 (1851);
=Coeomurus Gray, Nat. Arr. Brit. Pl. (London) 1: 541 (1821);
=Dichlamys Syd. & P. Syd., Annlsmycol. 17(2/6): 105 (1920), (1919);
=Groveola Syd., Annlsmycol. 19 (3–4): 173 (1921);
=Haplopyxis Syd. & P. Syd., Annlsmycol. 17 (2/6): 105 (1920), (1919);
=Haplotelium Syd., Annlsmycol. 20 (3/4): 124 (1922);
=
Hypodermium subgen. Uromyces Link, Mag. Gesell. naturf. Freunde, Berlin 8: 28
(1816), (1815);
=Klebahnia Arthur, Résult. Sci. Congr. Bot. Wien 1905: 345 (1906);
=Mapea Pat., Bull. Soc. Mycol. Fr. 22: 46 (1906);
=Nielsenia Syd., Annlsmycol. 19 (3–4): 171 (1921);

J. Fungi 2022,8, 633 3 of 99
=Nigredo (Pers.) Roussel, Fl. Calvados, Edn 2: 47 (1806);
=
Peridipes Buriticá& J.F. Hennen, RevtaAcad. Colomb. Cienc. Exact. Fís. Nat. 19 (no.
72): 50 (1994);
=Poliotelium Syd., Annlsmycol. 20 (3/4): 124 (1922);
=Puccinella Fuckel, Jb. nassau. Ver. Naturk. 15: 18 (1860);
=Pucciniola L. Marchand, Bijdr. Natuurk. Wetensch. 4: 47 (1829);
=Rubigo (Pers.) Roussel, Fl. Calvados, Edn 2: 46 (1806);
=Teleutospora Arthur & Bisby, Bull. Torrey bot. Club 48: 38 (1921);
=Telospora Arthur, Résult. Sci. Congr. Bot. Wien 1905: 346 (1906);
=Trichobasis Lév., in Orbigny, Dict. Univ. Hist. Nat. 12: 785 (1849);
=Trochodium Syd. & P. Syd., Annlsmycol. 17(2/6): 106 (1920) (1919);
=Uredo Pers., Syn. meth. fung. (Göttingen) 1: 214 (1801);
=Uromycopsis Arthur, Résult. Sci. Congr. Bot. Wien 1905: 345 (1906).
Classification:
Pucciniaceae,Pucciniales,Pucciniomycetes,Pucciniomycotina,Basidiomycota,Fungi
Type of species:
Uromyces appendiculatus Pers., Observ. Mycol. (Lipsiae) 1: 17 (1796).
The characteristic one-celled teliospores of the genus Uromyces distinguish it from Puc-
cinia, which has two-celled teliospores. Spermogonia subepidermal, Group V (type 4). Aecia
subepidermal, erumpent, Aecidium-type (with peridium and catenuate, mostly verrucose
aeciospores) or Uredo-type (with mostly echinulate aeciospores borne singly on pedicels).
Uredinia, mostly the Uredo type, is subepidermal and erumpent and consists of echinu-
late urediniospores with various pores, borne singly on pedicels. Telia are subepidermal,
erumpent, or remains covered by the epidermis with pedicellate, one-celled teliospores
borne singly on pedicels with a mostly pigmented wall. Basidia is external in nature. The
species of Uromyces produce macrocyclic and heteroecious modes of the life cycle, as well
as many endocyclic, microcyclic, or autoecious species [2,5].
The phylogenetic analyses of different DNA loci revealed that Uromyces, along with
the genus Puccinia in the family Pucciniaceae, is not monophyletic [
5
]. The generic name
of Uromyces was recommended to be protected over Uredo [
13
]. Uredo betae, as one of
the initially included species of this genus when described in 1801 by Persoon, is now
considered a synonym of Uromyces beticola. Since Uromyces appendiculatus and U. beticola
are regarded as congeneric, Uromyces and Uredo are synonyms. Previously, the name Uredo
was used in cases where sexual morph was lacking and the species could not be described
in the “correct” sexually typified genus. The reason why the name Uromyces was protected
over Uredo is due to two major reasons: Uredo belongs to many different rust genera, and
secondly, Uromyces is a commonly used generic name. The generic name Uromyces was
already conserved over Coeomurus Link ex Gray 1821 and Pucciniola L. Marchand 1829 [
13
].
3. Biology, Pathogenicity, and Life Cycle of Uromyces
Rusts represent one of the largest groups of plant pathogens and have historically
posed a major threat to farmers around the world. These diversified fungi are obligate
biotrophic parasites distributed in all geographical areas on a wide range of wild and
cultivated plants. The two major plant groups, cereals and legumes, suffer the most from
rust infections [
14
]. The genus Uromyces consists of a number of plant pathogens responsible
for many harmful plant diseases.
3.1. Disease Symptoms of Uromyces
Uromyces, as a pathogen, affects major hosts such as alfalfa (Medicago sativa), bean
(Phaseolus vulgaris), carnation (Dianthus caryophyllus), chickpea (Cicer arietinum), clover (Tri-
folium sp.), and pea (Pisum sativum) belonging to plant families, Fabaceae and
Poaceae [2,15].
The rust diseases caused by Uromyces spp. are characterized by numerous small, rust-like
orange/yellow or brown pustules formed on infected plant tissues. Uromyces, as a plant
pathogen, kills the diseased plant cells and forms a light yellow halo around diseased

J. Fungi 2022,8, 633 4 of 99
pustules. Dark brown to black-brown telia are usually formed by the rust fungi during
infection. The severity of the disease causes a loss of photosynthetic area in infected plants,
resulting in a reduction in overall plant performance. Since rust disease is affected by
low temperatures and heavy rain in spring and autumn, this can affect the severity of the
disease. The rust diseases caused by Uromyces are generally autoecious (one plant host)
and macrocyclic (produce five spore stages in the life cycle) in nature [
2
]. The appearance
of rust disease symptoms varies in different hosts. A single species of Uromyces can infect
multiple hosts and cause variable symptoms. The rust on pea (Pisum sativum) caused by
Uromyces pisi and Uromyces viciae-fabae produce minute, whitish, slightly raised spots that
enlarge and rupture the epidermis to produce reddish-brown, irregular pustules on the
stems, pods and lower surface of leaves. Initially, the pustules contain abundant powdery
urediniospores, but eventually, they turn dark brown to black as overwintering teliospores
are produced [
16
]. Similarly, Uromyces viciae-fabae is also reported to cause lentil rust with
disease symptoms of circular yellowish-white pycnidia and aecial cups on leaflets, which
later become oval to circular, brown uredial pustules; and dark brown to black, elongated
telia [
17
]. Such variability in disease symptoms caused by different Uromyces species can
also be observed in other crops (Figure 1).
J. Fungi 2022, 8, x FOR PEER REVIEW 5 of 88
Figure 1. Disease symptoms of species of Uromyces. (A,B) Uromyces ambiens on Buxus wallichiana;
(C–E) Uromyces fabae on Pisum sativum; (F,G) Uromyces geranii on Geranium sp.; (H,I) Uromyces trifolii
on Trifolium sp. Scale bar: A–I = 1 mm.
After the aecium is fully developed, the mature aeciospores infect the host surface by
germinating and producing infection structures. Eventually, this infection leads to the
production of uredia with urediospores. With the repeated infection of the host plant,
these urediospores are produced in large quantities during the summer. Surprisingly,
urediospores can disseminate thousands of kilometers with the help of the wind. When
suitable hosts are found, these spores differentiate into telia, which ultimately produce
unicellular diploid teliospores in the winter [17,18]. A general life cycle of the rust genus
Uromyces is presented in Figure 2 below.
3.3. The Rusts as Classical Biocontrol Agents
Uromyces is the causal agent of rust disease on numerous agricultural, horticultural,
and forest plantations. Rust fungi of this genus are considered a major economic threat
due to possible yield losses from reduced production. Rust diseases on pea, beans, lentils,
polyhouse flower crops, clover, and many more are some of the hosts infected by this
genus. In addition to their identity as the causative agent of rust, the Uromyces species also
shows potential for the biological control of various phytopathogenic fungi, weeds, etc.
Figure 1.
Disease symptoms of species of Uromyces. (
A
,
B
)Uromyces ambiens on Buxus wallichiana;
(
C
–
E
)Uromyces fabae on Pisum sativum; (
F
,
G
)Uromyces geranii on Geranium sp.; (
H
,
I
)Uromyces trifolii
on Trifolium sp. Scale bar: A–I= 1 mm.

J. Fungi 2022,8, 633 5 of 99
3.2. Life Cycle
The species of rust genus Uromyces are generally macrocyclic in nature, i.e., exhibit all
five spore forms known in Pucciniales. In addition, these fungi exhibit an autoecious life
cycle, meaning that all spore forms are produced on a single host. However, the endocyclic
or microcyclic nature with a heteroecious mode of the life cycle was observed in some
species of Uromyces [
2
,
5
]. The general life cycle process of Uromyces spp. involves the
germination of diploid teliospores in the spring with a metabasidium after overwintering
on plant debris. The metabasidium produces four haploid basidiospores after meiosis.
These basidiospores with two different mating types germinate and start infection by
producing different infection structures on the surface of the host plant. Once an infection
is established, the production of pycnia containing pycniospores of two mating types
and receptive hyphae takes place. After, the spermatization of pycniospores of pycnia
of different mating types and subsequent dikaryotization takes place in aecial primordia,
along with a subsequent exchange.
After the aecium is fully developed, the mature aeciospores infect the host surface
by germinating and producing infection structures. Eventually, this infection leads to the
production of uredia with urediospores. With the repeated infection of the host plant,
these urediospores are produced in large quantities during the summer. Surprisingly,
urediospores can disseminate thousands of kilometers with the help of the wind. When
suitable hosts are found, these spores differentiate into telia, which ultimately produce
unicellular diploid teliospores in the winter [
17
,
18
]. A general life cycle of the rust genus
Uromyces is presented in Figure 2below.
J. Fungi 2022, 8, x FOR PEER REVIEW 6 of 88
The biology and effectiveness of Uromyces heliotropii were studied by Hasan and Aracil
[19], where they found the rust to effectively control an annual weed heliotrope
(Heliotropium europaeum L.).
Figure 2. A general life cycle of rust genus Uromyces.
They evaluated the biocontrol potential of the rust in a greenhouse and field
inoculation experiments and observed that the rust rapidly killed infected plants and
reduced or prevented seed production. Similarly, Anderson et al. [20] conducted a study
to evaluate the potential of three rusts naturally infecting Chilean needle grass (Nassella
neesiana) in Argentina, Uromyces pencanus, Puccinia graminella, and P. nassellae, as
biocontrol agents. They found U. pencanus to be most effective due to the damage it inflicts
on its host in the field. A list of species of Uromyces studied as biocontrol agents is given
in Table 1.
Table 1. List of species of Uromyces with biocontrol potential.
Species of Uromyces Target Organism )Plants( Reference
Uromyces rumicis Rumex ckpus [21]
Uromyces galegae Galega officinalis [22]
Uromyces heliotropii Heliotropiu meuropaeum [19]
Uromyces pencanus Nassella neesiana [20,23–25]
Uromyces pisi-sativi Cytisus scoparius [26]
Uromyces pisi-sativi Chamaecytisus palmensis [26]
Uromyces pisi-sativi Lupinus polyphyllus [26]
Rust fungi as biocontrol agents are mainly studied in their use against plant weeds.
The biocontrol ability of some species of Puccinia was investigated, such as as P. abrupta
var. partheniicola on different growth stages of Parthenium hysterophorus [27,28], Puccinia
arechavaletae on Cardiospermum grandiflorum [29], Puccinia komarovii var. glanduliferae on
Impatiens glandulifera [30], and Prospodium transformans on Tecoma stans var. stans [31]. A
number of studies on Uromyces species have been carried out by a number of researchers
Figure 2. A general life cycle of rust genus Uromyces.
3.3. The Rusts as Classical Biocontrol Agents
Uromyces is the causal agent of rust disease on numerous agricultural, horticultural,
and forest plantations. Rust fungi of this genus are considered a major economic threat
due to possible yield losses from reduced production. Rust diseases on pea, beans, lentils,
polyhouse flower crops, clover, and many more are some of the hosts infected by this
genus. In addition to their identity as the causative agent of rust, the Uromyces species also
shows potential for the biological control of various phytopathogenic fungi, weeds, etc.

J. Fungi 2022,8, 633 6 of 99
The biology and effectiveness of Uromyces heliotropii were studied by Hasan and Aracil [
19
],
where they found the rust to effectively control an annual weed heliotrope (Heliotropium
europaeum L.).
They evaluated the biocontrol potential of the rust in a greenhouse and field inocula-
tion experiments and observed that the rust rapidly killed infected plants and reduced or
prevented seed production. Similarly, Anderson et al. [
20
] conducted a study to evaluate
the potential of three rusts naturally infecting Chilean needle grass (Nassella neesiana) in
Argentina, Uromyces pencanus,Puccinia graminella, and P. nassellae, as biocontrol agents.
They found U. pencanus to be most effective due to the damage it inflicts on its host in the
field. A list of species of Uromyces studied as biocontrol agents is given in Table 1.
Table 1. List of species of Uromyces with biocontrol potential.
Species of Uromyces Target Organism (Plants) Reference
Uromyces rumicis Rumex ckpus [21]
Uromyces galegae Galega officinalis [22]
Uromyces heliotropii Heliotropiu meuropaeum [19]
Uromyces pencanus Nassella neesiana [20,23–25]
Uromyces pisi-sativi Cytisus scoparius [26]
Uromyces pisi-sativi Chamaecytisus palmensis [26]
Uromyces pisi-sativi Lupinus polyphyllus [26]
Rust fungi as biocontrol agents are mainly studied in their use against plant weeds.
The biocontrol ability of some species of Puccinia was investigated, such as as P. abrupta
var. partheniicola on different growth stages of Parthenium hysterophorus [
27
,
28
], Puccinia
arechavaletae on Cardiospermum grandiflorum [
29
], Puccinia komarovii var. glanduliferae on
Impatiens glandulifera [
30
], and Prospodium transformans on Tecoma stans var. stans [
31
]. A
number of studies on Uromyces species have been carried out by a number of researchers
(see Table 1), but the assessment of species from other rust genera has not yet been well
established. Furthermore, the broad aspects of the rust fungi as biocontrol agents based on
biochemical and molecular approaches still need to be explored.
4. Data Collection and Molecular Analysis
4.1. Data Collection and Compilation
This paper was compiled based on the information obtained from an extensive search
of peer-reviewed publications, field guides, monographs, books, conference proceedings,
project reports, and other offline and online resources. This information was updated
as recently as December 2020. The information obtained was compiled firstly as a table
depicting names of species of Uromyces, their hosts along with the family, the locality of
occurrence, and the reference of scientific publication. The scientific names of the hosts and
fungi were then cross-verified for scientific entities. The host name given in the original
citation was sometimes changed to be consistent with the current taxonomy based on
The Plant List (http://www.theplantlist.org; accessed on 20 April 2022). The names of
rust fungi genus/species as reported in the cited publications were replaced by currently
accepted names according to the websites MycoBank (www.mycobank.org; accessed on
20 April 2022) and Species Fungorum (www.speciesfungorum.org; accessed on 20 April
2022). Fungal Databases, US National Fungus Collections, ARS, USDA, an important online
source of plant pathogens and their hosts, was also noted during the compilation [
32
]. An
attempt was made to summarize all available literature on diversity and distribution of
Uromyces spp.; only the most appropriate references were included in this study.
4.2. Analyses of Collected Data
After inserting the collected data into the primary database as a table, they were
analyzed for numerical and graphical summaries. First, the information was analyzed
by providing a comparative representation of the diversity and distribution of rust fungi

J. Fungi 2022,8, 633 7 of 99
(Uromyces) by country and continent. Thereafter, distribution patterns based on substrate
types (herb, shrub, and tree) were constructed to understand the host preference of these
rust fungi. In addition, the data of the host family were also presented. The publication
indices of Uromyces spp. in terms of year, decade, and era are analyzed and presented in
this paper. In addition, the references in other languages are translated into English so that
the scientific community can understand them easily.
4.3. Molecular Data Analyzing
DNA sequence data of Uromyces species from the LSU and ITS rDNA were down-
loaded from GenBank and through earlier published literature. A checklist of molecular
studies on Uromyces sp. along with the name of isolate and references were also prepared
and presented in Table 2. The relevant publications on molecular analyses were also con-
sulted [
5
,
12
,
33
,
34
]. Individual nucleotide sequences of LSU and ITS were aligned distinctly
using MAFFT 7 (http://mafft.cbrc.jp/alignment/server/; accessed on 1 April 2022) [
35
],
followed by manual checking and editing where necessary in BioEdit v. 7.0.9 [
36
]. The
sequences of taxa containing poorly aligned portions, incomplete data, missing sequence
data, and gaps were trimmed. The ITS and LSU sequences alignment was converted to
NEXUS format (.nxs) using ClustalX 2.1 (http://www.clustal.org/clustal2/; accessed on 1
April 2022) for Phylogenetic Analysis Using PAUP (PAUP) analysis. The aligned LSU and
ITS single-gene datasets and a concatenated dataset of LSU and ITS genes were analyzed
with PAUP 4.0b10 [
37
]. These datasets were run after completing the program output
tree in the Bootstrap.tre file. Maximum Likelihood bootstrap values greater than 60% are
considered good bootstrap supports and are given above each node. Phylogenetic trees are
visualized using the FigTree v1.4.0 program [
38
] and reorganized in Microsoft power point.
Table 2.
GenBank and voucher/culture collection accession numbers of Uromyces species included in
the phylogenetic study.
Taxa Isolate Name GenBank Accession No. References
ITS LSU
Uromyces acetosae DAOM 159824 HQ317557 HQ317557 [39]
Uromyces acuminatus DUCC506 - KC570451 [40]
CT-V080623-3 GU109282 - [41]
Uromyces aemulus HSZ0345 AF511081 AF511081 [42]
Uromyces aloes 2020-6-28-0005 MT136509 - Unpublished
WM 3290 - DQ917740 [43]
Uromyces alopecuri DAOM 234714 HM131363 HM131363 [44]
Uromyces alyxiae BPI 871816 - KY764186 Unpublished
Uromyces anthyllidis EB19 FJ468411 - [45]
Uromyces ari-triphylli BPI 871111 DQ354529 DQ354528 [12]
Uromyces betae BPI910289 KY764187 - Unpublished
Uromyces beticola PDD:101534 KX985757 KX985757 [46]
Uromyces bidenticola ZP-R1366 - MK518671 Unpublished
Uromyces
caricis-sempervirentis GZU 10-94 - DQ917714 Unpublished
Uromyces chenopodii-fruticosi K(M)107793 - MT053261 [47]
Uromyces ciceris-arietini BPI 879192 FJ468398 GU058030 [48]
- GQ914998 - [49]
Uromyces clignyi DAOM 192217 HM131364 - [44]

J. Fungi 2022,8, 633 8 of 99
Table 2. Cont.
Taxa Isolate Name GenBank Accession No. References
ITS LSU
Uromyces coloradensis DAOM 181557 HQ317565 - [39]
DAOM187619 HQ317564 - [39]
Uromyces coronatus DAOM 32991 HM131365 - [44]
BPI 910293 - KY764191 Unpublished
Uromyces dactylidis PRC692 KM667950 - [50]
TUB 14997 - DQ917745 [43]
Uromyces dianthi BPI 141961 - KY764192 Unpublished
Uromyces dolicholi DAOM 116149 HQ317563 - [39]
Uromyces durus IBAR11149 LC203758 - [51]
Uromyces eragrostidis BPI004474 - KY575113 Unpublished
Uromyces erythronii DE-soo-43285 MH205917 MH205917 [52]
Uromyces euphorbiae F425 AF180158 - [53]
BPI 863673 - KT750330 Unpublished
Uromyces ficariae WM 1398 - AF426204 [33]
Uromyces gageae HMJAU:8558 MG742207 MG742207 [54]
U242 DQ250133 DQ250133 [55]
Uromyces geranii BRIP 60100 - KX999898 [56]
Uromyces gladioli - - GU936633 [57]
Uromyces graminicola URO 3 KR264705 - [58]
Uromyces halstedii U1673 MG905913 - [59]
Uromyces hawksworthii
UB:22382 KR821137 -
[60]
UB:22875 KR821138 -
UB:22875 - NG059990
Uromyces hedysari-obscuri DAOM 189699 HQ317573 - [39]
DAOM 223015 - HQ317572
Uromyces ixiae RSA191 - DQ917738 [43]
RSA8 - DQ917739 Unpublished
Uromyces japonicus TNS:F:61987 LC203759 LC203759 [51]
Uromyces jonesii DAOM 137431 HQ317559 HQ317559 Unpublished
Uromyces junci GZU 11-98 - AF426203 [33]
Uromyces klotzschianus
BA91 MF044015 -
[61]
BA92 MF044016 -
BA93 - MF044017
Uromyces
lespedezae-procumbentis BPI 910294 KY764193 - Unpublished
Uromyces lomandracearum BRIP:59022 - KM249862 [62]
Uromyces lupini DAOM 233961 HQ317570 HQ317570 [39]
Uromyces magnusii PDD:94487 KX985755 - [46]
Uromyces minor MVAP50000151 MK045314 - Unpublished
Uromyces musae DAOM 192221 HQ317556 - [39]

J. Fungi 2022,8, 633 9 of 99
Table 2. Cont.
Taxa Isolate Name GenBank Accession No. References
ITS LSU
Uromyces muscari DAOM 75626 HQ317552 HQ317552 [39]
Uromyces neotropicalis MCA2533 - DQ021885 [63]
Uromyces novissimus R20 - EU851147 [57]
Uromyces orientalis BRIP 60934 KX999899 - [56]
Uromyces otaviensis NA 305 - DQ917715 [43]
Uromyces peglerae BPI 843311 - KY575116 Unpublished
Uromyces pisi-sativa - DQ521591 - Unpublished
BRIP 60151 - KX999900 [56]
Uromyces plumbarius 070P06138145 KP313731 - [64]
RK44 - JQ312671 [65]
Uromyces polygoni-avicularis DAOM 181565 - HQ317558 [39]
Uromyces probus BPI 893209 KT267172 - [66]
Uromyces rebeccae FDWSRU 15-022 MN386999 MN386999 [47]
Uromyces reichertii 6421 AF511084 AF511084 [42]
Uromyces rumicis PDD:93529 KX985763 - [46]
BPI 910298 - KY764197 Unpublished
Uromyces salsolae FDWSRU 05-090 MN381162 MN381162 [47]
Uromyces scaevolae BRIP:57517 KF690686 KF690686 [67]
Uromyces scutellatus E328 AF180200 AF180200 [53]
Uromyces setariae-italicae BPI 863744 KY575068 - Unpublished
Uromyces setariae-italicae BPI 863750 - KY575069 Unpublished
Uromyces socius BPI 910302 - KY764200 Unpublished
Uromyces sporobolicola DAOM 114294 HQ317560 HQ317560 [39]
Uromyces striatus U-675 HQ412651 - [68]
DAOM 240966 HQ317512 - [39]
Uromyces striolatus E405 AF180201 AF180201 [53]
Uromyces strobilanthis BPI 893253 - KY798375 Unpublished
Uromyces tenuicutis BRIP 60012 - KX999904 [56]
Uromyces transversalis
NZ11 PDD-99990 JQ777025 - Unpublished
CA1124 JQ776984 - Unpublished
PUR:N16743 - MF374730 [69]
Uromyces trifolii - - GU936634 [57]
Uromyces trifolii-repentis R217 EU014070 - [70]
BRIP 57653 - KX999905 [56]
Uromyces viciae-fabae TSH_R6320 AB115665 - [71]
WM 1365 - AF426199 [33]
Uromyces vignae
H92019 AB115731 - [72]
H50721 AB115726 -
BRIP 60213 - KX999906 [56]
BRIP 60213 - KX999906
Uromyces wedeliae BRIP 60000 - KX999907 [56]

J. Fungi 2022,8, 633 10 of 99
5. Distribution, Diversity, and Molecular Data Analysis of Uromyces
5.1. Phylogenetic Analyses
In the phylogenetic results, Uromyces were separated into two complexes in both ITS
and LSU sequence data. Both complexes of ITS and LSU share many similar sequences.
The incomplete sequences were mostly found in the Uromyces sequence dataset, e.g., ITS1
and 5.8S or ITS1, 5.8S complete, and ITS partial or 28S partially. Approximately 50% of the
sequences had up to 300 nucleotides, while the remaining sequences had up to 800 nucleotides.
Incomplete sequences can result in two complexes in a single genus. Therefore, complete gene
sequences from ITS and LSU are needed to analyze these complex clades (Figures 3and 4).
J. Fungi 2022, 8, x FOR PEER REVIEW 11 of 88
Figure 3. Phylogram generated from PAUP analysis of Uromyces species of ITS sequences. The scale
bar indicates 70 changes and outgroup Gymnosporangium przewalskii BJFCR01859_NR154073.
Bootstrap values of MP > 60% are given above branches, and 72 sequences are included in the
phylogenetic analyses. The best maximum parsimony (MP) dataset consists of 394 total characters,
of which 164 were constant, 156 parsimony-informative, and 74 parsimony-non-informative. The
parsimony analysis of the data matrix showed a thousand equally parsimonious trees with a length
of 917 steps in the first tree (CI = 0.408, RI = 0.702, RC = 0.286, HI = 0.592).
Figure 3.
Phylogram generated from PAUP analysis of Uromyces species of ITS sequences. The
scale bar indicates 70 changes and outgroup Gymnosporangium przewalskii BJFCR01859_NR154073.
Bootstrap values of MP > 60% are given above branches, and 72 sequences are included in the
phylogenetic analyses. The best maximum parsimony (MP) dataset consists of 394 total characters,
of which 164 were constant, 156 parsimony-informative, and 74 parsimony-non-informative. The
parsimony analysis of the data matrix showed a thousand equally parsimonious trees with a length
of 917 steps in the first tree (CI = 0.408, RI = 0.702, RC = 0.286, HI = 0.592).

J. Fungi 2022,8, 633 11 of 99
J. Fungi 2022, 8, x FOR PEER REVIEW 12 of 88
Figure 4. Phylogram generated from PAUP analysis of Uromyces species of LSU sequences. The scale
bars indicate 60 changes and outgroup Gymnosporangium przewalskii BJFCR01859_NG060667.
Bootstrap values of MP equal to or greater than 60% are given above branches, and 58 sequences
are included in the phylogenetic analyses. The best maximum parsimony (MP) dataset consists of
873 total characters, of which 600 were constant, 125 parsimony-informative, and 148 parsimony-
non-informative. The parsimony analysis of the data matrix showed 1000 equally parsimonious
trees with a length of 625 steps in the first tree (CI = 0.550, RI = 0.708, RC = 0.389, HI = 0.450).
Figure 4.
Phylogram generated from PAUP analysis of Uromyces species of LSU sequences. The
scale bars indicate 60 changes and outgroup Gymnosporangium przewalskii BJFCR01859_NG060667.
Bootstrap values of MP equal to or greater than 60% are given above branches, and 58 sequences
are included in the phylogenetic analyses. The best maximum parsimony (MP) dataset consists of
873 total characters,
of which 600 were constant, 125 parsimony-informative, and 148 parsimony-
non-informative. The parsimony analysis of the data matrix showed 1000 equally parsimonious trees
with a length of 625 steps in the first tree (CI = 0.550, RI = 0.708, RC = 0.389, HI = 0.450).
5.2. Morphological Diversity and Distribution
The results compiled on diversity and distribution revealed that the rust genus Uromyces
comprised a total of 1500 species that occurred worldwide as obligate parasitic fungi on
vascular plants [
3
,
4
]. After combing through the different online
databases [1,32,73],
a total of
988 species were included in this paper. Similar to all fungi, its distribution shows great
variations in different parts of the world. The tremendously changing climates across the
world lead to diversified flora, resulting in a wide diversity and distribution of rust fungi.
With regard to global diversity, it is pertinent to note here that Uromyces varies in diversity

J. Fungi 2022,8, 633 12 of 99
among countries and continents. These rust fungi generally show a macrocyclic nature and
autoecious mode of the life cycle [
2
,
5
], which confirms their morphological diversity on
specific hosts in particular regions. The broad host range in microcyclic and heteroecious
life cycles was also found in Uromyces species.
The genus Uromyces predominantly showed its great diversity in North America in
comparison to other continents. Almost 834 (30%) species are described here, which is
the highest among all continents. The diversity of the genus is known in other continents
of the world as follows: Asia, with 633 (23%) species described; Europe, with 622 (23%)
species; North America, with 321 (12%) species; Africa, with 313 (11%) described species;
and Australia, with 32 (1%) species described. The genus appears to be well-represented
in North American, Asian, and European counties. After dispersal by various modes,
such as wind, water, or insect vectors, the propagules of rust fungi germinate and infect
plant tissues of specific hosts. Entry of these pathogens takes place either by natural
openings such as lenticels and stomata or by wounds or injuries caused by various physical
agents [74].
Human anthropogenic activities also play an important role in the global distribution
of these organisms. So far, Uromyces species have been found on every land on earth
except Antarctica. To understand the distribution of the Uromyces species, we analyzed
their distributions across continents and terrestrial ecoregions. More than 150 countries
and territories or occupancies showed the distribution of this rust genus. Although only
73 sequences
of ITS and nLSU Uromyces species were identified based on molecular charac-
teristics, respectively, the majority of the species are still identified morphologically. This
may impact the number and distribution pattern of the Uromyces species as molecular-
based research on the rust fungi progress. In comparison to all continents, the highest
distribution of 834 species of Uromyces was recorded over 66 regions of different countries
and dependencies of North America. The distribution pattern observed in other continents
was observed as follows: 633 species in 26 countries and dependencies of Asia; 622 species
in
33 countries
and dependencies of Europe; 321 species in 17 countries and dependencies
of South America; 313 species in 27 countries and dependencies of Africa; and 37 species in
3 countries, islands, or dependencies of Oceania. The array of this global distribution of
Uromyces species reveals their vast diversity and justifies its position as the second-largest
genus of rust fungi (Figures 5and 6).
J. Fungi 2022, 8, x FOR PEER REVIEW 13 of 88
Human anthropogenic activities also play an important role in the global distribution
of these organisms. So far, Uromyces species have been found on every land on earth
except Antarctica. To understand the distribution of the Uromyces species, we analyzed
their distributions across continents and terrestrial ecoregions. More than 150 countries
and territories or occupancies showed the distribution of this rust genus. Although only
73 sequences of ITS and nLSU Uromyces species were identified based on molecular
characteristics, respectively, the majority of the species are still identified
morphologically. This may impact the number and distribution pattern of the Uromyces
species as molecular-based research on the rust fungi progress. In comparison to all
continents, the highest distribution of 834 species of Uromyces was recorded over 66
regions of different countries and dependencies of North America. The distribution
pattern observed in other continents was observed as follows: 633 species in 26 countries
and dependencies of Asia; 622 species in 33 countries and dependencies of Europe; 321
species in 17 countries and dependencies of South America; 313 species in 27 countries
and dependencies of Africa; and 37 species in 3 countries, islands, or dependencies of
Oceania. The array of this global distribution of Uromyces species reveals their vast
diversity and justifies its position as the second-largest genus of rust fungi (Figures 5 and
6).
Figure 5. Heat map to show the global distribution and species richness of Uromyces spp.
Distribution: America, Kazakhstan, India, Brazil, Syria, México, Russia, Cyprus,
Ecuador, Uzbekistan, New Guinea, Cape Province, Algeria, Tunisia, Japan, Norway,
China, Uganda, Hawaii, Argentina, Canada, Siberia, France, South Africa, Sri Lanka,
Ethiopia, Colombia, Romania, Costa Rica, Israel, Chile, Switzerland, Ivory Coast, Uganda,
Turkey, Kenya, Morocco, Britain, New Zealand, Malaysia, Philippines, Iran, Greece,
Madagascar, Romania, Australia, Yunnan, Tadzhikistan, Pakistan, Zimbabwe, Myanmar,
Namibia, Costa Rica, Taiwan, Nepal, Italy, Cuba, Afghanistan, Egypt, Iraq, Peru, Tibet,
Croatia, Sudan, Ecuador, Jamaica, Sweden, Bolivia, Spain, Austria, Tunisia, Kenya,
Norway, Kansas, Peru, Poland, and Venezuela.
Figure 5. Heat map to show the global distribution and species richness of Uromyces spp.

J. Fungi 2022,8, 633 13 of 99
J. Fungi 2022, 8, x FOR PEER REVIEW 14 of 88
Figure 6. Heat maps to show the continental distribution and species richness of Uromyces spp.
5.3. Distribution Patterns of Rust Fungi (Uromyces) by Substrate Types
Uromyces, being the second-largest rust genus after Puccinia, contains a number of
important plant pathogens. The species of Uromyces attacks nearly all categories of plants
and causes great damage to both plant and their products. Uromyces is a genus of rust
fungi that infects both monocots and dicots throughout the world. Analyses of the
available literature on host diversity of Uromyces revealed that a total of 647 plant genera
belonging to 95 plant families were found to be infected by these rust fungi. Uromyces is
particularly reported in plant families such as Amaranthaceae, Apiaceae, Asparagaceae,
Asteraceae, Caryophyllaceae, Cucurbitaceae, Cyperaceae, Euphorbiaceae, Fabaceae, Iridaceae,
Poaceae, and Rubiaceae. The highest occurrences of Uromyces spp. were observed in Poaceae
and Fabaceae, with 105 and 103 infected host genera, respectively. A number of other
families that have an infection in up to 50 plant genera have also been observed, including
Acanthaceae, Asparagaceae, Asteraceae, Caprifoliaceae, Cyperaceae, Euphorbiaceae, Fabaceae,
Lamiaceae, Liliaceae, Loranthaceae, Oleaceae, Poaceae, and Polygonaceae. A study on the
diversity and distribution of Uromyces in India by Gautam and Avasthi [75] revealed that
nearly 180 plant species belonging to 85 genera and 32 families were found to be infected
with Uromyces spp. Among all families, Fabaceae and Poaceae were found to be the most
infected with different species of Uromyces. A family-wise comparison of the genera of
infected host plants is shown in Figure 7.
Figure 6. Heat maps to show the continental distribution and species richness of Uromyces spp.
Distribution: America, Kazakhstan, India, Brazil, Syria, México, Russia, Cyprus,
Ecuador, Uzbekistan, New Guinea, Cape Province, Algeria, Tunisia, Japan, Norway, China,
Uganda, Hawaii, Argentina, Canada, Siberia, France, South Africa, Sri Lanka, Ethiopia,
Colombia, Romania, Costa Rica, Israel, Chile, Switzerland, Ivory Coast, Uganda, Turkey,
Kenya, Morocco, Britain, New Zealand, Malaysia, Philippines, Iran, Greece, Madagascar,
Romania, Australia, Yunnan, Tadzhikistan, Pakistan, Zimbabwe, Myanmar, Namibia, Costa
Rica, Taiwan, Nepal, Italy, Cuba, Afghanistan, Egypt, Iraq, Peru, Tibet, Croatia, Sudan,
Ecuador, Jamaica, Sweden, Bolivia, Spain, Austria, Tunisia, Kenya, Norway, Kansas, Peru,
Poland, and Venezuela.
5.3. Distribution Patterns of Rust Fungi (Uromyces) by Substrate Types
Uromyces, being the second-largest rust genus after Puccinia, contains a number of im-
portant plant pathogens. The species of Uromyces attacks nearly all categories of plants and
causes great damage to both plant and their products. Uromyces is a genus of rust fungi that
infects both monocots and dicots throughout the world. Analyses of the available literature
on host diversity of Uromyces revealed that a total of 647 plant genera belonging to 95 plant
families were found to be infected by these rust fungi. Uromyces is particularly reported
in plant families such as Amaranthaceae,Apiaceae,Asparagaceae,Asteraceae,Caryophyllaceae,
Cucurbitaceae,Cyperaceae,Euphorbiaceae,Fabaceae,Iridaceae,Poaceae, and Rubiaceae. The
highest occurrences of Uromyces spp. were observed in Poaceae and Fabaceae, with 105 and
103 infected host genera, respectively. A number of other families that have an infection
in up to 50 plant genera have also been observed, including Acanthaceae,Asparagaceae,
Asteraceae,Caprifoliaceae,Cyperaceae,Euphorbiaceae,Fabaceae,Lamiaceae,Liliaceae,Loranthaceae,
Oleaceae,Poaceae, and Polygonaceae. A study on the diversity and distribution of Uromyces
in India by Gautam and Avasthi [
75
] revealed that nearly 180 plant species belonging to

J. Fungi 2022,8, 633 14 of 99
85 genera
and 32 families were found to be infected with Uromyces spp. Among all families,
Fabaceae and Poaceae were found to be the most infected with different species of Uromyces.
A family-wise comparison of the genera of infected host plants is shown in Figure 7.
J. Fungi 2022, 8, x FOR PEER REVIEW 15 of 88
Figure 7. Family-wise comparison of genera of infected host plants.
5.4. Global Host Range of Uromyces Species
The host range of Uromyces species showed their occurrence in 647 plant genera of 95
plant families. They are reported to be infected by various species of Uromyces, such as:
Abrus, Abuilon, Acacia, Acalypha, Acantholimon, Acanthophyllum, Acetosa, Achyranthes,
Acidanthera, Acorus, Aconitum, Actinostemon, Adenostyles, Aegopogon, Aellenia, Aeluropus,
Agropyron, Agrostis, Aira, Albizia, Albuca, Alhagi, Allium, Aloe, Alopecurus, Alstroemeria,
Alysicarpus, Alyxia, Amblytropis, Ambrosia, Amischotolype, Amphicarpaea, Amphilophis,
Anabasis, Anaphalis, Anarthrophyllum, Ancrumia, Andropogon, Aneilema, Anemone, Anguria,
Anguria, Anisotome, Anomatheca, Anotis, Anthacanthus, Anthericum, Antholyza, Aphelandra,
Apios, Apluda, Ardisia, Arenaria, Argyrolobium, Arisaema, Aristida, Armeria, Arnica, Artemisia,
Arthrocnemum, Asclepias, Ascyrum, Aspalathus, Asperula, Aspilia, Aster, Astragalus, Astrebla,
Atriplex, Atylosia, Avenella, Avenula, Babiana, Baccharis, Bahia, Baltimora, Bartholina, Basella,
Bauhinia, Beckeropsis, Beckmannia, Bellevalia, Beloperone, Benedictella, Beta, Bidens, Blainvillea,
Boissiera, Bolboschoenus, Boltonia, Bomarea, Bonaveria, Bongardia, Borreria, Bothriochloa,
Bouvardia, Brachiaria, Bradburya, Briza, Brodiaea, Bromus, Bryzopyrum, Bufonia, Buforrestia,
Bulbine, Bupleurum, Buxus, Cacalia, Caccinia, Cachrys, Cajanus, Calamagrostis, Calandrinia,
Callicarpa, Callisia, Calopogonium, Caltha, Calycotome, Calyptridium, Camptosema,
Campylotropis, Canavalia, Capillipedium, Caragana, Carex, Cassia, Caucalis, Cayaponia, Celosia,
Celtis, Centropogon, Centrosema, Cerastium, Ceratocarpus, Ceratoides, Cestrum, Chaetobromus,
Chaetochloa, Chaetolimon, Chamaecrista, Chamaecytisus, Chamaesyce, Chasmanthe,
Chenopodium, Chesneya, Chiloglottis, Chionodoxa, Chloris, Chlorophytum, Chorizanthe, Cicer,
Figure 7. Family-wise comparison of genera of infected host plants.
5.4. Global Host Range of Uromyces Species
The host range of Uromyces species showed their occurrence in 647 plant genera of
95 plant
families. They are reported to be infected by various species of Uromyces, such as:
Abrus,Abuilon,Acacia,Acalypha,Acantholimon,Acanthophyllum,Acetosa,Achyranthes,Acidan-
thera,Acorus,Aconitum,Actinostemon,Adenostyles,Aegopogon,Aellenia,Aeluropus,Agropyron,
Agrostis,Aira,Albizia,Albuca,Alhagi,Allium,Aloe,Alopecurus,Alstroemeria,Alysicarpus,
Alyxia,Amblytropis,Ambrosia,Amischotolype,Amphicarpaea,Amphilophis,Anabasis,Anaphalis,
Anarthrophyllum,Ancrumia,Andropogon,Aneilema,Anemone,Anguria,Anguria,Anisotome,
Anomatheca,Anotis,Anthacanthus,Anthericum,Antholyza,Aphelandra,Apios,Apluda,Ardisia,
Arenaria,Argyrolobium,Arisaema,Aristida,Armeria,Arnica,Artemisia,Arthrocnemum,As-
clepias,Ascyrum,Aspalathus,Asperula,Aspilia,Aster,Astragalus,Astrebla,Atriplex,Atylosia,
Avenella,Avenula,Babiana,Baccharis,Bahia,Baltimora,Bartholina,Basella,Bauhinia,Beckeropsis,
Beckmannia,Bellevalia,Beloperone,Benedictella,Beta,Bidens,Blainvillea,Boissiera,Bolboschoenus,
Boltonia,Bomarea,Bonaveria,Bongardia,Borreria,Bothriochloa,Bouvardia,Brachiaria,Bradburya,
Briza,Brodiaea,Bromus,Bryzopyrum,Bufonia,Buforrestia,Bulbine,Bupleurum,Buxus,Cacalia,

J. Fungi 2022,8, 633 15 of 99
Caccinia,Cachrys,Cajanus,Calamagrostis,Calandrinia,Callicarpa,Callisia,Calopogonium,Caltha,
Calycotome,Calyptridium,Camptosema,Campylotropis,Canavalia,Capillipedium,Caragana,
Carex,Cassia,Caucalis,Cayaponia,Celosia,Celtis,Centropogon,Centrosema,Cerastium,Cerato-
carpus,Ceratoides,Cestrum,Chaetobromus,Chaetochloa,Chaetolimon,Chamaecrista,Chamae-
cytisus,Chamaesyce,Chasmanthe,Chenopodium,Chesneya,Chiloglottis,Chionodoxa,Chloris,
Chlorophytum,Chorizanthe,Cicer,Cicuta,Cimacoptera,Cirsium,Cissus,Cladostachys,Cladrastis,
Claytonia,Cleome,Climacoptera,Clitoria,Clutia,Cnidoscolus,Cnidoscolus,Colchicum,Collomia,
Cologania,Colutea,Combretum,Commelina,Commiphora,Convolvulus,Corallocarpus,Cordia,
Coronilla,Cosmos,Crepis,Crocosmia,Crocus,Crotalaria,Cruckshanksia,Cucubalus,Cucumis,
Cucurbita,Curculigo,Cyanotis,Cyathula,Cymbopogon,Cynosurus,Cyperus,Cypholepis,Cytisus,
Dactylis,Dactyloctenium,Dalbergia,Danthonia,Deeringia,Dendroseris,Deschampsia,Desmod-
ium,Desmostachya,Dianthus,Dichanthium,Dichromena,Dicliptera,Dierama,Digitaria,Diodia,
Dipcadi,Dipogon,Discaria,Distichlis,Dodecatheon,Dolicholus,Dolichos,Doronicum,Doryc-
nium,Dorycnopsis,Dorystaechas,Doyerea,Dracaena,Drimiopsis,Echinocephalum,Edwardsia,
Ehretia,Ehrharta,Eleocharis,Emex,Emmeorhiza,Enargea,Endymion,Engysiphon,Epicampes,
Eragrostis,Eremopogon,Eremopyrum,Eriochloa,Eriogonum,Eriophyllum,Eriosema,Eriosper-
mum,Erodium,Ervum,Erythrina,Erythronium,Eulophia,Euphorbia,Euryops,Excoecaria,
Exotheca,Fatoua,Ferraria,Ferula,Festuca,Ficaria,Fimbristylis,Flemmingia,Fleurya,Floscopa,
Freesia,Fritillaria,Gagea,Galactia,Galega,Galinsoga,Galium,Galphimia,Gamanthus,Gau-
dichaudia,Gaura,Gayophytum,Geissorhiza,Genista,Gentiana,Geranium,Gilliesia,Gladiolus,
Glaux,Glyceria,Glycyrrhiza,Gnaphalium,Gomphrena,Gossypium,Gouania,Grindelia,Guizotia,
Gurania,Gypsophila,Habrochloa,Haemanthus,Halenia,Halimodendron,Halocharis,Haloxylon,
Hardenbergia,Hedysarum,Helianthus,Helichrysum,Helictotrichon,Heliopsis,Heliotropium,
Helleborus,Helmontia,Hemarthria,Heracleum,Hesperantha,Heteranthera,Heteromorpha,Het-
eropogon,Hewittia,Hibiscus,Hippocrepis,Hippomarathrum,Holcus,Homogyne,Honckenya,
Hookera,Hordeum,Hosackia,Houstonia,Hyacinthoides,Hyacinthus,Hydrocotyle,Hymenocarpos,
Hyparrhenia,Hypericum,Hypoestes,Hypoxis,Hypsela,Indigofera,Inga,Ipomoea,Iresine,Isathne,
Ixia,Jacobinia,Jacquemontia,Jasminum,Jatropha,Jaumea,Juncus,Juniperus,Justica,Kalid-
ium,Karroochloa,Kochia,Koeleria,Krameria,Krascheninnikovia,Kummerowia,Kyllinga,Lablab,
Laburnum,Lachenalia,Lantana,Lapeirousia,Lasiacis,Lasiocorys,Lathyrus,Ledenbergia,Leersia,
Lembotropis,Lens,Leonotis,Leontice,Leopoldia,Leptochloa,Lerchenfeldia,Lespedeza,Leucocoryne,
Ligularia,Lilium,Limonium,Lloydia,Loeselia,Lomandra,Loranthus,Lotononis,Lotus,Lupinus,
Luzula,Lychnis,Lygeum,Lysimachia,Maackia,Maianthemum,Mallotus,Manettia,Manihot,
Massonia,Mattiastrum,Medicago,Meibomia,Melandrium,Melanolepis,Melanthera,Melas-
phaerula,Medica,Melilotus,Melothria,Menyanthes,Mercurialis,Merremia,Messerschmidia,
Microchloa,Microlespedeza,Microtis,Miersia,Mikania,Milium,Milla,Mimusops,Minuartia,
Modecca,Moehringia,Momordica,Monocymbium,Montanoa,Moraea,Mucuna,Muehlenbeckia,
Muhlenbergia,Mulinum,Muricauda,Musa,Muscari,Myosotis,Myrsine,Nassauvia,Nassella,
Noaea,Nothoscordum,Nymphoides,Ocimum,Odontonema,Oenanthe,Oenothera,Olsynium,
Onagra,Onobrychis,Ononis,Ophiorrhiza,Oreopolus,Ormosia,Ornithogalum,Ornithopus,
Orobus,Orthosiphon,Oryctanthus,Oryza,Oxytropis,Pachylophus,Panicum,Parietaria,Pas-
palum,Paspalum,Passiflora,Pavonia,Peireskia,Peltandra,Pennisetum,Pentace,Pentaschis-
tis,Peplis,Peracarpa,Perymenium,Petrorhagia,Petrosimonia,Peucedanum,Phaca,Phacelurus,
Phalaris,Phaseolus,Phleum,Phlogacanthus,Phlox,Phoradendron,Phragmites,Phrygilanthus,
Phthirusa,Physanthyllis,Phyteuma,Piptanthus,Pisum,Pittosporum,Plantago,Poa,Poinsettia,
Poiretia,Poitea,Polemannia,Polemonium,Pollia,Polycnemum,Polygala,Polygonum,Polymnia,
Polytrias,Polyxena,Pontederia,Pozoa,Prangos,Pratia,Primula,Priva,Pseudarthria,Pseuderan-
themum,Psoralea,Psychotria,Pteracanthus,Pulicaria,Quinchamalium,Ranunculus,Rapanea,
Rapanea,Ratibida,Rhinacanthus,Rhinopetalum,Rhynchospora,Rhynchosia,Ribes,Roepera,Ro-
mulea,Rosa,Rottboellia,Rudbeckia,Ruellia,Rumex,Rytidosperma,Sabinea,Salicornia,Salmea,
Salpichroa Salsola,Sapium,Saururus,Scaevola,Schismus,Schizachyrium,Schizeilema,Scilla,
Scirpus,Scleranthus,Scleria,Sclerochloa,Scribneria,Secale,Secamone,Securigera,Sedum,Selliera,
Selysia,Senecio,Sesbania,Seseli,Sessea,Setaria,Sigesbeckia,Silene,Silphium,Simethis,Sipho-
campylus,Siphocampylus,Sisyrinchium,Sium,Smilacina,Smilax,Snowdenia,Solanum,Solaria,

J. Fungi 2022,8, 633 16 of 99
Solidago,Sophora,Sorghum,Sparaxis,Sparganium,Spartina,Spergularia,Spermacoce,Sphaeral-
cea,Sphenostylis,Sporobolus,Spraguea,Stellaria,Stenorrhynchus,Stipa,Strobilanthes,Strumaria,
Struthanthus,Stylochaeton,Suaeda,Swainsona,Teramnus,Tessaria,Tetraena,Tetragonolobus,
Thalictrum,Thapsia,Thelymitra,Themeda,Thermopsis,Thyrsacanthus,Tithymalus,Tournefor-
tia,Toona,Tradescantia,Tragus,Triadenum,Trianosperma,Tribolium,Trichocline,Tricholaena,
Trichoneura,Trichosanthes,Tricoryne,Trifolium,Trigonella,Tripogandra,Tripogon,Tripsacum,
Trisetum,Triteleia,Tritonia,Trochomeriopsis Tulipa,Tylosema,Ulex,Uraria,Urginea,Urochloa,
Valerianella,Veratrum,Verbascum,Veronica,Vicia,Vigna,Vilfa,Viola,Viscaria,Viscum,Vitis,
Vossia,Vulpia,Watsonia,Wedelia,Wilbrandia,Wulffia,Zantedeschia,Zebrina,Zexmenia,Zigade-
nus,Zizania,Zizaniopsis, and Zygophyllum. This compiled information depicts the broad
host range of these rust fungi. The association of Uromyces species with some host plants is
given in Figure 8, while detailed information on the diversity, host range, and distribution
of Uromyces species is summarized in Table 3.
J. Fungi 2022, 8, x FOR PEER REVIEW 17 of 88
Thalictrum, Thapsia, Thelymitra, Themeda, Thermopsis, Thyrsacanthus, Tithymalus,
Tournefortia, Toona, Tradescantia, Tragus, Triadenum, Trianosperma, Tribolium, Trichocline,
Tricholaena, Trichoneura, Trichosanthes, Tricoryne, Trifolium, Trigonella, Tripogandra,
Tripogon, Tripsacum, Trisetum, Triteleia, Tritonia, Trochomeriopsis Tulipa, Tylosema, Ulex,
Uraria, Urginea, Urochloa, Valerianella, Veratrum, Verbascum, Veronica, Vicia, Vigna, Vilfa,
Viola, Viscaria, Viscum, Vitis, Vossia, Vulpia, Watsonia, Wedelia, Wilbrandia, Wulffia,
Zantedeschia, Zebrina, Zexmenia, Zigadenus, Zizania, Zizaniopsis, and Zygophyllum. This
compiled information depicts the broad host range of these rust fungi. The association of
Uromyces species with some host plants is given in Figure 8, while detailed information
on the diversity, host range, and distribution of Uromyces species is summarized in Table
3.
Figure 8. Occurrence of Uromyces species with some host plants. (A–G) Uromyces fabae on Pisum
sativum (pea). (A) Aecia; (B,C) Uredia; (D) Telia; (E) Aeciospores; (F) Urediospores; (G) Teliospores;
(H–J) Uromyces geranii on Geranium sp.; (H,I) Uredia; (J) Uredinospores; (K–N) Uromyces rumicis on
Rumex sp; (K,L) Uredia; (M) Urediospores; (N) Teliospores; (O–R) Uromycesviciae-fabae on Vicia faba;
(O–Q) Different rust sorus on natural host; (R) Aecia; (S) Urediospores and Teliospores; (T–W)
Uromyces trifolii on Trifolium sp.; (T–V) Uredia and Telia; (W) Teliospores. Scale bars: C,I,L,Q,R,U,V
= 1 mm; D = 0.5 mm; E,F,G,J,M,N,S,W = 20 µm.
Figure 8.
Occurrence of Uromyces species with some host plants. (
A
–
G
)Uromyces fabae on Pisum
sativum (pea). (
A
) Aecia; (
B
,
C
) Uredia; (
D
) Telia; (
E
) Aeciospores; (
F
) Urediospores; (
G
) Teliospores;
(
H
–
J
)Uromyces geranii on Geranium sp.; (
H
,
I
) Uredia; (
J
) Uredinospores; (
K
–
N
)Uromyces rumicis
on Rumex sp; (
K
,
L
) Uredia; (
M
) Urediospores; (
N
) Teliospores; (
O
–
R
)Uromycesviciae-fabae on Vicia
faba; (
O
–
Q
) Different rust sorus on natural host; (
R
) Aecia; (
S
) Urediospores and Teliospores; (
T
–
W
)
Uromyces trifolii on Trifolium sp.; (
T
–
V
) Uredia and Telia; (
W
) Teliospores. Scale bars:
C
,
I
,
L
,
Q
,
R
,
U
,
V
=
1 mm; D= 0.5 mm; E,F,G,J,M,N,S,W= 20 µm.

J. Fungi 2022,8, 633 17 of 99
Table 3. Diversity, host range, and distribution of species of Uromyces.
Species Host Locality Reference
Uromyces abbreviatus Arthur Psoralea purshii and P. physodes California, Idaho (Western USA), Nevada,
and Washington (USA) [76–78]
Uromyces acantholimonis Syd. & P. Syd. Acantholimon libanoticum Afghanistan, Iran, Israel, Kazakhstan,
Pakistan, and Turkey [79–81]
Uromyces acetosae J. Schröt. Acetosa pratensis,Rumex acetosa,R. acetosella,
R. acetoselloides,R. alpestris, and R. scutatus
Bulgaria, Czech Republic, China,
Czechoslovakia, Finland, Germany, Iceland,
México, Norway, Poland, Portugal,
Romania, Spain, Sweden, Russia, Taiwan,
Turkey, and the United Kingdom
[80,82–84]
Uromyces achrous Syd. & P. Syd. Dalbergia sissoo India, Pakistan [85,86]
Uromyces aconiti Fuckel
Aconitum apetalum,A. barbatum,A. carmichaelii,
A. delavayi,A. delavayi,A. lycoctonum,
A. macrorhynchum,A. monticola,A. umbrosum,
A. umbrosum, and A. volubile
China and Japan [87–91]
Uromyces aconiti-lycoctoni (DC.) G. Winter Aconitum apetalum California, Finland, Romania, and Spain [78,92]
Uromyces acori T.S. Ramakr. & Rangaswami Acorus calamus India, China, Japan, and Thailand [93–99]
Uromyces actinostemonis H.S. Jacks. & Holw. Actinostemon concolor Brazil [100–103]
Uromyces acuminatus Arthur
Arenaria lateriflora,Collomia linearis,
Dodecatheon pulchellum,Honckenya peploides,
Lysimachia ciliata,Maianthemum canadense,
Phlox glaberrima,P. pilosa,Polemonium reptans,
Smilacina stellate,Spartina alterniflora,S. alterniflora,
S. cynosuroides,S. gracilis,S. patens, and S. pectinata
Alberta, Bulgaria, California, Canada,
Colorado, Delaware, Florida, Indiana, Iowa,
Maine, Maryland, Massachusetts,
Michigan,
Montana, New Hampshire, New Jersey,
New York, North America, North Dakota,
Nova Scotia, Ontario, Quebec, Rhode
Island,
Saskatchewan, Wisconsin, and Wyoming
[41,104]
Uromyces acutatus Fuckel Gagea bohemica and G. villosa Germany [105]
Uromyces adelphicus Syd. Milium trichopodium Syria [106]
Uromyces aecidiiformis (F. Strauss) C.C. Rees Fritillaria pallidiflora,F. pineticola,
F. ussuriensis,Lilium bulbiferum, and L. candidum China, Greece, Germany, Norway, Sweden,
and the United Kingdom [91,105,107–109]
Uromyces aegopogonis Dietel & Holw. Aegopogon cenchroides,A. geminiflorus,
A. gracilis, and A. tenellus México [106,110]

J. Fungi 2022,8, 633 18 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces aeluropodinus Tranzschel Aeluropus littoralis Ukraine [111]
Uromyces aeluropodis-repentis Nattrass Aeluropus repens Cyprus, Turkey, and Ukraine [80,106]
Uromyces aemulus Arthur Allium acuminatum,A. brevistylum,
A. brandegei,A. douglasii,A. gooddingii,
A. tolmiei, and A. validum
Arizona, California, Colorado, Canada,
Idaho, Nevada, Utah, Oregon, Washington,
and Wyoming [112–116]
Uromyces affinis G. Winter
Alstroemeria aurantiaca,A. caryophyllaea,
A. inodora,A. isabellana,A. nemorosa,
Curculigo scorzoneraefolia,Hypoxis decumbens,
H. erecta, and H. hirsuta
Brazil, Colombia, Dominica, Dominican
Republic, Florida, Missouri, Mississippi,
and
Puerto Rico
[102,117–125]
Uromyces agnatus Arthur Jatropha stimulosa Florida [112]
Uromyces agropyri Barclay Agropyron sp. India [126]
Uromyces agrostidis (Gonz. Frag.) A.L.
Guyot Agrostis capillaris and Ranunculus repens Europe [39]
Uromyces aimeae Berndt Cucurbitaceae Ecuador [127]
Uromyces airae-flexuosae Ferd. & Winge Aira flexuosa,Avenella flexuosa,
Deschampsia beringensis,Deschampsia flexuosa,
and Lerchenfeldia flexuosa
Bulgaria, Czech Republic, Czechoslovakia,
Denmark, Finland, Germany, Poland, and
Russia [82,83,106,128–135]
Uromyces albidum Kirgizb. Acanthophyllum pungens Uzbekistan [106]
Uromyces albiziae Henn. Albizia procera Indonesia [136]
Uromyces albucae Kalchbr. & Cooke Albuca altissima,A. aurea,A. minor,
A. canadensis,A. juncifolia, and A. wakefieldii Kenya, Malawi, South Africa [137–142]
Uromyces albus (Clinton) Dietel & Holw. Vicia americana and V. linearis California and Nebraska [78,143]
Uromyces algeriensis P. Syd. & Syd. Scilla obtusifolia and Scilla sp. Algeria, Cyprus, and Tunisia [144,145]
Uromyces alhagi Szemb. Alhagi camelorum and A. sparsifolium Russia and Uzbekistan [146]
Uromyces allii-monanthi Y. Harada Allium monanthum Japan [147]
Uromyces allii-sibirici Gjaerum Allium sibiricum Norway [148,149]
Uromyces allii-victorialis Liou & Y.C. Wang Allium fistulosum,A. macrostemon, and
A. victorialis China [90]
Uromyces aloes (Cooke) Magnus Aloe spp. India, Japan, and South Africa [43,133,139,150–154]

J. Fungi 2022,8, 633 19 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces alopecuri Seym.
Alopecurus aequalis,A. amurensis,
A. arundinaceus,A. arundinaceus,A. geniculatus,
A. japonicas,A. pratensis,Ranunculus sceluratus,
R. sieboldii, and R. vernyi
China, Colorado, Japan, Iowa, Minnesota,
Nebraska, Texas, Turkey, and Wyoming [80,92,96,155,156]
Uromyces alpestris Tranzschel Euphorbia cyparissias Bulgaria, Germany, Spain, and Switzerland [82,157–160]
Uromyces alsinis Tranzschel Minuartia hamata and M. meyeri Turkey [80,161]
Uromyces alstroemeriae (Dietel) Henn. Alstroemeria revoluta,A. aurantiaca,
A. caryophyllaea,A. inodora,A. ligtu,
A. isabellana,A. nemorosa, and A. subrosulacea America, Argentina, Brazil, and Chile [143,162–167]
Uromyces alysicarpi Wakef. & Hansf. Alysicarpus glumaceus,A. monilifer,
A. vaginalis, and A. violaceus India, Malawi, Uganda, South Africa, and
Zambia [138,139,141,168–172]
Uromyces alyxiae Arthur Alyxia oliviformis Hawaii [173–176]
Uromyces ambiens Cooke Buxus sempervirens and B. wallichiana India and Pakistan [177,178]
Uromyces ambiguus (DC) Lév.
Allium ampeloprasum,A. angulosum,
A. atroviolaceum,A. babingtonii,A. descendens,
A. fistulosum,A. rotundum, A. schoenoprasum,
A. sibiricum,A. ursinum,A. vineale, and A. waldsteinii
Bulgaria, Finland, Germany, Greece, Japan,
Poland, Romania, Ukraine, and
the United Kingdom [82,105,133,135,145,179]
Uromyces americanus Speg.
Cicuta bulbifera,Hydrocotyle bonariensis,
H. umbellata,Oenanthe sarmentosa,Scirpus americanus,
S. acutus,S. californicus,S. lacustris,S. olneyi,
S. validus,Sium cicutifolium, and Sium suave
Alabama, Argentina, Canada, California,
Delaware, Indiana, Nebraska, Puerto Rico,
Texas, Uruguay, Virgin Islands, West Indies,
and
Wisconsin
[180–185]
Uromyces amoenus Syd. & P. Syd. Anaphalis alpicola,A. busua, A. contorta,
A. margaritacea, and Gnaphalia margaritacea
Canada, California, China, Idaho, Japan,
Michigan, Montana, Nepal, Oregon, Russia,
South Dakota, and Washington [79,94,186–190]
Uromyces amphidymus Syd. & P. Syd. Glyceria acutiflora,G. borealis, G. fluitans, and
G. septentrionalis America, Indiana, New Jersey, Rhode
Island, and Wisconsin [79,191–193]
Uromyces amphilophis-insculptae
T.S. Ramakr., Sriniv. & Sundaram Amphilophis insculpta India [194]
Uromyces amurensis Kom. Maackia amurensis,M. floribunda,M. hupehensis,
M. tashiroi, and M. reticulata China, Japan, Korea, and Russia [92,94,96,108,109]

J. Fungi 2022,8, 633 20 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces anabasis Kazenas Anabasis aphylla China [195]
Uromyces anagyridis Roum. Anagyris foetida,A. mongolicus,A. nanus,
Piptanthus concolor, and P. nepalensis China, Cyprus, Greece, Iraq, Israel, Spain,
and Turkey [80,81,96,108,109,196–198]
Uromyces andropogonis Tracy
Andropogon elliottii,A. emersus,A. glomeratus,
A. perforatus,A. ternarius,A. saccharoides,
A. virginicus,Sorghum halepense,
Viola papilionacea,V. pedata,V. pedata,
V. striata, and V. tricolor
Alabama, Arkansas, Bolivia, Carolina,
Chile,
Columbia, Connecticut, Delaware, Ecuador,
Florida, India, Indiana, Illinois, Jersey,
Kentucky, Louisiana, Maryland,
Mississippi,
Missouri, New North Georgia, New York,
Ohio, Tennessee, Texas, and Virginia
[171,172,199–205]
Uromyces andropogonis-annulati
Syd., P. Syd. & E.J. Butler
Andropogon annulatus,A. abyssinicus,A. brevifolius,
A. brevifolius,A. hirtiflorus,A. ischaemum,A. longipes,
A. multinervis,A. pilosellus,A. platyphyllus,A. schottii,
Bothriochloa insculpta,B. ischaemum,B. pertusa,
Cymbopogon giganteus,Dichanthium annulatum,
D. aristatum,D. caricosum,D. nodosum,
Exotheca abyssinica,Hemarthria altissima,
Hyparrhenia hirta,Monocymbium ceresiiforme,
and Themeda triandra
Barbados, Cuba, Ethiopia, Guatemala,
India, Kenya, Malawi, Mauritius, México,
New Guinea, Pakistann, Philippines,
Uganda, Sierra Leone, Sudan, Tunisia, and
West Indies
[85,87,88,122,139,168,171,172,201,
206–209]
Uromyces anguriae H.S. Jacks. & Holw. Anguria warmingiana,Gurania pycnocephala,
Helmontia cardiophylla, and Wilbrandia verticillata Brazil and French Guiana [127,210–212]
Uromyces anomathecae Cooke Anomatheca cruenta South Africa [123,138,169,213]
Uromyces anotidis Petch Anotis richardiana Sri Lanka [214]
Uromyces anotidis-monospermatis
T.S. Ramakr. & Sundaram Anotis monosperma India [215]
Uromyces anthacanthi H.S. Jacks. Anthacanthus spinosus Puerto Rico, Virgin Islands, and West Indies [125,184,216–218]
Uromyces anthemophilus Vestergr. Bauhinia longifolia and Bauhinia sp. Mato Grosso [102,219]
Uromyces antholyzae Syd. & P. Syd. Antholyza abyssinica Ethiopia and South Africa [138,142,169]

J. Fungi 2022,8, 633 21 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces anthyllidis (Grev.) J. Schröt.
Anagyris spp., Astragalus spp., Benedictella spp.,
Coronilla spp., Dorycnium spp., Hedysarum spp.,
Hippocrepis spp., Lathurus spp., Lotus spp.,
Lupinus spp., Medicago spp., Ononis spp.,
Trifolium spp., Trigonella spp., and Vicia spp.
Algeria, Armenia, Austria, Belarus,
Bulgaria,
Cyprus, France, Germany, Greece,
Hungary, Iran, Itlay, Kyrgyzstan, Libya,
Morocco, Palestine,
Poland, Portugal, Romania, Spain,
Switzerland, Syria, Tunisia, Turkey,
Ukraine, Ukraine,
and Yugoslavia
[81,220–222]
Uromyces antiguanus Cummins Desmodium orbiculare and Desmodium sp. Guatemala (USA) and México [171,172,223–225]
Uromyces antioquiensis Mayor Rhynchospora polyphylla and R. nervosa Colombia [119,226,227]
Uromyces antipae Săvul. & O. Săvul. Rosa lutea Romania [228]
Uromyces aphelandrae Syd. Aphelandra pectinata Costa Rica [218,224,229]
Uromyces apiosporus Hazsl. Primula minima and Primula suffrutescens Austria, Bulgaria, Germany, Poland,
Romania, and California [78,82,135,230,231]
Uromyces apludae Syd., P. Syd. & E.J. Butler Apluda mutica,Indigofera linifolia, and I. cordifolia India, Guinea, Pakistan, and Philippines [85,208,232,233]
Uromyces appelianus Gassner Passiflora foetida and Cayaponia sp. Brazil, India, and Uruguay (USA) [127,211,212,234]
Uromyces appendiculatus (Pers.) Link Amphicarpaea spp., Cajanus sp., Dolichos spp.,
Lablab sp., Phaseolus spp., and Vigna spp. Worldwide [92,195,235–238]
Uromyces appendiculatus var. azukicola
Hirata Phaseolus angularis Japan [239]
Uromyces aquiriensis Berndt Cucurbitaceae Acre (Israel) [127]
Uromyces araucanus Dietel & Neger Senecio otites La Araucania (Chile) [162,240]
Uromyces archerianus Arthur & Fromme Chloris elegans Africa, China, Kenya, Malawi, and México
(USA) [108–110,171,172,241–245]
Uromyces arenariae Tranzschel Arenaria graminea, A. gypsophiloides, and
A. serpyllifolia Armenia, Romania, and Turkey [80,161,246]
Uromyces arenariae-grandiflorae Mayor Arenaria saponarioides Turkey [80]
Uromyces argaeus Maire Rumex tuberosus Asia [247]
Uromyces argutus F. Kern Spartina alterniflora and S. glabra Florida [106,242,248]
Uromyces argyrolobii Doidge Argyrolobium amplexicaule and Sesbania sp. KwaZulu-Natal and Zimbabwe (South
Africa) [138,249,250]

J. Fungi 2022,8, 633 22 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces ari-triphylli (Schwein.) Seeler Arisaema spp., Peltandra virginica, and
Zantedeschia sp.
Britain, California, Canada, Florida,
Indiana,
Illinois, Iowa, Minnesota, Mississippi,
México, Washington, and Virginia
[185,187,251–256]
Uromyces arizonicus Tracy & Galloway Eriogonum racemosum Arizona (USA) [257]
Uromyces armeriicola Speg. Armeria chilensis North, Central, and South America, and
West Indies [120,125,258–265]
Uromyces asclepiadis Cooke Asclepias spp. Argentina, Brazil, Bolivia, Colombia, Cuba,
California, Florida, Maine, Peru, Puerto
Rico, México, and Texas [100,118,119,123,124,262,266]
Uromyces asperulae McAlpine Asperula conferta and A. oligantha Australia [267,268]
Uromyces aspiliae H.S. Jacks. & Holw. Aspilia phyllostachya,Aspilia sp., and Wedelia saltensis Argentina and Rio de Janeiro (Brazil) [102,210,269]
Uromyces aspiliellus Vienn. -Bourg. Aspilia latifolia Ivory Coast [270]
Uromyces aspiliicola Cummins Aspilia asperifolia,A. africana,A. kotschyi,
Guizotia sp., and Wedelia sp. Malawi, Sudan, Tanzania, and Uganda [168,271–275]
Uromyces astragali-alopecuri Gjaerum Astragalus alopecurus Turkey [276]
Uromyces astragali-atropilosuli Gjaerum Astragalus atropilosulus var. bequaertii Kenya [276]
Uromyces astragalicola Henn. Astragalus adsurgens North Dakota and Utah [277,278]
Uromyces astragali-pseudoutrigeris Gjaerum Astragalus pseudoutriger Turkey [276]
Uromyces atlanticus Guyot & Malençon Hippocrepis scabra Morocco [279]
Uromyces atriplicis McAlpine Atriplex confertifolia,A. paludosa,A. semibaccata,
and A. vesicaria Australia, Britain, and Colorado [280–282]
Uromyces atropidis Tranzschel Atropidis distantis Turkey [161]
Uromyces aureus Dietel & Holw. Allium trinulatum,A. validum,Allium sp., and
Chlorogalum pomeridianum California and Washington (USA) [78,186,187]
Uromyces auriculae (Magnus) A. Buchheim Primula auricula Austria and Germany [160,283]
Uromyces azorellae Cooke Pozoa trifoliate and Schizeilematrifolio latum New Zealand [213,284]
Uromyces babianae Doidge Babiana disticha Western Cape Province (South Africa) [249,285]
Uromyces baccarinii Syd. & P. Syd. Wedelia sp. Eritrea 1937 [286,287]

J. Fungi 2022,8, 633 23 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces badius Syd. Haemanthus coccineus,H. pumilio,H. rotundifolius,
and H. sanguineus South Africa [138,167,288]
Uromyces baeumlerianus BubákMelilotus spp.
Austria, Bulgaria, China, Czech Republic,
Hungary, India, Israel, Japan, Poland,
Spain,
Romania, Turkey, and Ukraine
[81,82,96,289,290]
Uromyces bahiensis Perd. Sánch. Loranthaceae Panama [15]
Uromyces basellae Syd. & P. Syd. Basella rubra Malaysia [15]
Uromyces bauhiniae Henn. Bauhinia bongardii,B. cuyabensis,B. hiemalis,
B. longiflora,B. pauletia, and B. ungulata Brazil and México [102,291,292]
Uromyces bauhiniicola Arthur Bauhinia chlorantha and B. pringlei México [291]
Uromyces beckeropsidis E. Castell. Beckeropsis nubica Eritrea and Euthopia [106,293,294]
Uromyces beckmanniae H.S. Jacks. Beckmannia eruciformis and B. syzigachne Oregon [295]
Uromyces behenis (DC.) Unger Silene spp. Worldwide [296,297]
Uromyces belemensis F.C. Albuq. &
Figueiredo Ormosia nobilis Brazil [219,298]
Uromyces beloperones G.F. Laundon Beloperone californica,B. purpusii, and
Jacobinia mexicana Arizona, California, and México [116,218,299,300]
Uromyces bermudianus Cummins Cyperus paniculatus Bermuda [224,233]
Uromyces bethelii Arthur Silene verecunda California [301]
Uromyces beticola
(Bellynck) Boerema, Loer. & Hamers Beta vulgaris Bulgaria, Israel, and Turkey [80–82,302]
Uromyces bicolor Ellis Allium spp.
California, Idaho, Kansas, Massachusetts,
Missouri, Montana, Maine, New York,
Ohio,
and Texas
[186,191,303–305]
Uromyces bidenticola (Henn.) Arthur Bidens spp. and Cosmos caudatus Africa, Asia, Central and South America,
and the Southern United States [96,121,306–310]
Uromyces bidentis Lagerh. Bidens spp. and Cosmos caudatus Central and South America, Southern
United States, and West Indies [118,119,152,184,264,311–313]
Uromyces bisbyi Savile Eriogonum parvifolium California [314]

J. Fungi 2022,8, 633 24 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces blainvilleae Berk. Blainvillea spp. Brazil, Chinas, India, Nigeria, Sri Lanka,
and Tanzania [152,195,210,315–317]
Uromyces blandus Syd. Phragmites vulgaris China and Philippines [318,319]
Uromyces boissierae Vienn. Bourg. Boissiera pumilio Iran [320]
Uromyces bolusii Massee Aspalathus pachyloba South Africa [138,321,322]
Uromyces bomareae Henn. Bomarea sp. Brazil [167,261,323]
Uromyces bonaerensis Speg. Gomphrena elegans Buenos Aires [324]
Uromyces bonae-spei BubákTritonia scillaris and Acidanthera pallida Southern Africa [325]
Uromyces bonaveriae P. Syd. Bonaveria securidaca and Securigera securidaca Greece [326,327]
Uromyces borealis Peck Hedysarum boreale,H. mackenzii, and Rumex arifolius Canada and Finland [328,329]
Uromyces bornmuelleri Magnus Bongardia chrysogonum and Leontice armeniaca Cyprus, Iran, Iraq, Israel, and Turkey [81,196,330]
Uromyces borreriae Henn. Borreria verticillata Rio de Janeiro (Brazil) [331]
Uromyces bosseri Vienn. Bourg. Trochomeriopsis diversifolia Madagascar [127,332]
Uromyces bothriochloae-intermediae Gorlenko Bothriochloa intermedia China [108,109]
Uromyces bouvardiae Syd. & P. Syd. Bouvardia spp. Guatemala and México [281,292,333]
Uromyces bradburyae H.S. Jacks. & Holw. Bradburya pubescens,B. virginiana,
Centrosema pubescens, and C. virginianum Brazil [101,102]
Uromyces brandzae Săvul. Lathyrus aureus,L. venetus, and Orobus vernus Romania and Ukraine [111,126,228]
Uromyces brasilianus Speg. Senecio brasiliensis Buenos Aires [334]
Uromyces brasiliensis Trotter Jacquemontia spp. Brazil, China, and México [102,335,336]
Uromyces bravensis Cummins Sporobolus argutus and S. pyramidalis The Dominican Republic and Texas [171,172,337]
Uromyces bresadolae Tranzschel Euphorbia sp. Itlay [157]
Uromyces briardii Har. Euphorbia sp. Germany [338]
Uromyces brizae Gäum., E. Müll. & Terrier Briza media France [339]
Uromyces brodiaeae Ellis & Harkn Brodiaea spp., Hookera hyacinthina, and
Triteleia ixioides California, Oregon, and Washington (USA) [187,340]

J. Fungi 2022,8, 633 25 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces bromicola Arthur & Holw. Bromus coloratus and B. lithobius Chile [199]
Uromyces brominus Guceviˇc Bromus riparius Ukraine and Russia [106,111]
Uromyces buforrestiae Cummins Buforrestia imperforata Ghana [341]
Uromyces bugranae A.L. Guyot Ononis columnae and O. striata France, Switzerland [342]
Uromyces bulbinicola Doidge Bulbine bulbosa South Australia [249,343]
Uromyces bulbinus Thüm. Bulbine sp. Australia and New Zealand [344]
Uromyces bunsteri (Neger) H.S. Jacks. &
Holw. Sisyrinchium cuspidatum,S. graminifolium, and
Sisyrinchium sp. Chile [166]
Uromyces bupleuri Magnus Bupleurum spp. China, Greece, Iran, Morocco, and Tibet [91,190,220,345,346]
Uromyces bylianus Doidge Liliaceae KwaZulu-Natal [249]
Uromyces cacaliae (DC.) Unger Adenostyles spp. and Cacalia spp. Austria, Germany, Japan, Poland, Romania,
Russia, and Switzerland [92,135,159,160]
Uromyces cacciniae Jørst. Caccinia strigosa and Mattiastrum sp. Afghanistan and Iran [347,348]
Uromyces cachrydis Har. Cachrys pterochlaena,Hippomarathrum boissieri,
H. crassilobum,H. siculum, and Prangos sp. Cyprus, Israel, Libya, Morocco, and
Portugal [81,349]
Uromyces caladii (Schwein.) Farl. Arisaema spp., Peltandra spp., and
Muricauda dracontium
Carolina, Georgia, Iowa, Minnesota,
México,
Missouri, Oklahoma, Texas, and West
Virginia
[133,155,253,255]
Uromyces calamagrostidis Uljan. Calamagrostis arundinacea The former Soviet Union [106]
Uromyces callicarpae (Petch) Fujik. ex S. Ito Callicarpa spp. China, Japan, and Taiwan [94,96,97]
Uromyces calopogonii Cummins Calopogonium galactioides Guatemala [171,172,224,350]
Uromyces calotheus Syd. Urginea Sierra Leone [351]
Uromyces calycotomes Gäum. & Terrier Calycotome spinosa France [220]
Uromyces calystegiae de Bary ex Fuckel Convolvulus arvensis America, Australia, Italy, and Germany [352]
Uromyces camphorosmae (Castagne) Har. Camphorosma monspeliaca France [353]
Uromyces canavaliae J.M. Yen Canavalia microcarpa China and Taiwan [97,108,109]
Uromyces capensis Doidge Oenothera biennis Western Cape Province [138]
Uromyces capitatus Syd. & P. Syd. Desmodium yunnanensis China, India, and Pakistan [96,156,354–357]

J. Fungi 2022,8, 633 26 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces caraganae (Thüm.) Magnus Caragana arborescens Portugal [358,359]
Uromyces caraganicola Henn. Caragana chamlagu Asia, Africa, and Europe [81,225,236,360–363]
Uromyces caricis-brunneae Morim. Carex brunnea Japan [94]
Uromyces caricis-dolichocarpae Azbukina Carex dolichocarpa Sakhalin (Russia) [364]
Uromyces caricis-rafflesianae Mayor Carex rafflesiana Philippines [365]
Uromyces caricis-schmidtii Tomilin Carex schmidtii Khabarovsk [297,366]
Uromyces caricis-sempervirentis E. Fisch. Carex sempervirens,C. sempervirens,C. stenophylla,
Phyteuma betonicifolium, and P. orbiculare Austria, Europe, Germany, Poland,
Romania, Switzerland, and Turkey [43,80,160]
Uromyces carneus (Nees) Har. Astragalus alpinus and Oxytropis campestris Finland and Wyoming [329,367,368]
Uromyces carpathicus Namysł. Geranium phaeum Poland [369,370]
Uromyces carthagenensis Speg. Manihot spp. Argentina, Brazil, and Uruguay [371,372]
Uromyces cassiae-mimosoidis (Doidge)
Doidge Cassia mimosoides and Chamaecrista mimosoides South Africa [138,171,172,285]
Uromyces castaneus P. Syd. & Syd. Desmodium incanum,D. purpureum, and
Desmodium sp. Argentina and Brazil [171,269]
Uromyces cayaponiae Henn. Cayaponia sp. Santa Catarina [127]
Uromyces cearensis Berndt & F.O. Freire Ipomoea sp. Brazil [373]
Uromyces cedrelae (Henn.) Henn. Toona serrata Indonesia [177,374]
Uromyces celosiae Dietel & Holw. Celosia latifolia Argentina, Brazil, Costa Rica, Cuba,
Jamaica,
and México [167,263,269,308,375,376]
Uromyces celtidis Dietel Celtis sp. Brazil [102,375]
Uromyces cenisiae A.L. Guyot Ononis cenisia France [342]
Uromyces ceratocarpi Syd. & P. Syd. Ceratocarpus arenarius China and Russia [91,144]
Uromyces cestricola Speg. Cestrum pubeens and C. strigilatum Argentina and Bolivia [334,377]
Uromyces chaetobromi Gjaerum Chaetobromus dregeanus and C. schraderi South Africa [244,378]
Uromyces chaetolimonis Korbonsk. Chaetolimon sogdianum Tajikistan [379]
Uromyces charmelii Liou Phyteuma charmelii France [380]

J. Fungi 2022,8, 633 27 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces chenopodii J. Schröt Chenopodium sp., Salsola spp., and Suaeda spp. Around the world [381]
Uromyces chenopodii-fruticosi (DC.) M.
Abbasi & Aime Suaeda maritima England and Germany [47]
Uromyces chesneyae Tranzschel & Erem. Chesneya astragalina Central Asia and Russia [382,383]
Uromyces chevalieri A.L. Guyot Drimiopsis sp. Chari and Tropical Africa [384,385]
Uromyces chilensis Dietel & Neger Lathyrus magellanicus and L. multiceps Chile [162,386]
Uromyces chiovendae Bacc. Cissus sp. Somalia [287]
Uromyces chloridis Doidge Chloris myriostachya,C. pilosa, and C. virgate Uganda and South Africa [138,249]
Uromyces chlorogali Dietel & Holw. Chlorogalum pomeridianum California and Washington (USA) [186,187,387]
Uromyces chorizanthis Ellis & Harkn. Chorizanthe pungens California [340,388]
Uromyces christensenii J. Anikster & I. Wahl Muscari parviflorum and Hordeum bulbosum Israel [389,390]
Uromyces chubutensis Speg. Poa chubutensis Chubut [391]
Uromyces ciceris-arietini (Grognot) Jacz. & G.
Boyer Cicer arietinum,Trigonella polycerata, and Vicia spp. Brazil, India, Kenya, Libya, México,
Morocco,
and Pakistan [81,221,357,392]
Uromyces ciceris-soongaricae S. Ahmad Cicer songaricum Pakistan [393]
Uromyces circinalis Kalchbr. & Cooke Scilla prasina South Africa [138]
Uromyces circumscriptus Neger Loranthus spp., Struthanthus complexus, and
Phrygilanthus spp. Argentina, Brazil, and Chile [394]
Uromyces cisneroanus Speg. Excoecaria biglandulosa var. serrata and Sapium spp. Argentina, Brazil, Paraguay, and Venezuela [100,101,103,269,324]
Uromyces cladomanes Traverso Vitis sp. Somalia [287]
Uromyces cladrastidis Kusano Cladrastis shikokiana Japan [92]
Uromyces clarus H.S. Jacks. & Holw. Iresine angustifolia,I. celosia and Iresine sp. Bolivia, Brazil, Cuba, Venezuela, and
West Indies [167,263,376]
Uromyces clavatus Dietel Lathyrus spp. and Vicia spp. Argentina, Chile, and Uruguay [143,240,395]
Uromyces claytoniae Cooke & Peck Claytonia caroliniana Canada and New York [104,185,396]

J. Fungi 2022,8, 633 28 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces clignyi Pat. & Har.
Andropogon spp., Bothriochloa spp., Capillipedium sp.,
Cymbopogon sp., Dichanthium spp., Eremopogon sp.,
Exotheca sp., Hemarthria spp., Heteropogon sp.,
Monocymbium sp., Schizachyrium sp., Sorghum sp.,
and Themeda spp.
Uganda, Australia, Cameroon, China,
Ethiopia, Ghana, Guatemala, India, Japan,
Kenya, México, Nepal, Nigeria, Pakistan,
South Africa, Tanzania, Uganda, and
Zambia
[110,152,171,172,243,244,356,397]
Uromyces clignyioides Gjaerum Monocymbium ceresiiforme Zimbabwe [244]
Uromyces clitoriae Arthur Clitoria mexicana México [172,398]
Uromyces clivalis Mitter Argyrolobium flaccidum India [357,399]
Uromyces clutiae Kalchbr. & Cooke Clutia sp. Kenya [400]
Uromyces cnidoscoli Henn. Cnidoscolus vitifolius and Jatropha sp. Argentina and Brazil [103,282,291]
Uromyces cobresiae Korbonsk. Carex sp. Uzbekistan [146]
Uromyces colchici Massee Colchicum spectabile and C. bavaricum Great Britain and Turkey [133,401,402]
Uromyces collinus J.F. Hennen & Cummins Bauhinia sp. México [133,292,403]
Uromyces cologaniae Arthur Cologania spp. and Teramnus uncinatus Costa Rica, Guatemala, México, Puerto
Rico, and Venezuela [259,281,306,310,398]
Uromyces coloradensis Ellis & Everh. Astragalus sp. and Vicia spp. Arizona, California, Canada, Colorado,
Iowa,
Utah, Washington, and Wisconsin (USA) [225,404]
Uromyces columbianus Mayor Melanthera spp. Florida, West Indies, and Central and South
America [76,77,120,122,226,306]
Uromyces coluteae Arthur Colutea arborescens Austria [299]
Uromyces combreti Thaung Combretum Myanmar [405]
Uromyces comedens P. Syd. & Syd. Jasminum pubescens India and Dominican Republic [118,119,144,224,406]
Uromyces commelinae Cooke
Amischotolype sp., Aneilema spp., Callisia spp.,
Commelina spp., Cyanotis spp., Cyanotis spp.,
Pollia spp., Tradescantia spp., Tripogandra sp.,
and Zebrina sp.
Worldwide [222,407]
Uromyces compactus Peck Aster spinosus, Boltonia diffusa, and Baccharis sp. Arizona, México, and Texas [116,337,408,409]
Uromyces comptus P. Syd. & Syd. Ipomoea bipinnatipartita Namibia, South Africa, and Tanzania [126,138,165,384]
Uromyces congoensis Syd. & P. Syd. Bauhinia sp. The Democratic Republic of the Congo [410]

J. Fungi 2022,8, 633 29 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces conicus Jørst. Cleome sp. Bolivia [411]
Uromyces coordinatus Arthur Euphorbia spp. and Tithymalus palmeri California and Utah [103,114,404,412]
Uromyces corallocarpi W.T. Dale Corallocarpus emetocatharticus and
Doyerea emetocathartica Trinidad and Tobago, México, and West
Indies [127,211,212,308]
Uromyces cordiae Henn. Cordia sp. Rio de Janeiro [413]
Uromyces coronatus Yoshin. Helictotrichon milanjianum,Oryza sativa,
Zizania spp., and Zizaniopsis sp. China, Florida, Hong Kong, Japan, Korea,
Taiwan, Thailand, and Uganda [39,94–97,262,414]
Uromyces coronillae Vienn.-Bourg. Coronilla rostrate and C. varia Israel and Switzerland [81,220]
Uromyces correntinus J.C. Lindq. Rhynchospora tenuis Argentina [415]
Uromyces corrugatus Speg. Vicia patagonica Argentina and Brazil [126,180]
Uromyces costaricensis Syd. Lasiacis sp. and Panicum altissimum Brazil, Colombia, Costa Rica, Florida,
México, Puerto Rico, and Venezuela [106,229,294]
Uromyces costesianus Speg. Sphaeralcea velutina Chile [386,416]
Uromyces crassipes Dietel & Neger Rumex spp. Argentina, Chile, and Peru [269,281,376,417]
Uromyces crassivertex Dietel Lychnis miqueliana and Lychnis sp. China and Japan [92,94,418]
Uromyces crepidis-fraasii Kaps. Gotsi Crepis fraasii Greece [419]
Uromyces cretensis Petr. Coronilla parviflora and C. rostrata Greece [420,421]
Uromyces cristatulus Tranzschel Euphorbia sp. Australia, Germany, Hungary, and Russia [157]
Uromyces cristatus J. Schröt. & Niessl Dianthus caryophyllus,Lychnis viscaria,
Viscaria alpina, and V. vulgaris
Portugal, Czechoslovakia, Denmark,
Finland, Greece, Norway, Poland, and
Sweden [105,128,129,135,158]
Uromyces cristulatus Tranzschel Euphorbia barrelieri,E. petrophila,E. salicifolia, and
E. seguieriana Bulgaria and Ukraine [82,111,157,422]
Uromyces croci Pass. Crocus spp. Bulgaria, Israel, Romania, Russia, and
Ukraine [81,82,165,422]
Uromyces crotalariae-nitens Salazar-Yepes &
BuriticáCrotalaria nitens Colombia [423]
Uromyces cruchetii Mayor Borreria tenella Colombia [226]
Uromyces cruckshanksiae Cummins & Bonar Cruckshanksia bustillosii,C. palma,Oreopolus glacialis,
and O. palmae Argentina and Chile [233,240,415]

J. Fungi 2022,8, 633 30 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces cucubali Hirats. & Hashioka Cucubalus baccifer China, Japan, and Taiwan [89,96,97]
Uromyces cucullatus Syd. & P. Syd. Baltimora recta,Perymenium spp., Wedelia spp.,
and Zexmenia spp. Cuba, Costa Rica Guatemala, México, and
Panama [124,281,398,424,425]
Uromyces cucumivorus Berndt Cucumis melo var. flexuosus Iraq [127]
Uromyces cuenodii Maire Silene eriocalycina Iraq [196]
Uromyces cuspidatus G. Winter Festuca spp., Melica laxiflora,
Muhlenbergia spp., and Poa chubutensis Argentina, Bolivia, and Chile [106,199,240]
Uromyces cyanotidis Cummins Cyanotis capitata Papua New Guinea [233,426]
Uromyces cyathulae Henn. Cyathula globulifera Eritrea [287,427]
Uromyces cyperi Henn. Cyperus sp. Eritrea [287,427]
Uromyces cypericola Gjaerum Cyperus albostriatus and C. cyperoides Kenya, South Africa, and Uganda [428,429]
Uromyces cyprius Vienn.-Bourg. Rumex cyprius Iran [430]
Uromyces cystopiformis Lagerh. Centropogon sp. and Siphocampylus sp. Costa Rica, Ecuador [165,310,431]
Uromyces cytisi J. Schröt. Caragana arborescens,Cytisus polytrichus,
Genista aucheri,G. tinctoria, and Laburnum
anagyroides Russia, Ukraine, and Turkey [80,111,432]
Uromyces dactylidis G.H. Otth Agrostis spp., Alopecurus spp., Anemone sp., Avenula
sp., Dactylis spp., Festuca spp., Poa spp., Ranunculus
spp., and Trisetum sp.
Arizona, Armenia, Australia, Bulgaria,
Pakistan, Canada, China, Checz Republic,
Czechoslovakia, Denmark, Finland,
Germany,
Hungary, Japan, New Zealand, Norway,
Oklahoma, Pennsylvania, Romania, Spain,
Sweden, Switzerland, Ukraine, Virginia,
Vermont, and other temperateregions of the
world
[43,80,82,105,113,368,409,433–441]
Uromyces dactyloctenii Wakef. & Hansf. Dactyloctenium aegyptium China, Japan, Kenya, Namibia, Samoa,
Tanzania, Tonga, and Uganda [140,168,274,442,443]
Uromyces dactylocteniicola (Speg.) J.C.
Lindq. Dactyloctenium aegyptium Central Africa, the Philippines, and South
America [106]
Uromyces danthoniae McAlpine Danthonia spp. and Rytidosperma spp. Australia and New Zealand [245,280,313]

J. Fungi 2022,8, 633 31 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces decoratus Syd. & P. Syd. Crotalaria spp.
China, Costa Rica, Guinea, Ghana, India,
Japan, Madagascar, Myanmar, Nigeria,
Pakistan, Sri Lanka, Taiwan, Thailand, and
Venezuela
[85,86,94,96,97,152,171,172,281,290,
312,356,357,444,445]
Uromyces deeringiae Syd. & P. Syd. Cladostachys polysperma and Deeringia spp. China, Japan, India, Indonesia, Philippines,
and Taiwan [92,167,232,309,446,447]
Uromyces delagoënsis BubákLapeirousia delagoensis Mozambique [249]
Uromyces dendroseridis Keissl. Dendroseris micrantha Chile [240]
Uromyces densus Arthur Bidens pilosa Puerto Rico [76,77]
Uromyces desmodii Cooke Desmodium canescens South Carolina and Brazil [448]
Uromyces desmodiicola Jørst. Desmodium albiflorum Brazil [102,171,449]
Uromyces desmodii-leiocarpi Henn. Desmodium leiocarpum Brazil [171,450]
Uromyces devoluensis Gäum. Senecio doronicum France [451]
Uromyces dianthi (Pers.) Niessl
Arenaria leptoclados,Bufonia spp.,
Cerastium brachypetalum,Dianthus spp.,
Euphorbia spp., Gypsophila spp., Petrorhagia spp.,
Silene sp., Vaccaria sp., and Vicia sp.
Australia, Bermuda, Bolivia, Bulgaria,
Brazil,
California, Chile, Cuba, Germany, Japan,
Kenya, Madagascar, New York, Poland,
Romania, Spain, Thailand, Texas, Turkey,
Ukraine, and
Washington (USA)
[80,82,94,105,111,135,140,452]
Uromyces dianthi-caryophylli Monchot Dianthus sp. France [453]
Uromyces dichromenae W.T. Dale Dichromena radicans Jamaica, Trinidad and Tobago, and West
Indies [264,308]
Uromyces dictyospermae Ellis & Everh. ex
Tranzschel Euphorbia spp., Jatropha spp., and Tithymalus sp. California, Florida, Minnesota, Montana,
Oklahoma, and Washington (USA) [103,157,187,254,262,278,388,454–
456]
Uromyces didymae Gapon. Veronica polita Uzbekistan [457]
Uromyces dieramatis Doidge Dierama spp. South Africa [123,138,139,273,458]
Uromyces dietelianus Pazschke Bauhinia spp. Argentina, Brazil, and Uruguay [100,101,269,297]
Uromyces digitariae-adscendentis Y.C. Wang Digitaria spp. China and Taiwan [96,97]
Uromyces dilucidus Cummins Sisyrinchium striatum Argentina [415,459]

J. Fungi 2022,8, 633 32 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces diniensis A.L. Guyot Ononis fruticosa France [460]
Uromyces dinteri Mennicken, W. Maier &
Oberw. Tetraena spp. Namibia and Egypt [385]
Uromyces dipcadi Gjaerum Dipcadi viride Kenya [461,462]
Uromyces discariae G. Cunn. Discaria toumatou New Zealand [284,313,463]
Uromyces dispersus Hirats. f. Apios fortunei Japan [222]
Uromyces dobremezii Durrieu Euphorbia stracheyi Nepal [464]
Uromyces doebbeleri Berndt Hypericum irazuense Costa Rica [310]
Uromyces dolichi Cooke Dolichos axillaris Brazil, China, Madagascar, Puerto Rico,
Zimbabwe, and Uganda [108,109,125,171,172,250,465]
Uromyces dolicholi Arthur Cajanus spp., Dolicholus spp. and
Rhyncosia spp.
Canada, China, Colombia, Costa Rica,
Florida, India, Japan, Jamaica, Kenya,
Panama, Puerto Rico, South Africa, Taiwan,
Texas, Venezuela, and West Indies
[96,97,138,227,310,357,466–469]
Uromyces dolichosporus Dietel & Holw. Tournefortia spp. Argentina, Brazil, Bolivia, Cuba, Ecuador,
Jamaica, México, West Indies, and
Venezuela [100,101,263,269,308,375,456]
Uromyces doricus Maire Silene spp. Bulgaria and Greece [82,308,327,470]
Uromyces dorystaechadis Gjaerum & Bahç. Dorystaechas hastata Turkey [471]
Uromyces drimiopsidis Doidge Drimiopsis maculata South Africa [138]
Uromyces dubiosus Henn. Lantana sp. Goiás[291]
Uromyces ducellieri Maire Anabasis aphylla China [91,96,472]
Uromyces dusenii Dietel & Neger Gilliesia graminea,G. monophylla, Miersia chilensis,
and Ornithogalum biflorum Chile [166,240,417]
Uromyces echinodes Kunze ex Henn. Asclepiadaceae Suriname [473]
Uromyces ecklonii BubákFreesia refracta Kenya and South Africa [138,140]
Uromyces eclipsis Berndt Zygophyllum morgsana South Africa [234]
Uromyces edwardsiae G. Cunn. Edwardsia spp. and Sophora spp. New Zealand [284,463,474]
Uromyces ehrhartae McAlpine Ehrharta stipoides Australia and New Zealand [106,280,313,474]

J. Fungi 2022,8, 633 33 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces ehrhartae-giganteae Doidge Ehrharta spp. South Africa [138,249,378,429]
Uromyces elegans (Berk. & M.A. Curtis)
Lagerh. Trifolium spp. Florida, Kenya, Mississippi, Oklahoma,
South Carolina, Texas, and Uganda [168,261,455,475]
Uromyces eleocharidis Arthur Eleocharis spp. Iowa, Kansas, North and South Dakota,
and
Washington (USA) [155,278,301,466,476]
Uromyces ellipticus Dietel & Neger Glycyrrhiza astragalina Chile [240,477]
Uromyces ellisianus Henn. Euphorbia marginata Minnesota [277]
Uromyces emmeorhizae Syd. Emmeorhiza umbellata Venezuela [478]
Uromyces epicampis Dietel & Holw. Epicampes macroura,Melica laxiflora, and
Muhlenbergia spp. Arizona, California, Chile, Ecuador,
Guatemala, México, and Texas [106,116,199,240,281,337,479]
Uromyces eragrostidicola Gjaerum Eragrostis rigidior and Eragrostis sp. Ethiopia, Kenya, Tanzania, and Zimbabwe [244]
Uromyces eragrostidis Tracy Anthericum torreyi,Cypholepis yemenica,
Desmostachya bipinnata,Eragrostis spp., and
Tripogon chinensis
Arizona, Argentina, Australia, Botswana,
Brazil, China, Georgia, Ghana, India, Israel,
Kenya, Malawi, Malaysia, Mississippi,
México, Nebraska, Oklahoma, Pakistan,
Puerto Rico, South Africa, Texas, West
Indies,
and Venezuela
[90,110,116,141,171,172,199,208,
243,244,250,480]
Uromyces eriochloae
(Syd. & P. Syd.) Syd., P. Syd. & E.J. Butler Eriochloa spp. China, Indonesia, Japan, Madagascar, and
Philippines [85,90,232,309,465]
Uromyces eriogoni Ellis & Harkn. Eriogonum virgatum California [340,388]
Uromyces eriospermi Kalchbr. & Cooke Eriospermum spp. South Africa, Tanzania, and Zimbabwe [137,138,250,274]
Uromyces ermelensis Doidge Indigofera sp. South Africa [138]
Uromyces ervi (Wallr.) Westend. Ervum spp., Lens spp., Pisum sativum, and Vicia spp.
Algeria, Austria, Belgium, Bulgaria, China,
Czech Republic, Denmark, Finland, France,
Germany, Hungary, Italy, Japan, Morocco,
Netherlands, Pakistan, Poland, Portugal,
Romania, Russia, Spain, Switzerland,
Sweden, Taiwan, and Ukraine
[82,96,128,220,393,481,482]
Uromyces erythrinae Lagerh. Erythrina sp. Ecuador [165]
Uromyces erythronii (DC.) Pass. Erythronium sp., Lilium spp., and
Tulipa spp. Bulgaria, China, Germany, Greece, Japan,
Japan, Korea, and Spain [82,92,96,158,179,483]

J. Fungi 2022,8, 633 34 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces eugenei-mayorii M. Morelet Ulex europaeus Italy [451]
Uromyces eugentianae Cummins Gentiana spp. and Halenia guatemalensis Canada, Iowa, Norway, and Washington
(USA) [129,187,252,484–486]
Uromyces eulophiae Gjaerum Eulophia paivaena subsp. borealis India, Rwanda, and Uganda [318,487,488]
Uromyces euphaeus Syd. Hypoxis glabella Australia [489]
Uromyces euphlebius Syd. & P. Syd. Phoradendron calyculatus, and Phoradendron sp. México [15,490]
Uromyces euphorbiae Cooke & Peck Acalypha communis, Chamaesyce sp.,
Euphorbia spp., and Poinsettia heterophylla
Afghanistan, Argentina, Azirona, Brazil,
California, Canada, China, Cuba,
Columbia,
Costa Rica, Cyprus, India, Israel, Jamaica,
Japan, Mississippi, Mauritius, New York,
Pakistan, Panama, South Africa, Texas,
Uganda, Venezuela, West Indies, and
Zambia
[96,102,203,209,227,255,263,269,
300,308,310,318,348,469,475,488,
491,492]
Uromyces euphorbiae-connatae Speschnew Euphorbia sp. Russia [493]
Uromyces euphorbiae-javanicae E. Fisch. Euphorbia javanica Indonesia [309]
Uromyces euphorbiae-lunulatae Liou & Y.C.
Wang Euphorbia esula,E. kansui, and E. lunulata China [96,108,109]
Uromyces euphorbiae-nicaeensis Unamuno Euphorbia nicaeensis Spain [494]
Uromyces euphorbiae-polytimeticae Zenkova Euphorbia polytimetica Tajikistan [495]
Uromyces euphorbiicola
(Berk. & M.A. Curtis) Tranzschel Euphorbia spp. Brazil, Colombia, Madagascar, South
Africa,
and Zimbabwe [102,138,157,250,465]
Uromyces eurotiae Tranzschel Ceratoides latens,Krascheninnikovia ceratoides, and K.
latens China, Turkey, and Uzbekistan [80,91,146,161]
Uromyces euryopsidicola A.R. Wood & M.
Scholler Euryops empetrifolius and E. tenuissimus South Africa, North and Western Cape
Province [496]
Uromyces evastigatus Cummins Phthirusa pyrifolia El Salvador [15]
Uromyces fallens (Arthur) Barthol. Trifolium pratense Iran [497]
Uromyces fatouae Henn. Fatoua pilosa Japan [498]
Uromyces fedtschenkoi Faizieva Rumex fedtschenkoi Uzbekistan [457]
Uromyces ferganensis Tranzschel & Erem. Stipa caucasica,S. holosericea, and S. lessingiana Iran, Siberia, and Russia [106,382,499]

J. Fungi 2022,8, 633 35 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces ferrariae Doidge Ferraria sp. South Africa and Zimbabwe [138,250]
Uromyces ferulae Juel Ferula spp. and Heracleum cachemiricum Australia, Iran, Israel, Pakistan, and
Morocco [81,500]
Uromyces ferulaginis Lindr. Feruladinis silvaticae Poland [501]
Uromyces festucae-nigricantis Gonz. Frag. Festuca nigricans Morocco and Spain [279,502]
Uromyces ficariae (Schumach.) Lév. Ficaria spp. and Ranunculus spp.
Bulgaria, Czech Republic, Czechoslovakia,
Denmark, Finland, Germany, Iran, Norway,
Poland, Russia, Sweden, Turkey, and
Ukraine
[80,82,83,105,111,128,129,132,432,
438,503]
Uromyces fiebrigii Henn. & Vestergr. Bauhinia sp. Paraguay [216]
Uromyces fiorianus Sacc. Peucedanum fraxinifolium and Peucedanum sp. South Africa [138,504]
Uromyces fischerianus Mayor Ranunculus glacialis Switzerland [119]
Uromyces fischeri-eduardi Magnus Vicia spp. Bulgaria, Mongolia, Poland, Romania, and
Turkey [80,82,135,505]
Uromyces flavicomae Liou Euphorbia flavicoma France [506]
Uromyces flemmingiae Henn. Flemmingia sp. Uganda [507]
Uromyces fleuryae J.M. Yen Fleurya podocarpa Gabon [508]
Uromyces floralis Vestergr. Bauhinia hiemalis,B. cuyabensis,B. holophylla,
B. rufa, and Bauhinia sp. Brazil [102,216]
Uromyces floscopae Syd. & P. Syd. Floscopaperuviana and Floscopa sp. Brazil [102,509]
Uromyces fontii Gonz. Frag. Peplis acutangula Morocco [510]
Uromyces formosus Syd. & P. Syd. Dianthus libanotis Israel and Iran [81]
Uromyces foveolatus Juel Bauhinia hirsuta,B. mirandina, and Bauhinia sp. Brazil [100–102,511]
Uromyces fragilipes Tranzschel Agropyron squarrosum,Agrostis spp.,
Deschampsia spp., Eremopyrum bonaepartis,
Festuca spp., Hordeum spp., and Scribneria bolanderi California, Iran, Pakistan, and Russia [106,161,347,412,512,513]
Uromyces fraserae Arthur & Ricker Frasera speciosa Wyoming [514]
Uromyces fremonti Syd. & P. Syd. Oenothera fremontii Kansas [79]
Uromyces fulgens BubákChamaecytisus spp., Cytisus spp., and
Lembotropis nigricans
Bulgaria, Czech Republic, France, Greece,
Hungary, Itlay, Poland, Romania, Russia,
Ukraine, and Yugoslavia [82,220,421]

J. Fungi 2022,8, 633 36 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces fuscatus Arthur Polygonum alpinum Idaho and Utah [281]
Uromyces fusisporus Cooke & Massee Acacia neriifolia,A. salicina, and Acacia sp. Australia [177,515]
Uromyces gaeumannii Mayor & Vienn.
Bourg. Hippocrepis spp. France, Greece, Libya, Morocco, Palestine,
Portugal, Spain, Tunisia, and Yugoslavia [220,516]
Uromyces gageae Beck Gagea spp., Lloydia triflora, and
Ornithogalum umbellatum
Austria, Bulgaria, Czech Republic, China,
Denmark, Finland, Germany, Japan,
Norway,
Poland, Romania, Russia, Turkey, Sweden,
and Ukraine
[54,80,82,128,135,222,228,440]
Uromyces galactiae Rezende & Dianese Galactia pedunculata Brazil [517]
Uromyces galegae Sacc. Astragalus glycyphylloides and Galega spp. Czech Republic, France, Greece, Hungary,
Italy, Poland, Russia, Turkey, and
Yugoslavia [80,82,220,421]
Uromyces galegicola Woron. Galega orientalis Armenia, Romania, and Turkey [80,228,518,519]
Uromyces galii Dietel Galium aparine and G. spurium Japan [222,414]
Uromyces galii-californici Linder Galium californicum and Galium sp. California [520]
Uromyces galphimiae Dietel & Holw. Galphimia glauca and G. humboldtiana México [456,479]
Uromyces garanbiensis
(Hirats. f. & Hashioka) Sawada Ehretia dicksonii Taiwan [521]
Uromyces gaubae Petr. Caltha introloba Australia [346,522]
Uromyces gausseni Mayor & Vienn.-Bourg. Dorycnopsis gerardii France [516]
Uromyces geissorhizae Henn. Geissorhiza sp. Western Cape Province [523]
Uromyces gemmatus Berk. & M.A. Curtis Convolvulus parviflorus and
Convolvulus sp.
Brazil, China, Cuba, Ecuador, India, Japan,
Jamaica, México, Taiwan, West Indies,
and Venezuela
[94,96,97,100,101,122,264,306,319,
524,525]
Uromyces genistae Fuckel Cytisus capitatus,Genista tinctoria, Genista sp., and
Tithymalus cyparissias Czech Republic, Morocco, and Turkey [80,83,220,525]
Uromyces geranii (DC.) G.H. Otth & Wartm. Erodium spp. and Geranium spp.
Australia, Bulgaria, China, Chile,
Czech Republic, Denmark, Finland, France,
Germany, India, Japan, Kenya, Korea,
Norway, Pakistan, Poland, Portugal,
Romania, Russia, Spain, Sweden, Turkey,
Ukraine, and Zimbabwe
[80,82,87,88,94,96,128,179,208,436,
526]

J. Fungi 2022,8, 633 37 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces geraniicola Speg. Geranium patagonicum Chile [240,258]
Uromyces ghaznicus Petr. Limonium sp. Afghanistan [527]
Uromyces giganteus Speg. Kalidium foliatum and Suaeda spp. Australia, Arizona, China, California,
Idaho,
Texas, and Ukraine [47,96,300,337,421]
Uromyces gigantiformis Salazar-Yepes &
BuriticáBidens sp. Colombia [423]
Uromyces gilgitae S. Ahmad Sophora alopecuorides Pakistan [528]
Uromyces gladioli Henn. Babiana spp., Geissorhiza spp., Gladiolus spp.,
Moraea ramose, and Romulea spp. Cuba, Kenya, Malawi, Nigeria,
South Africa, and Uganda [138,140,168,273,291,458,529,530]
Uromyces globosus Dietel & Holw. Sapium spp. México [103,456,479]
Uromyces glyceriae Arthur Glyceria spp. Illinois, Indiana, New Jersey, Rhode Island,
and Wisconsin (USA) [39,191,299]
Uromyces glycyrrhizae (Rabenh.) Magnus Glycyrrhiza spp.
Armenia, Algeria, Azerbaijan, Bulgaria,
Caucasus, Czech Republic, China, Greece,
Iraq, Israel, Itlay, Japan, Kazakhstan,
Pakistan, Palestine, Portugal, Romania,
Spain, Turkey, and Uzbekistan
[39,78,80–82,91,96,116,146,196,278,
317,327,356,393,409,531–536]
Uromyces gnaphalii Ellis & Everh. Gnaphalium sp. Colorado [388]
Uromyces gouaniae F. Kern Gouania domingensis,G. lupuloides,
Ledenbergia macrantha, and Ledenbergia sp. Guatemala and México [281,486,537]
Uromyces goyazensis Henn. Bauhinia sp. Brazil [291,538]
Uromyces graminis (Niessl) Dietel Caucalis platycarpos,Ferula communis,
Melica spp., and Thapsia garganica
Afghanistan, Armenia, Bulgaria, Greece,
Italy, Morocco, Romania, Spain, Portugal,
and Ukraine
[82,111,158,228,348,421,519,534,
539]
Uromyces grandiotii Gäum. Ancrumia cuspidata Chile [540]
Uromyces greenstockii Doidge Ipomoea greenstockii South Africa [138]
Uromyces guatemalensis Vestergr. Bauhinia inermis,B. ungulata, and Bauhinia sp. Costa Rica, El Salvador, Guatemala, and
Venezuela [216,281,444,541]
Uromyces guayacuru Speg. Statice brasiliensis Buenos Aires [542]

J. Fungi 2022,8, 633 38 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces gueldenstaedtiae Liou & Y.C. Wang Populus sp. China [543]
Uromyces guerkeanus Henn. Lotus spp. and Tetragonolobus biflorus Belarus, Egypt, France, Greece, Italy, Iraq,
Malta, Morocco, Palestine, Portugal, Spain,
Syria, Tunisia, and Yugoslavia [196,220,422,426,544]
Uromyces guraniae Mayor Gurania sp. Brazil and Colombia [127,212,226]
Uromyces gypsophilae Cooke Gypsophila sp. Asia, Europe, Turkey, Iran, and Iraq [80,82,165,196,317,346,545–547]
Uromyces habrochloae Gjaerum Habrochloa bullockii Malawi [244]
Uromyces hainanicus J.Y. Zhuang & S.X. Wei Ipomoea sumatrana China [319]
Uromyces halimodendri Solkina Halimodendron argenteum and H. halodendron China and Uzbekistan [198,220]
Uromyces handelii BubákLotus gebelia Iraq [220,548]
Uromyces haraeanus Syd. & P. Syd. Scirpus spp. China, Japan, and Russia [90,92,297,549]
Uromyces hardenbergiae McAlpine Hardenbergia monophylla Australia [280]
Uromyces hariotianus Lagerh. Odontonema callistachyum,
Pseuderanthemum cuspidatum,Pseuderanthemum sp.,
andThyrsacanthus strictus Costa Rica and México [218,281,550]
Uromyces harmsianus (Henn.) Doidge Crotalaria spp. India, Kenya, Malawi, Rwanda, South
Africa,
Tanzania, and Uganda [138,168,171,172,551]
Uromyces haussknechtii Tranzschel Euphorbia thamnoide and E. pilosa India and Syria [157,350]
Uromyces hawksworthii
É.S.C. Souza, Z.M. Chaves, W.R.O. Soares,
Pinho & Dianese
Phthirusa stelis Brazil [60]
Uromyces hedysari-obscuri (DC.) Carestia &
Picc. Hedysarum spp.
Alaska, Austria, Canada, Czech Republic,
China, Germany, France, Finland, Hungary,
India, Italy, Kazakhstan, Mongolia,
Romania,
Russia, Switzerland, and Tajikistan
[94,96,220,357,436,535,552,553]
Uromyces hedysari-paniculati (Schwein.) Farl. Desmodium spp. and Meibomia spp.
America, Argentina, Bolivia, Brazil,
Canada,
Colombia, Costa Rica, Cuba, Indiana,
Jamaica, Massachusetts, México, Trinidad
and Tobago,
Venezuela, West Indies, and Vermont
[87,88,171,172,185,191,310,415]

J. Fungi 2022,8, 633 39 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces heimerlianus Magnus Ervum spp., Euphorbia spp., Lathyrus spp.,
and Vicia spp.
Armenia, Austria, Belarus, Bulgaria, China,
Czech Republic, France, Germany,
Hungary,
Italy, Japan, Portugal, Romania, Russia,
Serbia, Spain, Switzerland, Ukraine,
and Yugoslavia
[220,519,554]
Uromyces heimii Mayor & Vienn. Bourg. Medicago arborea France [516]
Uromyces helichrysi Lagerh. Helichrysum plicatum,Helichrysum rupestre,
and Helichrysum siculum Algeria, Greece, and Turkey [80,421,435]
Uromyces heliotropii Sred. Heliotropium spp. Australia, Bulgaria, Cyprus, Greece, Israel,
Pakistan, Ukraine, and Uzbekistan [80,81,111,145,393,421,555,556]
Uromyces hellebori-thibetani J.Y. Zhuang &
S.X. Wei Helleborus thibetanus China [195]
Uromyces hemmendorfii Vestergr. Bauhinia forficata São Paulo [219]
Uromyces hermonis Magnus Euphorbia herniariifolia and E. peplus Greece and Iraq [196,345,421]
Uromyces herterianus Dietel Borreria verticillata,Diodia sp. and
Spermacoce verticillata Argentina, French Guiana, and Uruguay [395,415,557]
Uromyces hessii Berndt Zantedeschia angustiloba Angola [558]
Uromyces heterantherae (Henn.) P. Syd. &
Syd. Heteranthera reniformis Brazil [102,144]
Uromyces heterodermus Syd. & P. Syd. Erythronium spp. Alberta, California, Canada, Columbia,
Colorado, Idaho, Montana, Oregon, Texas,
Utah, and Washington (USA) [79,185,187,559]
Uromyces heterogeneus Cooke Hibiscus syriacus India [126,560]
Uromyces heteromallus Syd. Haloxylon recurvum Pakistan [208,356,561–563]
Uromyces heteromorphae Thüm. Heteromorpha spp. and Peucedanum spp. Eritrea, Malawi, and South Africa [138,273,287,564]
Uromyces hewittiae Syd. & P. Syd. Hewittia bicolor and Hewittia sp. Philippines [565]
Uromyces hidakaensis Muray. & Takeuchi Pisum sativum Japan [222,566]
Uromyces himalaicus Y. Ono, Adhikari &
Rajbh. Lilium sp. Nepal [567]
Uromyces hippocrepidis Syd. & P. Syd. Hippocrepis ciliate and H. comosa Greece, France, Spain, and Switzerland [220,421,568]

J. Fungi 2022,8, 633 40 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces hippomarathri Lindr. Hippomarathrum crispum,H. libanotis, and
H. microcarpum Iran and Morocco [279,317,569]
Uromyces hippomarathricola Sousa da
Câmara Hippomarathrum pterochlaenum Portugal [570]
Uromyces hobsonii Vize Jasminum spp. Euthopia, India, Kenya, and Sri Lanka [140,571,572]
Uromyces holci Jørst. Holcus setiger,Karroochloa sp., Schismus barbatus,
S. scaberrimus, and Tribolium echinatum Namibia and South Africa [124,378]
Uromyces holubii Doidge Dracaena sp. Gauteng [573]
Uromyces holwayi Lagerh. Chlorophytum sp. and Lilium spp. China, California, Canada, Idaho, Iowa,
Japan, New York, Nebraska, New Jersey,
Oregon, Pakistan, and Washington (USA) [94,96,187,412,559]
Uromyces hordeastri A.L. Guyot Bellevalia flexuosa,Hordeum spp., and
Muscari parviflorum France and Israel [81,126]
Uromyces houstoniatus J. Sheld. Houstonia spp. and Sisyrinchium spp.
Connecticut, Illinois, Kansas,
Massachusetts,
Missouri, Mississippi, New York,
Pennsylvania, Vermont, and Wisconsin
(USA)
[191,251]
Uromyces howei (Peck) De Toni Asclepias curassavica,A. guatemalensis, and A. nivea Bermuda, Cuba, Guatemala, Jamaica, and
Peru [260,281,308,574]
Uromyces huallagensis Henn. Desmodium sp. Peru [575]
Uromyces hyacinthi W. Schneid. Hyacinthus fastigiatus Canada [115]
Uromyces hyalinus Peck Sophora nuttalliana,S. sericea, and S. stenophylla Arizona, Colorado, México, and Wyoming [116,225,292,559,576]
Uromyces hybridi W.H. Davis Trifolium hybridum Connecticut, India, Massachusetts, Maine,
and Vermont [191,577,578]
Uromyces hyderabadensis
Ramachar, K.N. Rao & Bagyan. Atylosia scarabaeoides India [579]
Uromyces hydrocotylicola J.Y. Zhuang Hydrocotyle sp. China [96,580]
Uromyces hymenocarpi Jaap Hymenocarpos circinnatus Croatia, Greece, Israel, and Turkey [80,81,421,581]
Uromyces hyparrheniae Gjaerum Hyparrhenia spp. Angola, Cameroon, Ethiopia, and Uganda [243,244]
Uromyces hyparrheniicola Gjaerum Hyparrhenia dregeana South Africa [243]

J. Fungi 2022,8, 633 41 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces hyperici (Schwein.) M.A. Curtis Ascyrum hypericoides,Elodea sp., and Hypericum spp.
China, California, Florida, Georgia, Indiana,
Iowa, Japan, Massachusetts, Mississippi,
Missouri, Maine, New York, Texas, Uganda,
Vermont, and Wisconsin (USA)
[89,90,186,202,251,262,582,583]
Uromyces hyperici-frondosi (Schwein.)
Arthur Hypericum spp. and Triadenum virginicum Brazil, Chile, California, Colombia,
Guatemala, Maine, Massachusetts,
Pennsylvania, South Africa, and Vermont
[78,101,118,119,138,191,281,520,
584]
Uromyces hypericinus Speg. Hypericum brasiliense Formosa [180]
Uromyces hypoestis Tarr & G.F. Laundon Hypoestes verticillaris South Africa, Sudan, Tanzania, and
Zimbabwe [218,250,274]
Uromyces hypsophilus Speg. Euphorbia sp. Mendoza [391]
Uromyces ictericus Cummins Iresine celosia and Iresine sp. Costa Rica and Guatemala [167,233,310]
Uromyces ignobilis (Syd. & P. Syd.) Arthur Muhlenbergia sp. and Sporobolus spp.
Barbados, Cuba, Dominica, Grenada,
Guyana,
Indonesia, Hawaii, México, Pakistan, Texas,
and Venezuela
[76,122,171,172,306,309,356,430,
431,445,456,559,585,586]
Uromyces illotus Arthur & Holw. Mucuna andreana,M. rubro-aurantiacea,M. sloanei,
and M. urens The Dominican Republic, Ethiopia, and
Guatemala [281,406,571]
Uromyces imperfectus Arthur Bauhinia spp. Jamaica, Nicaragua, and Venezuela [225,308,444,586]
Uromyces inaequialtus Lasch Cerastium sp., Melandrium sp., and Silene sp.
Argentina, Bulgaria, Canada, Chile, China,
Finland, Greece, Japan, Nepal, Norway,
Romania, Taiwan, South Africa, Sweden,
and Ukraine
[82,91,92,97,111,128,135,142,185,
188,240,415,421,587]
Uromyces inayatii Syd. & P. Syd. Apluda aristata and A. mutica India and Papua New Guinea [85,426]
Uromyces indicus Pat. Sporobolus indicus Barbados [588,589]
Uromyces indigoferae Dietel & Holw. Indigofera spp. Argentina, Arizona, Costa Rica, China,
Guatemala, Florida, India, México, Panama,
Texas, and Venezuela
[90,116,171,172,262,281,292,357,
375,415,425,492,559,590]
Uromyces induratus Syd., P. Syd. & Holw. Dicliptera spp., Jacobinia sp., and Justicia racemosa Argentina, Bolivia, Brazil, Costa Rica, and
México [102,218,292,310,333]
Uromyces infarctus Berndt Cayaponia sp. Costa Rica [127]
Uromyces inflatus (Cooke) McKenzie Anisotome sp. New Zealand [591]

J. Fungi 2022,8, 633 42 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces ingicola Henn. Inga sp. Amazonas [575]
Uromyces ingiphilus Speg. Inga edulis Argentina [592]
Uromyces insignis P. Syd. & Syd. Echinocephalum latifolium and Melanthera latifolia Brazil [102,219]
Uromyces insularis Arthur Clitoria cajanifolia Puerto Rico [466]
Uromyces intricatus Cooke Chorizanthe sp., Eriogonum spp., and
Gayophytum ramosissimum
Arizona, Canada, California, Idaho,
Oregon,
Montana, and Washington (USA) [185,187,300,412,448]
Uromyces invisus (Speg.) Speg. Solanum sisymbriifolium Argentina [415,592]
Uromyces ipatingae F.A. Ferreira & Y. Hirats. Clitoria fairchildiana Brazil [219,593]
Uromyces iresines Lagerh. Iresine spp. Argentina, Colombia, Dominican Republic,
Ecuador, Guatemala [118,119,144,167,281]
Uromyces isachnes Petch Isathne kunthiana Sri Lanka [594]
Uromyces itoanus Hirats. f. Kummerowia stipulacea, K. striata, and
Microlespedeza spp. China, Korea, Japan, and Taiwan [89,90,483,595,596]
Uromyces ixiae (Lév.) G. Winter
Acidanthera spp., Antholyzasp., Babiana spp.,
Engysiphon sp., Geissorhiza sp., Gladiolus spp.,
Hesperantha spp., Ixia spp., Lapeirousia spp.,
Melasphaerula spp., Romulea sp., and Sparaxis spp.
Kenya and South Africa [123,140]
Uromyces jacksonii Arthur & Fromme Agrostis spp., Deschampsia spp., Festuca spp.,
and Hordeum spp. California, Idaho, Oregon, Michigan, and
Washington (USA) [241,559]
Uromyces jamaicensis Vestergr. Bauhinia spp. Cuba, El Salvador, Jamaica, México,
Puerto Rico, and Venezuela [184,216,292,306,308,444,541]
Uromyces janiphae (G. Winter) Arthur Jatropha spp. and Manihot sp.
Brazil, Colombia, Costa Rica, Cuba,
Ecuador,
Panama, Puerto Rico, México, and
Venezuela
[76,118,119,125,431,456,511,597]
Uromyces japonicus Berk. & M.A. Curtis Orchidaceae Bulgaria, China, Italy, Japan, Poland,
Russia, and Ukraine [51,82,96,135,160,598,599]
Uromyces jatrophae Dietel & Holw. Jatropha multifida Brazil, Colombia, Cuba, México, Jamaica,
Puerto Rico, and Virgin Islands [102,103,184,227,308,479,597]
Uromyces jatrophicola Henn. Cnidoscolus sp. and Jatropha sp. Brazil [103,600]
Uromyces joffrinii Delacr. Vanilla planifolia France [476]

J. Fungi 2022,8, 633 43 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces johowii Dietel & Neger Vicia macraei,V. nigricans, and Vicia sp. Chile [282,494]
Uromyces jonesii Peck Ranunculus sp. California, Colorado, Montana, and
Wyoming [187,300,368,412,601]
Uromyces jordianus BubákEuphorbia cyparissias
Austria, Bulgaria, Czech Republic,
Germany,
Hungary, Romania, Russia, Switzerland,
and Ukraine
[82,220,602]
Uromyces junci Tul.
Ambrosia psilostachya,Arnica spp., Aster spp.,
Bahia dissecta,Cirsium spp., Eriophyllum sp.,
Grindelia sp., Helianthus spp., Jaumea spp.,
Juncus spp., Juniperus litoralis,Luzula alopecurus,
and Pulicaria dysenterica
Arizona, Bolivia, Bulgaria, Canada,
California, Chile, Colorado, Florida,
Germany, Idaho,
Israel, Japan, North and South Dakota,
Nebraska, Montana, Oklahoma, Oregon,
Pennsylvania, Poland, Romania, Spain,
Wyoming, and Wisconsin (USA)
[81,82,105,116,135,185,187,262,377,
386,559,603]
Uromyces juncicola Speg. Juncus stipulatus Mendoza [180]
Uromyces junci-effusi P. Syd. & Syd. Juncus spp.
Arizona, California, Canada, Idaho, Florida,
Georgia, Maine, Missouri, Montana,
Oregon,
Utah, Washington, and Wyoming
[114,116,144,185,187,191,202,262,
559]
Uromyces juncinus Thüm. Junco acutiflori Italy [604]
Uromyces kaernbachii Henn. Abrus precatorius Papua New Guinea [136]
Uromyces kaimontanus Hirats. f. & S. Sato Veratrum album,V. grandiflorum,V. nigrum,
and V. puberulum China, Japan [222,605]
Uromyces kalmusii Sacc. Euphorbia sp. China, Germany, Poland, and Russia [105,135,606,607]
Uromyces karjaginii Uljan. Seseli cuneifolium Azerbaijan [608]
Uromyces kawakamii Syd. & P. Syd. Euphorbia serrulata China, Japan, and Taiwan [94,97,108,446]
Uromyces kentaniensis Doidge Antholyza aethiopica and Chasmanthe aethiopica South Africa [249,322]
Uromyces kenyensis J.F. Hennen Chloris gayana,C. myriostachya, and C. roxburghiana Kenya, Uganda, Zimbabwe [171,242,460]
Uromyces kigesianus Cummins Pittosporum abyssinicum Uganda [609]
Uromyces kisantuensis Henn. Dolichos sp. The Democratic Republic of the Congo [610]
Uromyces klebahnii E. Fisch. Astragalus spp. and Oxytropis sp. Austria, Bulgaria, France, Italy, Romania,
Russia, Switzerland, and Turkey [80,82,220]
Uromyces klotzschianus B. Ali Rumex dentatus Pakistan [61]
Uromyces kochiae Syd. & P. Syd. Kochia prostrata China, Russia, Turkey, and Uzbekistan [144,146,198,402]

J. Fungi 2022,8, 633 44 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces kochianus Gäum. Geranium nodosum Switzerland [611]
Uromyces koeleriae Uljan. Koeleria caucasica Russia [106]
Uromyces komarowii BubákSolidaginis virgaureae Czech Republic [612]
Uromyces kondoi Miura Amblytropis multiflora China, Japan, and Pakistan [89,156,356]
Uromyces krameriae Arthur Krameria spp. Florida and Texas [262,281]
Uromyces krantzbergensis Doidge Anthericum sp. Namibia [138]
Uromyces kunigamiensis Shimab. Fimbristylis dichotoma and Fimbristylis sp. China and Japan [108,222]
Uromyces kurtzii Henn. Senecio spp. Argentina [415,613]
Uromyces kwangensis Henn. Justicia sp. Congo [218,610]
Uromyces kwangsianus Cummins Fimbristylis sp. China and Taiwan [614,615]
Uromyces lachenaliae Doidge Lachenalia spp. and Polyxena ensifolia South Africa [123,138]
Uromyces laevigatus Syd. & P. Syd. Aneilema spp. Ghana and Tanzania [318,549,616]
Uromyces laevis Körn. Euphorbia spp. Bulgaria, Germany, Iran, Romania, Turkey,
and Ukraine [80,82,105,111,228,617,618]
Uromyces langtangensis Durrieu Anaphalis nepalensis Nepal [463]
Uromyces lapeyrousiae Petr. Lapeyrousia Tanzania [231]
Uromyces lapponicus Lagerh. Astragalus spp. and Oxytropis spp.
Alaska, Canada, China, Colorado, Finland,
Idaho, India, Iran, Japan, Mongolia,
Norway,
Oregon, Pakistan, Russia, Sweden, and
Turkey
[80,96,128–
131,357,368,436,449,535,619]
Uromyces largus Arthur & Cummins Chamaesyce lata Colorado [87,88]
Uromyces laserpitii-graminis E. Fisch. Laserpitii sileris and Melica ciliata Southern Europe and Northern Africa [106]
Uromyces lasiocorydis Henn. Lasiocorys abyssinica Eritrea [287,426]
Uromyces lathyrinus Speg. Lathyrus spp. and Vicia spp. Argentina, Brazil, Chile, and Paraguay [100–102,219,620]
Uromyces latimammatus J.Y. Zhuang & S.X.
Wei Ipomoea sumatrana China [621]
Uromyces lazistanicus Petr. Orobus roseus Armenia and Turkey [220]
Uromyces lenticola Petr. Lens esculenta Iran [430]
Uromyces leonotidis Bagyan., Gjaerum & M.
Raju Leonotis nepetifolia India [622]

J. Fungi 2022,8, 633 45 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces leptaleus Syd. Stellaria laxa China, Japan, Philippines, and Taiwan [89,90,97,565,623]
Uromyces leptochloae Wakef. Leptochloa obtusiflora Kenya and Uganda [140,168]
Uromyces lereddei Dupias Colutea arborescens France [220,624]
Uromyces lespedezae (Thüm.) Peck Lespedeza capitata Vermont [583]
Uromyces lespedezae-bicoloris F.L. Tai & C.C.
Cheo Lespedeza bicolor and L. formosa China [108,109,625]
Uromyces lespedezae-macrocarpae Liou & Y.C.
Wang Campylotropis macrocarpa,Lespedeza bicolor,
and L. formosa China [108,109]
Uromyces lespedezae-procumbentis
(Schwein.) Lagerh. Campylotropis spp., Kummerowia spp.,
and Lespedeza spp. Canada, China, Georgia, India, Japan,
Korea, Missouri, Taiwan, and Pennsylvania [94,108,109,185,202,255,357,482,
596]
Uromyces lespedezae-sericeae S. Ahmad Lespedeza sericea Pakistan [208]
Uromyces libycus Trotter Lotus pusillus Libya [197,626]
Uromyces ligulariae Hirats. & Hoshioka Ligularia tussilaginea var. formosana China, Japan, Taiwan [94,97,109,627]
Uromyces limonii-caroliniani Savile &
Conners Limonium carolinianum Canada, Mississippi, and Texas [185,559,628]
Uromyces limosellae F. Ludw. Limosella sp. Australia [280]
Uromyces loculatus Cummins Kyllinga alba Zambia [341]
Uromyces loculiformis T.S. Ramakr. & K.
Ramakr. Chlorophytum attenuatum China, India, and Nepal [96,464,629]
Uromyces lomandracearum J. Walker & van
der Merwe Lomandra longifolia and Lomandra sp. Australia [62,630]
Uromyces longipedicellaris Ramachar & A.
Sudh. Rao Rumex vesicarius Pakistan [631]
Uromyces longipes (Lasch) Tranzschel Pedicularis sp. Russia [382]
Uromyces loranthi H.S. Jacks. Loranthus Brazil [376]
Uromyces lotononidicola Berndt Lotononis cytisoides South Africa [632]
Uromyces lugubris Kalchbr. Euphorbia cyparissias South Africa [137]
Uromyces lupini Berk. & M.A. Curtis Lupinus spp. Arizona, California, Canada, Colorado,
México, Montana, Oregon, Washington,
and Wyoming [116,185,187,368,412,454,598]

J. Fungi 2022,8, 633 46 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces lupinicola BubákLupinus spp. Bulgaria, Finland, Lithuania, Norway,
and Spain [158,363,587,612]
Uromyces lychnidis Tracy & Earle Lychnidis drummondii Utah [633]
Uromyces lygei P. Syd. & Syd. Lygeum spartum Sardegna [325]
Uromyces macnabbii Cummins Chionochloa spp. and Danthonia spp. New Zealand [284,591]
Uromyces macounianus Ellis & Everh. Euphorbia maculata var. affinis and E. serpyllifolia Canada and British Columbia [103,388,634]
Uromyces macowanii BubákScilla prasina Africa [612]
Uromyces magnatus (Arthur) Arthur Polygonatum sp. and Vagnera sp. Illinois, Iowa, Minnesota, Montana,
Nebraska, North and South Dakota, and
Wisconsin [306,307]
Uromyces magnusii Kleb. Medicago minima China, Turkey, and Ukraine [80,96]
Uromyces maireanus P. Syd. & Syd. Ornithogalum spp. Morocco, South Africa, Tanzania [142,144,318]
Uromyces major Arthur Muhlenbergia reverchonii,Muhlenbergia sp.,
and Sporobolus indicus Belize, México, Texas, Trinidad, and Tobago [112,117,635]
Uromyces malloti Henn. Mallotus moluccanus and Melanolepismulti glandulosa Guinea, New Papua, and the Philippines [136,177]
Uromyces mangenotii Mayor & Vienn.
Bourg. Vicia pubescens France [516]
Uromyces manihoticola Henn. Manihot spp. Brazil [102,103,291]
Uromyces manihotis Henn. Manihot spp. Brazil, Colombia, Cuba [102,227,291]
Uromyces manihotis-catingae Henn. Manihot spp. Brazil [102,600]
Uromyces marinus Guyot & Malençon Medicago marina Morocco [220,279]
Uromyces martinii Farl. Melanthera spp., Bidens spp. Florida [225,262,636]
Uromyces massoniae Doidge Massonia latifolia South Africa [138]
Uromyces mayorii Tranzschel Euphorbia spp. California and Colombia [227,412]
Uromyces mbelensis Henn. Indigofera The Democratic Republic of the Congo [610]
Uromyces megalosporus Speg. Tessaria absinthioides Tucumán[371]
Uromyces meimerlianus Magnus Vicia sp. Germany [531]
Uromyces melandrii Dietel & Neger Melandrium cucubaloides Los Lagos [162]
Uromyces melantherae Cooke Melanthera brownii Ghana, South Africa, and Uganda [138,168,616]

J. Fungi 2022,8, 633 47 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces melasphaerulae Syd. & P. Syd. Melasphaerula graminea Western Cape Province [424]
Uromyces melothriae Henn. Melothria tomentosa Eritrea [287,427]
Uromyces menyanthis Azbukina & Zenkova Menyanthes trifoliata Primorye 1966 [637]
Uromyces mercurialis Henn. Mercurialis leiocarpa China, Japan, and Taiwan [94,96,97]
Uromyces mexicanus Dietel & Holw. Desmodium Arizona, Costa Rica, México [116,171,310,479]
Uromyces meygounensis Petr. Euphorbia bungei Iran [638]
Uromyces microchloae Syd. & P. Syd. Microchloa setacea Bolivia, Brazil, Central Africa, Sudan, and
Uganda [106,168,199,394]
Uromyces microsorus Kalchbr. & Cooke Prunus armeniaca South Africa [137]
Uromyces microtidis Cooke Microtis porrifolia Australia and New Zealand [245,313,639]
Uromyces miersiae Gäum. Miersia chilensis Chile [540]
Uromyces mikaniae Viégas Mikania sp. Brazil [219,640]
Uromyces mimusops Cooke Mimusops sp. South Africa [641]
Uromyces minimus Davis Muhlenbergia sylvatica Canada, Michigan, Oregon, and Wisconsin [185,635]
Uromyces minor J. Schröt. Medicago denticulata,Pisum sativum,
and Trifolium spp.
Armenia, Austria, Bulgaria, California,
Caucasus, Chile, China, Czech Republic,
Estonia, Finland, France, Germany,
Hungary, India,
Montana, Pakistan, Poland, Romania,
Russia, Spain, Sweden, Switzerland, Texas,
Ukraine,
New Zealand, Oregon, Washington,
and Wyoming
[220,313,355,357,559,642]
Uromyces minutus Dietel Carex spp. Alabama, Central and Eastern United
States,
Iowa, Mississippi, Texas, and Wisconsin [255,475,559,643,644]
Uromyces miurae Syd. & P. Syd. Fritillaria kamtschatcensis Alaska, Canada, Idaho, Japan, Russia, and
Washington (USA) [94,185,187,645,646]
Uromyces moehringiae S. Ito & Hirats. f. Moehringia lateriflora Japan [94,297]
Uromyces moesiacus Lindtner & Vienn.
Bourg. Lathyrus nissolia Yugoslavia [451]
Uromyces mogianensis BubákFritillaria graeca,F. rhodocanakis,
Rhinopetalum arianum,R. gibbosum, and R. karelinii Afghanistan, Greece, and Uzbekistan [421,555,612]

J. Fungi 2022,8, 633 48 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces mongolicus U. Braun & G. Hirsch Euphorbia kozlovii Mongolia [535,647]
Uromyces monspessulanus Tranzschel Euphorbia serrata Balearic Islands, Libya, and Spain [129,157,197,648]
Uromyces montanoae Arthur & Holw. Montanoa dumicola,M. pittieris,M. hibiscifolia,
and Montanoa sp. Costa Rica and Guatemala [225,281,310]
Uromyces montanus Arthur Lupinus mexicanus Guatemala and México [281,398]
Uromyces montis-ferrati Maire Euphorbia luteola Northern Africa [649]
Uromyces moraeae Syd. & P. Syd. Moraea spathacea South Africa [138,144]
Uromyces mucunae Rabenh. Mucuna pruriens
China, Cambodia, Japan, India, Indonesia,
Madagascar, Malawi, Mauritius, Taiwan,
Tanzania, South Africa, Guinea,
Philippines,
Thailand, and Uganda
[97,209,273,309,465,565,605,650–
652]
Uromyces muhlenbergiae S. Ito Muhlenbergia japonica Japan [635]
Uromyces mulgedii Lindr. Lactuca sp. The Czech Republic and Turkey [569]
Uromyces mulini J. Schröt. Mulinum integrifolium Argentina and Chile [240,331,415]
Uromyces musae Henn. Musa spp. Congo, Fiji, and Philippines [565,610,653,654]
Uromyces muscari (Duby) Niess
Bellevalia spp., Chionodoxa spp.,
Dipcadi erythraeum,Endymionspp., Gagea sp.,
Hyacinthoides spp., Leopoldia spp., Muscari spp.,
Ornithogalum sp., Scilla spp., and Urginea spp.
Afghanistan, Bulgaria, California, Canada,
Germany, Greece, India, Israel, Iran, Italy,
Morocco, New Zealand, Norway, Scotland,
Sweden, and Turkey
[80–82,105,123,124,179,317,318,389,
534,655]
Uromyces mussooriensis Syd. & P. Syd. Stipa sibirica India [106,656]
Uromyces myosotidis Bahç. Myosotis sp. Turkey [471]
Uromyces myristica Berk. & M.A. Curtis Euphorbia bicolor Texas [657]
Uromyces myrsines Dietel Ardisia compressa,Icacorea sp., Myrsine spp.,
and Rapanea spp. Bolivia, Brazil, Costa Rica, and Uruguay [100,101,143,310]
Uromyces mysticus Arthur Hordeum jubatum California, Canada, Colorado, Idaho, Israel,
and Utah [81,112,114,185,412,559]
Uromyces namaqualandus
Mennicken, W. Maier & Oberw. Roepera cordifolia Namibia [384]
Uromyces nassauviae J.C. Lindq. Nassauvia lagascae Argentina [415]
Uromyces nassellae Cummins Nassella pubiflora Bolivia [106]

J. Fungi 2022,8, 633 49 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces natalensis Magnus Euphorbia spp. Madagascar and South Africa [169,465]
Uromyces natricis A.L. Guyot Ononis rotundifolia France [342]
Uromyces nattrassii Cummins Statice spicata Cyprus [350]
Uromyces naucinus Berndt Cayaponia sp. Ecuador [127]
Uromyces necopinus Cummins Hypoxis hirsuta Connecticut, Massachusetts, and New York [167,233]
Uromyces neotropicalis J.R. Hern. & Aime Cayaponia rigida,C. selysioides, and Cucurbita sp. Guiana and Guyana [63,557]
Uromyces nevadensis Harkn. Primula suffrutescens California [78,658]
Uromyces nidificans Tranzschel Aellenia subaphylla,Cimacoptera korshinskyi,
Salsola arbuscula, and Salsola sp. Central Asia, Iran, and Uzbekistan [146,161,346]
Uromyces niteroyensis Rangel Panicum spp. and Setaria spp. Argentina, Brazil, Colombia, India,
French Guiana, and México [102,110,395,415,659]
Uromyces nordenskjoldii Dietel Vicia darapskyana and Vicia sp. Argentina and Chile [415]
Uromyces notabilis Wakef. Cyperus sp. and Kyllinga sp. Uganda [168,462]
Uromyces nothoscordi Syd. & P. Syd. Nothoscordum striatum Texas [660]
Uromyces novissimus Speg. Trianosperma ficifolia Argentina, Brazil, Colombia, Panama,
México,
and West Indies [48,324,469]
Uromyces numidicus Maire Geranium atlanticum Northern Africa [649]
Uromyces nyikensis Syd. & P. Syd. Gladiolus nyikensis Malawi [424]
Uromyces nymphoidis Săvul. Nymphoides peltata Romania [228]
Uromyces oaxacanus Dietel & Holw. Jatropha urens Arizona, Belize, Guatemala, and México [103,116,281]
Uromyces oberwinklerianusBerndt Acalypha sp. Costa Rica [310]
Uromyces obesus Durrieu Heteropogon contortus Nepal and Uganda [244,464]
Uromyces oblectaneus H.S. Jacks. & Holw. Rhynchospora corymbosa,R. exaltata, and
Rhynchospora sp. Brazil [166,219]
Uromyces oblongisporus Ellis & Everh. Artemisia tridentata China and Wyoming [225,661]
Uromyces oblongus Vize Medicago polymorpha and Trifolium spp. California and Washington [78,454,662]
Uromyces obscurus Dietel & Holw. Phaseolus sp. México [456,479]

J. Fungi 2022,8, 633 50 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces occidentalis Dietel Euphorbia spp., Lupinus spp., and Tithymalus spp. Arizona, California, Colorado, México,
Montana, Nevada, Nigeria, Oregon, and
Utah [114,116,187,292,316,412,663]
Uromyces occultus J.C. Lindq. Juncus densiflorus var. pohlii Argentina, Brazil, Chile, and México [102,486,664]
Uromyces ocimi Hansf. Ocimum menthifolium Uganda [168]
Uromyces oedipus Dietel Sophora japonica Japan [665]
Uromyces oenotherae Burrill Oenothera linifolia Illinois [582]
Uromyces oliveirae J. Anikster & I. Wahl Bellevalia eigii Israel [390]
Uromyces ononidis Pass. Euphorbia spp. and Ononis spp.
Bulgaria, Cyprus, Greece, Israel, Italy,
Libya,
Poland, Morocco, Romania, Spain, and
Turkey
[80–82,105,111,220,532,666,667]
Uromyces ophiorrhizae Gäum. Ophiorrhiza longiflora Indonesia [309]
Uromyces orbicularis Dietel Desmodium spp. Argentina, Brazil, and Bolivia [143,171,269]
Uromyces orchidearum Cooke & Massee Chiloglottis spp. Australia [268]
Uromyces orientalis Syd. & P. Syd. Indigofera spp. Australia, Ethiopia, Ghana, India, Japan,
Kenya, Pakistan, Malawi, Philippines,
Tanzania, Uganda, and Zambia [85,168,171,172,357]
Uromyces ornatipes Arthur Phrygilanthus sonorae México [15,76,77]
Uromyces ornithogali (Wallr.) Niessl Gagea arvensis,Ornithopus nanum, and Ornithopus sp. Romania, Portugal, Spain, and Ukraine [111,158,179,228]
Uromyces ornithopodioides Gonz. Frag Ornithopus isthmocarpus and O. compressus Portugal [317]
Uromyces orobi-tuberosi (Pers.) Liro Gladiolus sp. Finland [668]
Uromyces orthosiphonis T.S. Ramakr. &
Sriniv. Orthosiphon glabratus India [152,669]
Uromyces otakou G. Cunn. Poa spp. New Zealand [591,670]
Uromyces otaviensis Mennicken, W. Maier &
Oberw. Ipomoea verbascoidea Namibia [385]
Uromyces ovalis Dietel Leersia oryzoides Japan [671]
Uromyces ovirensis Jaap Primula wulfeniana Austria [672]
Uromyces pachyceps Lagerh. Ipomoea sp. Brazil, Ecuador [126]
Uromyces pallidus Niessl Chamaecytisus spp., Cytisus spp., and
Lembotropis nigricans
Austria, Belarus, Bulgaria, Czech Republic,
Germany, Hungary, Italy, Poland, Romania,
Ukraine, and Yugoslavia [105,135,220,673]

J. Fungi 2022,8, 633 51 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces panici-sanguinalis Rangel Digitaria horizontalis,Panicum sanguinale, and
Panicum sp. Brazil, Cuba, Grenada, and Venezuela [171,674,675]
Uromyces pannosus Vestergr. Bauhinia candicans Brazil [216]
Uromyces papillatus Kalchbr. & Cooke Heteromorpha arborescens South Africa [137]
Uromyces paradoxus Syd. & P. Syd. Commiphora zimmermannii and Commiphora sp. Kenya and Mozambique [138,140,676]
Uromyces parilis Syd. Rumex occultans Israel [81]
Uromyces paspalicola Arthur & Holw. Paspalum racemosum Ecuador [199,294]
Uromyces paulshoekensis
Mennicken, W. Maier & Oberw. Roepera foetida South Africa and Northern Cape Province [384]
Uromyces pavgii R.N. Goswami & Ngachan Achyranthes aspera India [677]
Uromyces pavoniae Arthur Pavonia racemosa Puerto Rico [259,678]
Uromyces pazschkeanus Henn. Vigna sp. Eritrea [287,427]
Uromyces peckianus Farl. Aristida spp., Atriplex spp., Bryzopyrum sp.,
Chenopodium spp., Distichlis spp., Houstonia spp.,
Plantago spp., and Salicornia spp.
Alabama, California, Canada, Florida,
Oklahoma, Massachusetts, Missouri,
Montana, New York, Texas, and Wisconsin
(USA)
[185,187,262,455,475,559,636]
Uromyces peglerae Pole-Evans Digitaria spp.
Kenya, India, Malavi, Mauritius, New
Guinea,
Pakistan, Philippines, South Africa,
Uganda,
and Zimbabwe
[171,172,232,441,447]
Uromyces peireskiae Dietel Peireskia grandifolia,P. sacharosa, and Peireskia sp. Argentina and Brazil [394,415]
Uromyces pencanus
(Dietel & Neger) Arthur & Holw. Nassella spp. and Stipa spp. Argentina, Australia, Bovilia, Chile,
andNew Zealand [24,199,415]
Uromyces penniseti S. Ahmad Pennisetum lanatum Pakistan [536,679]
Uromyces pentaceae D.K. Agarwal Pentace burmanica India [680]
Uromyces pentaschistidis Gjaerum Pentaschistis airoides South Africa [243]
Uromyces peracarpae S. Ito & Tochinai Peracarpa carnosa and P. circaeoides Japan and Russia [94,297]
Uromyces peraffinis Dietel Bauhinia sp. Brazil [219]
Uromyces pereskiae H.S. Jacks. & Holw. Pereskia aculeata,P. grandifolia, and Pereskia sp. Argentina and Brazil [269,681]

J. Fungi 2022,8, 633 52 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces perigynius Halst. Carex spp., Ratibida columnaris,Rudbeckia spp., and
Solidago spp.
Bermuda, Canada, Idaho, Iowa, Maine,
Missouri, Montana, Oregon, Pennsylvania,
Washington, and Wisconsin [185,187,191,252,260]
Uromyces perlebiae Vestergr. Bauhinia spp. Brazil [102,216]
Uromyces permeritus Cummins Tournefortia sarmentosa and Tournefortia sp. Papua New Guinea [223,426]
Uromyces persicus Syd. & P. Syd. Astragalus spp., Oxytropis spp., and
Phaca spp.
Alaska, Alberta, Austria, Canada,
Colorado,
India, Iran, Japan, Kazakhstan, Norway,
Oregon, Russia, Siberia, Sweden, Tajikistan,
and Turkmenistan
[220,357,682]
Uromyces petitmenginii Maire Minuartia globulosa and M. mereyi Greece, Turkey [80,421,470]
Uromyces phacae-frigidae (Wahlenb.) Har. Astragalus spp. and Phaca spp. Alaska, Canada, Caucasus, Kyrgyzstan,
Pakistan, Norway, Russia, and Sweden [367,436,553,646]
Uromyces phalaridicola Katajev Phalaris minor Turkmenistan [106]
Uromyces phaseolicola Speg. Phaseolus prostratus Argentina [126,180]
Uromyces phlei-michelii Cruchet Phleum alpinum,P. phleoides, and
P. michelii Morocco and Switzerland [126,279]
Uromyces phlogacanthi Gäum. Phlogacanthus celebicus Indonesia [309,683]
Uromyces phtirusae Mayor Phthirusa pyrifolia Colombia [15,226]
Uromyces phyllachoroides Henn. Cynosurus elegans Tunisia [684]
Uromyces physanthyllidis Vienn.-Bourg. Physanthyllis tetraphylla Greece [544]
Uromyces phyteumatum (DC.) Niessl Phyteuma spp. The Czech Republic, Denmark, France,
Germany, Norway, Poland, Romania, and
Spain [128,158,159,228,440]
Uromyces pianhyensis Henn. Wedelia spp. Brazil, Ethiopia, India, Puerto Rico,
Virgin Islands, and West Indies [264,312,571,600]
Uromyces pictus Thüm. Abutilon elaeocarpoides and Abutilon sp. Galapagos Islands and Ethiopia [571,685,686]
Uromyces pieningii Cummins Ipomoea argentaurata and Ipomoea pes-caprae Ghana and Indonesia [126,341]

J. Fungi 2022,8, 633 53 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces pisi-sativi (Pers.) Liro
Astragalus spp., Colutea sp., Cytisus scoparius,
Euphorbia sp., Galega officinalis,Genista spp.,
Lathyrus spp., Lotus spp., Medicago spp.,
Onobrychis spp., Pisum spp., Trifolium pratense,
Ulex europaeus, and Vicia sp.
Finland, Germany, Hong Kong, Israel,
Libya,
Turkey, Serbia, and Uzbekistan [80,81,329]
Uromyces pittospori Henn. Pittosporum abyssinicum Eritrea [287,687]
Uromyces planiusculus (Mont.) Jørst. Rumex frutescens Tristan da Cunha [688]
Uromyces plantaginis Vestergr. Plantago barbata and P. tubulosa Argentina [165,415]
Uromyces plumbarius Peck Gaura spp., Oenothera spp., Onagra biennis,
and Pachylophus marginatus
California, Canada, Colorado, Connecticut,
Florida, Idaho, Louisiana, Iowa, México,
Minnesota,
Mississippi, Oregon, Texas, and Utah
[114,155,185,187,262,475,559]
Uromyces poae-alpinae Rytz Poa alpina and Ranunculus montanus Germany and Poland [135,160]
Uromyces poinsettiae Speg. Poinsettia heterophylla Argentina [689]
Uromyces poiretiae Syd. Poiretia scandens Venezuela [431,690]
Uromyces polemanniae Kalchbr. & Cooke Polemannia spp. South Africa [137,142]
Uromyces poliotelis Syd. Anguria sp., Gurania sp., and
Selysia prunifera Costa Rica [127,229]
Uromyces politus (Berk.) McAlpine Muehlenbeckia cunninghamii Australia [280,343]
Uromyces polycnemi McAlpine Polycnemum pentandrum Bulgaria, Britain, Iran, and Ukraine [82,280,317,422]
Uromyces polygalae Grove Polygala spp. Mongolia and Uganda [535,691]
Uromyces polygoni-avicularis (Pers.) G.H.
Otth Polygonum nepalense Nepal [692,693]
Uromyces polymniae (Henn.) Dietel & Holw. Polymnia spp. Argentina, Brazil, Colombia, Guatemala,
and México [219,269,281,375]
Uromyces polymorphus Peck & Clinton Lathyri sp. New York [694]
Uromyces polytriadicola Arthur & Cummins Polytrias amaura Philippines [232]
Uromyces pontederiae W.R. Gerard Pontederia cordata
Argentina, Brazil, Delaware, Florida,
Georgia,
Missouri, New York, Pennsylvania, Texas,
and Virginia
[415,559,695]

J. Fungi 2022,8, 633 54 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces pontederiicola Speg. Pontederia sagittata Argentina and India [34,592]
Uromyces poonensis W.D. More & Moniz Sesbania aegyptiaca,S. grandiflora, and
S. sesban India [357,696]
Uromyces porcensis Mayor Inga ingoides Colombia [226]
Uromyces porosus (Peck) H.S. Jacks. Vicia americana and V. sparsifolia Iowa [155,295,454]
Uromyces pozoae Dietel & Neger Pozoa hydrocotylifolia Chile [240,417]
Uromyces praetextus Vestergr. Bauhinia sp. Brazil [216]
Uromyces prangi Har. Hippomarathrum cristatum and Prangos sp. Bulgaria and Iran [82,347]
Uromyces pratensis Juel Poa pratensis,Ranunculus auricomus, and R.
cassubicus Finland [329]
Uromyces pratiae Speg. Hypsela reniformis and Pratia repens Argentina, Brazil, and Ecuador [123,219,415]
Uromyces pretoriensis Doidge Commelina africana Ghana, Uganda, Namibia, and South Africa [138,168,443]
Uromyces primaverilis Speg. Allium striatellum Argentina, Illinois, Kansas, Missouri,
Michigan, Missouri, Texas, Oklahoma, and
Uruguay [305,415,455,542,559]
Uromyces primulae-integrifoliae (DC.) Niessl Primula deorum and P. integrifolia Bulgaria and Switzerland [82,159]
Uromyces prismaticus Vienn. Bourg. Secale montanum Iran [320]
Uromyces privae Syd. & P. Syd. Priva lappulacea Cuba and Venezuela [697,698]
Uromyces probus Arthur Olsynium grandiflorum and
Sisyrinchium spp. Canada, California, Idaho, Oregon, Texas,
Utah, and Washington [66,112,185,187,559]
Uromyces procerus J.C. Lindq. Festuca procera Chile [240]
Uromyces propinquus P. Syd. & Syd. Desmodium sp. and Rhopalotria mollis Mexico [325]
Uromyces prosopidis (Jacz.) Jacz. Prosopis farcta Iran [699]
Uromyces pseudarthriae Cooke Pseudarthria robusta South Africa [700]
Uromyces psoraleae Peck Psoralea lanceolata
Arizona, Canada, Colorado, Idaho,
Montana,
Oregon, South Africa, Utah, Washington,
and Wyoming
[142,185,187,300,368,559,701]
Uromyces psychotriae Henn. Psychotria sp. Brazil [219,575]
Uromyces pteroclaenae Lindr. Cachryde sp. Algeria [501]

J. Fungi 2022,8, 633 55 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces pulchellus Ellis & Everh. Silene douglasii and Silene sp. California and Washington (USA) [78,388]
Uromyces pulvinatus Kalchbr. & Cooke Euphorbia inaequilatera South Africa [702]
Uromyces punctiformis Syd. & P. Syd. Vigna strobiliphora Mexico [703]
Uromyces purpureus Lagerh. Liliaceae Angola [704]
Uromyces pustulatus Wakef. Bauhinia fassoglensis Uganda and Kenya [140,168]
Uromyces puttemansii Rangel Panicum spp. and Setaria spp. Argentina, Brazil, Cuba, Honduras,
Jamaica, México, Panama, Uruguay, and
Venezuela [126,171,674]
Uromyces pyriformis Cooke Acorus calamus Iowa, Illinois, Japan, Massachusetts, Maine,
Minnesota, Mississippi, New York, and
Taiwan [92,155,574,596]
Uromyces quaggafonteinus Mennicken &
Oberw. Ehrharta calycina South Africa [378,429]
Uromyces quinchamalii Neger Quinchamalium spp. Argentina, Bolivia, and Chile [415,705]
Uromyces ramacharii Ravinder & Bagyan. Ocimum sp. India [706]
Uromyces ranunculi-distichophylli Semadeni Ficaria sp. and Ranunculus sp. Africa, America, China, Canada, Europe,
Iran,
Japan, and Russia [106]
Uromyces rapaneae Henn. Rapanea sp. São Paulo [450]
Uromyces ratoides Jørst. Cayaponia spp. Ecuador [124,212]
Uromyces ratus H.S. Jacks. & Holw. Cayaponia spp. Brazil [210,212]
Uromyces rayssiae J. Anikster & I. Wahl Scilla hyacinthoides Israel [390]
Uromyces rebeccae
Bruckart, M. Abbasi
& Aime Suaeda californica California [47]
Uromyces regius Vestergr. Bauhinia candicans Brazil [216,219]
Uromyces reichei Dietel Milla bivalvis and Triteleia gaudichaudiana Chile [240,415]
Uromyces reichertii J. Anikster & I. Wahl Scilla hyacinthoides and Hordeum bulbosum Israel [81]
Uromyces renovatus P. Syd. & Syd. Lupinus sp. Czech Republic, Greece, Israel, Kenya,
Finland, Portugal, and Spain [81,131,140,144,158,707]

J. Fungi 2022,8, 633 56 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces reticulatus (Thüm.) BubákAllium victorialis Portugal and Spain [158,612]
Uromyces reynoldsii Thaung Modecca bracteata and Trichosanthes spp. Myanmar [212]
Uromyces rhinacanthi Cummins Rhinacanthus nasutus Ghana, West Africa [218,609]
Uromyces rhodesicus Wakef. Bauhinia galpinii and B. macrantha South Africa and Zimbabwe [138,250]
Uromyces rhynchosporae Ellis Rhynchospora spp.
Brazil, Bermuda, California, Cuba, Florida,
Georgia, Hawaii, Louisiana, Japan,
Massachusetts, Michigan, New Jersey,
Puerto Rico, West Indies, and Vermont
[94,166,191,202,306,307,388,412,
559]
Uromyces ribicola H.S. Jacks. & Holw. Ribes albifolium and R. andicola Bolivia, Brazil, and Colombia [101,126,227]
Uromyces rickerianus Arthur Polygonum spp. and Rumex spp. Colorado, Idaho, Utah, and Wyoming [187,368]
Uromyces riloensis Hinkova Doronicum cordifolium Bulgaria [82]
Uromyces rostratus Henn. Eriosema sp. Rio de Janeiro [331]
Uromyces rottboelliae Arthur Rottboellia compressa,R. exaltata, and R. speciosa Congo, India, and Philippines [171,708]
Uromyces rubidus Arthur & Holw. Andropogon condensatus Brazil [199]
Uromyces rudbeckiae Arthur & Holw. Rudbeckia spp. and Solidago spp.
Canada, China, Idaho, Florida, Japan,
Korea, Iowa, Missouri, Mississippi,
Montana, Taiwan,
Texas, Wyoming
[94,108,109,185,483,559]
Uromyces ruelliae Holw. Beloperone californica,Beloperone sp.,
Justicia brandegeana, and Ruellia sp. Arizona, California, Florida, México,
and Nevada [218,262,409,412]
Uromyces rugosus Arthur Lupinus sp. México [398]
Uromyces rugulosus Pat. Campylotropis spp. and Lespedeza spp. China [96]
Uromyces ruiz-leali J.C. Lindq. Anarthrophyllum elegans Argentina [415]
Uromyces rumicis (Schumach.) G. Winter Emex australis,Ficaria spp., Medicago spp.,
Ranunculus ficaria, and Rumex spp.
Argentina, Armenia, Australia, Brazil,
Bulgaria, Chile, Denmark, Finland,
Germany, Greece, India, Iran, Italy, Japan,
Malawi,
New Zealand, Norway, Pakistan, Poland,
Portugal, Sicily, South Africa, Spain,
Romania, Sweden, Tanzania, Turkey,
Uganda, Ukraine,
Uzbekistan, and Russia
[75,80,82,128,135,142,158,245,279,
313,468,519,534]
Uromyces rumicis (Schumach.) G. Winter Rumicis sp. Australia, Morocco, and Switzerland [709]

J. Fungi 2022,8, 633 57 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces rumicum (DC.) Fuckel Rumicis hydrolapathi Egypt and Minnesota [179,710]
Uromyces rzedowskii J.F. Hennen &
Cummins Ledenbergia macrantha México [292,403]
Uromyces sabineae Arthur Poitea spp. and Sabinea punicea Cuba, Dominican Republic, Puerto Rico [184,486]
Uromyces saginatus Syd. Urginea altissima Namibia, Zimbabwe [138,250]
Uromyces sakawensis Henn. Solidago virgaurea Japan [711]
Uromyces salicorniae (DC.) de Bary Arthrocnemum glaucum and Salicornia spp. China, Germany, Finland, Poland, Portugal,
Romania, and United Kingdom [105,133,135,195,228,329]
Uromyces salmeae Arthur & Holw. Galinsoga sp. and Salmea scandens Costa Rica, Dominican Republic,
Guatemala, Puerto Rico, Virgin Islands,
and West Indies [125,184,225,264,281,282,406]
Uromyces salpichroae H.S. Jacks. & Holw. Salpichroa diffusa and Salpichroa sp. Bolivia and Ecuador [210]
Uromyces salsolae Rabenh. Climacoptera sp., Gamanthus gamocarpus,
Gossypium hirsutum,Halocharis hispida,Noaea spp.,
Petrosimonia spp., and Salsola spp.
Algeria, China, Cyprus, Finland, Israel,
Japan, Mongolia, Morocco, Pakistan,
Russia,
Romania, Turkey, and Ukraine
[80,81,111,146,198,228,329,435,535]
Uromyces sasaensis Gjaerum Valeriana kilimandscharica and V. volkensii Uganda [317,712]
Uromyces satarensis P.B. Chavan & Bakare Blainvillea acmella and B. latifolia China and India [319,713]
Uromyces saulensis Berndt Selysia prunifera France [127]
Uromyces saururi Henn. Saururus chinensis and S. loureiroi China, Japan, and Taiwan [96,97,222]
Uromyces saussureae P. Karst. Saussurea sp. Japan, Siberia [714]
Uromyces savulescui Rayss Limonium sinuatum and Limonium sp. The Canary Islands, Greece, and Israel [81,327]
Uromyces scaberulus L. Guo & Y.C. Wang Lespedeza bicolor,L. cuneate,L. cyrtobotrya,
and L. formosa China [156]
Uromyces scaevolae G. Cunn. Scaevola albida,S. calendulacea,S. spinescens,
and S. radicans Australia and New Zealand [67,313]
Uromyces schanginiae Thüm. Suaeda spp. California and Egypt [710,715]
Uromyces schinzianus Henn. Bauhinia fassoglensis,B. reticulate, and
B. thonningii Somalia, South Africa, and Uganda [138,168]
Uromyces schismi Jørst. Schismus scaberrimus South Africa [411]
Uromyces schoenanthi Syd. & P. Syd. Andropogon schoenanthus,Apluda mutica,Cymbopogon
schoenanthus,Polytrias amaura, and P. diversifolia India, New Guinea, Pakistan, and the
Philippines [152,201,318]

J. Fungi 2022,8, 633 58 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces schweinfurthii Henn. Acacia ehrenbergiana,A. flava, and A. seyal South Yemen and Ethiopia [287,687]
Uromyces scillinus (Durieu & Mont.) Har. Scilla autumnalis Algeria and Europe [451]
Uromyces scirpi-maritimi Hirats. f. & Yoshin. Bolboschoenus maritimus,Glaux maritima,
Scirpus fluviatilis, and S. maritimus Japan and Russia [222,297]
Uromyces scirpinus Syd. Scirpus supinus Philippines [232]
Uromyces scleranthi Rostr. Minuartia spp. and Scleranthus spp. Australia, Bulgaria, Denmark, Finland,
Norway, Sweden, Turkey, and Ukraine [80,107,128,280,329,422]
Uromyces scleriae Henn. Scleria spp.
Barbados, Brazil, Colombia, Cuba,
Dominica,
Grenada, Guyana, Nigeria, Rio de Janeiro,
Saint Lucia, Puerto Rico, Venezuela,
and West Indies
[122,184,219,264,323,431]
Uromyces sclerochloae Tranzschel Sclerochloa dura Central Asia (Turkmenistan and Iran) [161,383]
Uromyces scleropoae Baudyš & Picb. Scleropoa ridiga Croatia [451]
Uromyces scrophulariae (DC.) Fuckel Scrophularia auriculata Carmarthenshire and Wales [716]
Uromyces scutellatus (Schrank) Niessl Euphorbia spp.
Bulgaria, Chile, Germany, Iran, Iraq,
Poland,
Romania, Russia, Serbia, Spain, Turkey,
Ukraine, and Uzbekistan
[43,80,82,105,111,146,179,196,228,
236,347,386]
Uromyces secamones Wakef. Secamone platystigma Uganda [168]
Uromyces sedi Gäum. Sedum anacampseros France [717]
Uromyces seditiosus F. Kern Aristida spp. and Plantago spp.
Alabama, Arkansas, California, Colorado,
Illinois, Indiana, Kansas, Iowa, Missouri,
Nebraska, Nebraska, New York, Oklahoma,
Texas, Wisconsin, and Virginia
[537,559,718]
Uromyces seligeri Tranzschel & Erem. Lathyrus grandiflorus and L. sylvestris Greece and Russia [220,382,421]
Uromyces sellierae G. Cunn. Selliera radicans New Zealand [67]
Uromyces semnanensis Gjaerum Astragalus fridae Iran [276]
Uromyces senecionicola Arthur Cacalia sp., Senecio roldana, and Senecio sp. México [292,398]
Uromyces senecionis-gigantis Gjaerum Senecio gigas Ethiopia [719]
Uromyces senorensis J.F. Hennen &
Cummins Compositae Mexico [703,720]

J. Fungi 2022,8, 633 59 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces sepultus Mains Setaria tenax America, México [721]
Uromyces seseli-graminis E. Fisch. Arrhenatherum elatius,Melica ciliate, and Poae sp. Europe [106]
Uromyces seselis Sousa da Câmara Seseli tortuosum Portugal [722]
Uromyces sesseae Lagerh. Sessea sp. Ecuador [325]
Uromyces setariae-italicae Yoshino Brachiaria spp., Chaetochloa spp., Eriochloa spp.,
Lasiacis spp., Setaria spp., and Urochloa spp. Worldwide [294]
Uromyces shahrudensis Petr. Onobrychis sp. Iran [723]
Uromyces shearianus Arthur Arizona, California, Colorado, México,
Utah,
and Wyoming [114,292,412,559]
Uromyces shikokianus Kusano Cladrastis platycarpa and C. shikokiana Japan [222]
Uromyces sii-latifolii P. Karst. Sium latifolium Britain and Russia [714]
Uromyces silenes (Schltdl.) Fuckel Arenaria glabrescens,Arenaria sp., and Silene spp.
California, Colorado, Iowa, Kansas,
Montana, Pennsylvania, Portugal, Turkey,
Utah,
and Washington (USA)
[80,114,454,559]
Uromyces silenes-chloraefoliae Vienn.-Bourg. Silene chlorifolia Iran [320]
Uromyces silenes-ponticae Const. Silene spp. Bugaria, Iraq, Libya, Romania, Portugal,
and Turkey [80,82,196,228,532]
Uromyces silksvleyensis Mennicken &
Oberw. Bartholina burmanniana Western Cape Province [378]
Uromyces silphii (Syd. & P. Syd.) Arthur Aster macrophyllus,Helianthus spp.,
Heliopsis helianthoides,Juncus spp., and
Silphium spp.
Argentina, Canada, Czech Republic, Chile,
Florida, Georgia, Idaho, Illinois, Iowa,
México, Minnesota, Missouri, New York,
Oklahoma,
Oregon, Pennsylvania, Poland, Texas,
Uruguay, Washington, Wisconsin, and
Vermont
[185,187,191,292,415,559]
Uromyces simulans Peck Vilfa sp. Colorado [724]
Uromyces siphocampyli-gigantei Berndt Siphocampylus giganteus Ecuador [725]
Uromyces sisyrinchiicola Speg. Sisyrinchium iridifolium Chile [258]
Uromyces skottsbergii Jørst. Enargea marginata Argentina and the Falkland Islands [726,727]
Uromyces smilacis Mayor Smilax sp. Colombia [226,227]

J. Fungi 2022,8, 633 60 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces snowdeniae Cummins Snowdenia scabra Kenya [140]
Uromyces socius Arthur & Holw. Loranthus spp. and Struthanthus spp. Guatemala and México [15,281]
Uromyces solani Dietel & Holw. Solanum appendiculatum Colombia, Guatemala, and México [227,281,456,479]
Uromyces solariae Dietel Solaria miersioides Chile [240]
Uromyces solidaginis (Sommerf.) Fuckel Solidago spp. Colorado, Finland, Idaho, Japan, Montana,
Oregon, Poland, Russia, Washington, and
Wyoming [92,329,454,559]
Uromyces solidaginis-caricis Arthur Carex varia Indiana [728]
Uromyces solidus Berk. & M.A. Curtis Desmodium strictum North Carolina [657]
Uromyces sommerfeltii Hyl., Jørst. & Nannf. Solidago spp.
China, Canada, Germany, Finland, Idaho,
Japan, Russia, Mississippi, Montana, Nepal,
Norway, Oregon, Poland, Russia, Sweden,
and Turkey
[80,94,129–
131,135,185,187,436,475,567]
Uromyces sonorensis J.F. Hennen &
Cummins Merremia palmeri México [729]
Uromyces sophorae Peck Sophora alopecuroides,S. sericea, and S. mollis Pakistan and México [563,730]
Uromyces sophorae-flavescentis Kusano Sophora alopecuroides,S. angustifolia,
S. flavescens,S. jaubertii, and S. japonicum China, Japan, Korea, Turkey, Russia,
and Uzbekistan [96,146,297,483]
Uromyces sophorae-japonicae Dietel Sophora japonica Japan [92,731]
Uromyces sophorae-vicifoliae F.L. Tai Sophora viciifolia China [732]
Uromyces sparaxidis Syd. & P. Syd. Sparaxis lineata,S. tricolor, and Sparaxis sp. South Africa [142]
Uromyces sparganii Cooke & Peck Acorus calamus,Hypericum virginicum, and
Sparganium eurycarpum
Canada, Iowa, Indiana, India, Michigan,
Mississippi, New York, Nebraska, and
Wisconsin [152,185,475]
Uromyces sparsus (Kunze & J.C. Schmidt)
Lév. Arenaria marina,Spergularia spp. and Stellaria patens Bulgaria, Denmark, Finland, Germany,
Nepal, Norway, Romania, and Ukraine [82,111,128,135,179,188,228,329]
Uromyces spartii-juncei P. Syd. & Syd. Spartium junceum France, Greece, Portugal, Spain,
Switzerland,
and Yugoslavia [220]
Uromyces speciosus Holw. Frasera macrophylla Colorado and New México [559,733]

J. Fungi 2022,8, 633 61 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces spegazzinii (De Toni) Arthur Commelina angustifolia,C. elegans,C. erecta,
C. nudiflora, and C. virginica Florida, Taiwan, Texas, and the Virgin
Islands [299,447]
Uromyces spermacoces (Schwein.) Thüm. Diodia teres,D. virginiana,Diodia sp.,
and Spermacoce tenuior
Brazil, Carolina, Florida, Georgia, Indiana,
Missouri, Mississippi, Oklahoma,
Pennsylvania, and Texas [202,203,262,455,584]
Uromyces sphaericus H.S. Jacks. & Holw. Perymenium ecuadoricum Ecuador [210,219]
Uromyces sphaerocarpus Syd. & P. Syd. Indigofera potaninii,I. pseudotinctoria,
I. tinctoria, and Indigofera sp. China and Japan [96,734]
Uromyces sphaerophysae Pospelov ex Nevod. Swainsona salsula China [198]
Uromyces splendens A. Blytt Astragalus oroboides Norway [735]
Uromyces sporoboli Ellis & Everh. Allium spp., Sporobolus spp.
Bolivia, Chile, Indiana, Kansas, Iowa,
Missouri, Nebraska, Puerto Rico, Texas,
and
Wisconsin
[199,386,559,635]
Uromyces sporobolicola J.C. Lindq. Sporobolus marginatus,S. pyramidalis, and
S. pyramidatus Argentina, India, Pakistan, and México [171,415,488,736]
Uromyces sporoboloides Cummins Sporobolus berteroanus Ecuador [737]
Uromyces spragueae Harkn. Calyptridium umbellatum,C. umbellatum var.
caudiciferum,Spraguea umbellate, and Spraguea sp. California, Oregon, and Wyoming [186,559,658]
Uromyces standleyanus Arthur Gaudichaudia schiedeana El Salvador [301]
Uromyces statices Berk. & M.A. Curtis Statice sp. California [598]
Uromyces statices-mucronatae Malençon Statice mucronata Morocco [279]
Uromyces statices-sinensis Liou & Y.C. Wang Statice sinensis China [543]
Uromyces steironematis Arthur Spartina michauxiana Nebraska [738]
Uromyces stellariae Syd. & P. Syd. Stellaria kotschyana Iran [739]
Uromyces stellariae-saxatilis L. Guo & Y.C.
Wang Stellaria media,S. saxatilis,S. vestita, and Stellaria sp. China [96]
Uromyces stenorrhynchi Henn. Stenorrhynchus sp. Peru [740]

J. Fungi 2022,8, 633 62 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces stipinus Tranzschel & Erem. Stipa rubens Russia [106]
Uromyces strauchii Doidge Clutia daphnoides Southern Africa [138]
Uromyces striatellus Tranzschel Euphorbia esula,E. leptocaula,E. sieboldiana,
and Euphorbia sp. China, Iran, Japan, Ukraine, and Russia [94,108,109,157,297]
Uromyces striatus J. Schröt.
Argyrolobium spp., Cicer arietinum,Ervum lens,
Euphorbia spp., Hosackia sp.,
Hymenocarpos circinnatus,Lathyrus odoratus,
Lens culinaris,Lotus spp., Manihot esculenta,
Medicago lupulina,Melilotus spp., Pisum spp.,
Trifolium spp., and Vicia spp.
Worldwide [104,165,220,222,225]
Uromyces striolatus Tranzschel Euphorbia aff-esula,E. boissieriana,E. cyparissias,
E. esula, and Euphorbia sp. Bulgaria, China, Iran, and Pakistan [82,108,157,317,356]
Uromyces strobilanthis Barclay Pteracanthus urticifolius,Strobilanthes alata,
S. dalhousieana, and Strobilanthes sp. India, Pakistan, Taiwan, and the
Philippines [218,563,565]
Uromyces strumariae A.R. Wood Strumaria gemmata Western Cape Province [741]
Uromyces struthanthi Perd.-Sánch. Struthanthus sp. Panama [15]
Uromyces stylochaetonis Doidge Stylochaeton natalense KwaZulu-Natal [249]
Uromyces sublevis Tranzschel Euphorbia baetica,E. petrophila, and E. tictoria Lebanon, Portugal, and Ukraine [157,158,422,449]
Uromyces substriatus Syd. & P. Syd. Lupinus argenteus Montana [79]
Uromyces suksdorfii Dietel & Holw. Lychnis drummondii,Lychnis sp., Silene oregano,
S. pacifica,S. scouleri, and Silene sp. Arizona, California, Idaho, Oregon, Utah,
México, and Washington [114,116,187,454,559]
Uromyces superfixus Vestergr. Bauhinia mirandina,B. mollis, and Bauhinia sp. Argentina, Brazil, Bolivia, and Venezuela [216,394,431]
Uromyces superfluus P. Syd. & Syd. Panicum antidotale and P. depauperatum America, India, and Pakistan [144,171,294]
Uromyces superstomatalis Berndt Cayaponia rigida France [127]
Uromyces suzukii Sawada ex Hirats. f. Sigesbeckia orientalis China, Japan, and Taiwan [94,108,319]
Uromyces symaethidis W. Schneid. Simethis bicolor Italy [742]
Uromyces tairae Hirats. f. Messerschmidia argentea Japan [743]
Uromyces tarapotensis Henn. Camptosema sp. Peru [575]
Uromyces teheranicus Petr. Trifolium retense Iran [744]

J. Fungi 2022,8, 633 63 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces tehuelches Speg. Alstroemeria patagonica Argentina [167,592]
Uromyces tener J. Schröt. Manettia gracilis Brazil [219,331]
Uromyces tenuicutis McAlpine Parietaria judaica and Sporobolus spp.
Australia, China, Fiji, Ghana, Guyana,
Grenada, India, Japan, Jamaica, Kenya,
Malawi, México, Malaysia, Nigeria,
Pakistan,
Philippines, Puerto Rico, South Africa,
Sierra,
Taiwan, Uganda, Virgin Island, West Indies,
and Zimbabwe
[97,171,172,184,243,244,280,615]
Uromyces tenuistipes Dietel & Holw. Desmodium spp. and Meibomia sp. Bolivia, Colombia, Cuba, and México [172,479,597,745]
Uromyces teodorescui Rayss Onobrychis crista-galli Israel and Palestine [81]
Uromyces tepicensis J.F. Hennen & Cummins Loeselia amplectens México [292]
Uromyces tessariae (Speg.) J.C. Lindq. Tessaria absinthioides Spain [746]
Uromyces thapsi Opiz ex BubákVerbascum spp. Greece, Romania, Spain, Turkey, Ukraine,
and Uzbekistan [111,146,158,228,421]
Uromyces thellungi Maire Rumex roseus,R. simpliciflorus, and R. vesicarius Canary Islands, Iran, Israel, and Pakistan [472]
Uromyces thelymitrae McAlpine Thelymitra antennifera and T. flexuosa Australia [81,280,317,356]
Uromyces thermopsidicola Shimab. Thermopsis chinensis Japan [222]
Uromyces tinctoriicola Magnus Euphorbia spp. Armenia, Germany, Iraq, Morocco, Spain,
Turkey, Romania, and Ukraine [80,196,228,422,519]
Uromyces tingitanus Henn. Rumex spp. The Canary Islands, Libya, and Morocco [197,279]
Uromyces tolerandus H.S. Jacks. & Holw. Manihot esculenta and Manihot sp. Brazil [219]
Uromyces tomentellus Cooke Leguminosae sp. California [747]
Uromyces tordillensis Speg. Euphorbia serpens and E. ovalifolia Argentina, Chile, Córdoba, and Uruguay [371,386]
Uromyces tosensis Henn. Commelina communis Japan [711]
Uromyces tournefortiae Henn. Tournefortia sp. Brazil [600]
Uromyces tragi Wakef. & Hansf. Tragus berteronianus Malawi and Uganda [168,273]
Uromyces transcaspicus Petr. Astragalus angustidens Turkmenistan [748]

J. Fungi 2022,8, 633 64 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces transversalis (Thüm.) G. Winter Crocosmia spp., Freesia refracta,Gladiolus spp.,
Tritonia spp., and Watsonia spp.
Argentina, Australia, Brazil, California,
Cuba, England, Florida, France, Malwai,
Martinique, Mauritius, México,
New Zealand, South Africa, Tanzania,
Uganda, Venezuela, Zambia, and
Zimbabwe
[69,142,209,250,273,487,530,650]
Uromyces tranzschelii Syd. & P. Syd. Euphorbia crenulata,E. lucida,E. montana,
E. palmeri,E. robusta, and E. robusta Azirona, Colorado, Montana, Oregon, Utah,
and Wyoming [103,157,187,292,368,412]
Uromyces traucoensis Monteal. & Oehrens Selliera radicans Chile [749]
Uromyces triandrae T.S. Ramakr. & Sriniv. Themeda triandra India [669]
Uromyces trichoclines Henn. Trichocline polymorpha Brazil [219,323]
Uromyces tricholenae Gonz. Frag. & Cif. Tricholaena rosea Dominican Republic [750]
Uromyces trichoneurae Doidge Astrebla elymoides,A. lappacea,A. pectinate,
A. squarrosa,Trichoneura grandiglumis, and T. lisboae Australia, India, South Africa [169,488,751]
Uromyces tricorynes McAlpine Tricoryne elatior Australia [245,267]
Uromyces trifolii-megalanthi
(Dietel & Neger) H.S. Jacks. & Holw. Trifolium peruvianum and Trifolium sp. Brazil, Chile, and Peru [101,126]
Uromyces trifolii-purpurei Const. Trifolium campestre,T. eriosphaerum,T. purpureum,
T. scabrum,T. stellatum, and Trifolium sp.
Bulgaria, Greece, France, Israel, Italy,
Russia,
Romania, and Spain [81,82,126,228,327]
Uromyces trifolii-repentis Liro Trifolium spp. Worldwide [225,361]
Uromyces trigonellae Pass. Trigonella spp. Bulgaria, Israel, Romania, and Turkey [80–82,197,228,357]
Uromyces trigonellae-occultae Henn. Trigonella occulta Egypt [752]
Uromyces tripogonicola Payak & Thirum. Tripogon lisboae India [753]
Uromyces tripogonis-sinensis Y.C. Wang Tripogon chinensi and T. lisboae China and India [106,108,109]
Uromyces tripsaci F. Kern & Thurst. Tripsacum dactyloides Ecuador and Venezuela [431,754]
Uromyces triquetrus Cooke Ascyrum hypericoides,Hypericum spp.,
and Triadenum japonicum
Argentina, Brazil, California, Canada,
China,
Colombia, Indiana, Indonesia, Iowa, Maine,
México, Missouri, Mississippi, Taiwan,
Oregon, Japan, and Wisconsin
[96,97,187,219,252,255,292,412,415,
449,475]
Uromyces triseti Katajev Trisetum cavanillesii Turkmenistan [106]

J. Fungi 2022,8, 633 65 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces triteleiae Dietel & Neger Brodiaea porrifolia,Leucocoryne alliacea,
and Triteleia porrifolia Chile [162,166,240,415]
Uromyces trollii-caroli Ulbr. Trifolium sp. Germany and India [34,755]
Uromyces trollipii Kalchbr. & MacOwan Roepera foetida and Zygophyllum foetidum South Africa [138,385]
Uromyces tropaeoli Ranoj. Tropaeolum major Yugoslavia [212]
Uromyces truncatulus Trotter Geranium versicolor Greece [421]
Uromyces truncicola Henn. & Shirai Sophora japonica and Sophora sp. China, Japan, and Korea [468,483,734]
Uromyces tuberculatus Fuckel Euphorbia spp.
The Balearic Islands, Canary Islands, China,
Finland, Germany, Iran, Malaysia,
Romania,
Pakistan, Portugal, Spain, and Uzbekistan
[105,129,130,228,317,318,329,356]
Uromyces tulipae Dietel Tulipa edulis Japan [756]
Uromyces tungurahuensis Syd. Aspilia lanceolata Ecuador [757]
Uromyces turcomanicus Katajev Muscari leucostomum,Hordeum bulbosum,
H. marinum, and H. spontaneum Iraq, Jordan, Libya, Russia, and
Turkmenistan [126,243,244]
Uromyces tylosemae Gjaerum Tylosema fassoglensis Uganda and Sudan [428]
Uromyces uleanus Dietel Euphorbia sp. Brazil [143,219]
Uromyces umiamensis Berndt & Baiswar Cucumis sp. and Momordica cochinchinensis India [127,212,234]
Uromyces undulatoparietis B. Li Ligularia hookeri China [108,109]
Uromyces undulatus Tranzschel Euphorbia condylocarpa and Euphorbia sp. Armenia and Iran [157,318]
Uromyces unioniensis Viégas Desmodium sp. and Meibomia sp. Brazil [394,640]
Uromyces unitus Peck Calandrinia leeana Washington (USA) [758]
Uromyces urariae B. Li Uraria lagopodioides China [759]
Uromyces urbanianus Henn. Oryctanthus spicatus Argentina, Brazil, Colombia, Guatemala,
Honduras, México, and Trinidad and
Tobago [15,227,473]
Uromyces urgineae M.S. Patil Urginea indica India [397]
Uromyces ushuwaiensis Speg. Impatiens hochstetteri Ethiopia [258,760]
Uromyces ustalis Tranzschel Ranunculus repens Siberia [761]

J. Fungi 2022,8, 633 66 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces usterianus Dietel Myrtaceae Colombia and São Paulo [762]
Uromyces valens F. Kern Carex aestivalis,C. lupulina, and C. utriculata Indiana, North Carolina, and Wisconsin [763–765]
Uromyces valerianae (Schumach.) Fuckel Valerianella spp.
Bulgaria, China, Czech Republic,
Czechoslovakia, Denmark, Finland, France,
Germany, Greece, Japan, Mongolia,
Norway,
Poland, Romania, Scotland, Spain,
South Africa, Sweden, Switzerland, Taiwan,
Turkey, and Ukraine
[80,82,90,94,128,132,133,135,138,
228,283,421,440]
Uromyces valerianae-microphyllae Berndt Valeriana microphylla Ecuador [766]
Uromyces valerianae-wallichii
(Dietel) Arthur & Cummins Valeriana spp. China, Japan, Taiwan, India, and Pakistan [87,88,94,108,109,356]
Uromyces valesiacus E. Fisch. Vicia ervilia and V. onobrychioides Albania, Bulgaria, France, Greece, Romania,
Switzerland, Turkey, and Yugoslavia [82,126,220]
Uromyces vanderystii Henn. Teramnus labialis The Democratic Republic of the Congo [610]
Uromyces vankyorum Berndt Atriplex lampa Argentina [282]
Uromyces venustus Dietel & Holw. Cestrum nitidum México [292,375]
Uromyces veratri (DC.) J. Schröt. Adenostyles spp., Homogyne alpine,Ligularia spp.,
and Veratrum spp.
Austria, Bulgaria, China, Germany, Japan,
Korea, Poland, Romania, Russia, Spain,
Switzerland, and Turkey [82,135,158,222,228,231,283,402]
Uromyces verbasci Niessl Verbascum spp. Bulgaria, Germany, Poland, and Ukraine [82,105,135]
Uromyces verrucosae-craccae Mayor Vicia cracca France and Switzerland [126,220]
Uromyces verruculosus J. Schröt. Arenaria serpyllifolia,Cucubalus baccifer,
Dianthus armeria,Lychnis spp., Melandrium spp.,
and Silene spp.
Armenia, Bulgaria, Czech Republic,
Denmark, Greece, Germany, Indiana, Iran,
Michigan,
New York, Norway, Poland, Romania,
Sweden, Texas, Turkey, Ukraine,
and Wisconsin
[105,107,135,228,317,422,519]
Uromyces verus H.S. Jacks. & Holw. Bauhinia rufa Brazil [101]
Uromyces vesicatorius (Bubák) Nattrass Lens esculenta and L. leontopetalum Cyprus, Iraq, Israel, and Turkey [80,81,196,599]
Uromyces vesiculosus G. Winter Zygophyllum spp. Australia [245]

J. Fungi 2022,8, 633 67 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces vestergrenii P. Syd. & Syd. Bauhinia tomentosa India and Sri Lanka [165]
Uromyces viciae-craccae Const. Ervum lens,Euphorbia sp., Lathyrus aphaca,
Lens spp., and Vicia spp.
Armenia, Austria, Azerbaijan, Bulgaria,
China, Czech Republic, Finland, France,
Germany,
Georgia, Greece, Hungary, Iran, Itlay,
Kazakhstan, Poland, Romania, Russia,
Siberia, Switzerland, Ukraine, and
Yugoslavia
[82,135,220,767,768]
Uromyces viciae-fabae (Pers.) J. Schröt. Vicia spp., Lathyrus spp., and Pisum spp. Worldwide [71,222,236,238,769]
Uromyces viciae-unijugae S. Ito Vicia unijuga Japan [92]
Uromyces vicinus H.S. Jacks. & Holw. Ipomoea sp. Brazil [219]
Uromyces vicosensis R.T. Almeida Bauhinia sp. Brazil [219]
Uromyces viegasii R.T. Almeida Bauhinia forficata and Bauhinia sp. Brazil [219,394]
Uromyces viennot-bourginii J. Anikster & I.
Wahl Bellevalia eigii and Hordeum spontaneum Israel [81]
Uromyces vignae Barclay Desmodium spp., Dipogon spp., Lablab sp.,
Phaseolus sp., Sphenostylis sp., and Vigna vexillata Worldwide [71,222,225]
Uromyces vignae-luteolae Henn. Vigna luteola Congo [610]
Uromyces vignae-sinensis Miura Vigna sinensis China and Japan [89,96]
Uromyces visci T. Majewski & K.A. Nowak Viscum congolense Rwanda [551]
Uromyces volkartii Gäum. & Terrier Trisetum flavescens Switzerland [770]
Uromyces vossiae Barclay Phacelurus speciosus,Rottboellia sp., and
Vossia speciosa India and Pakistan [106,294,771]
Uromyces vulpiae Sousa da Câmara Vulpia broteri Portugal [772]
Uromyces waipoua McNabb Hypericum gramineum and H. japonicum New Zealand [313]
Uromyces wartoensis Petr. Astragalus wartoensis Armenia and Turkey [220,276]
Uromyces wedeliae Henn. Wedelia bicolour,W. biflora,W. chinensis,
W. menotriche, and W. prostrata China, Eritrea, Fiji, Japan, Micronesia,
Philippines, Taiwan, and Tonga [94,97,108,109,287,565,654]
Uromyces wedeliae-biflorae Boedijn Wedelia biflora Indonesia [309]
Uromyces wellingtonicus T.S. Ramakr. & K.
Ramakr. Sporobolus indicus India [629]

J. Fungi 2022,8, 633 68 of 99
Table 3. Cont.
Species Host Locality Reference
Uromyces wiehei Cummins Thalictrum rhynchocarpum Kenya, Malawi, and Uganda [318,609,712]
Uromyces wolfii Cummins Borreria laevis Venezuela [609]
Uromyces wulffiae Henn. Wulffia baccata,W. maculata,W. scandens,
W. stenoglossa, and Wulffia sp. Brazil [219,394,575]
Uromyces wulffiae-stenoglossae Dietel Wulffia baccata,W. maculata,W. stenoglossa,
and Wulffia sp.
Brazil, French Guiana, Guyana, Trinidad
and
Tobago, Venezuela, and West Indies [120,219,264,431,557]
Uromyces yakushimensis Hirats. f. & Katsuki Trichosanthes bracteata,T. multiloba, and T. palmata Japan and Myanmar [94,127,212]
Uromyces yoshinagae Henn. Pisum sativum Honshu [360]
Uromyces yurimaguasensis Henn. Clitoria guianensis and Clitoria sp. Belize and Peru [172,575,721]
Uromyces zeyheri BubákIxia scillaris,I. scillaris, and Tritonia pallida South Africa [123,138,165]
Uromyces zigadeni Peck Zigadenus paniculatus Utah [328]
Uromyces zizaniae-latifoliae Sawada Zizania latifolia Taiwan [773]

J. Fungi 2022,8, 633 69 of 99
5.5. Endemic/Native Uromyces Species
The distribution of rust fungi is governed by various ecological factors that affect their
host range and endemism. As with other biotrophic pathogens, rust fungi are also found
to be co-evolved with their host plants, which possibly justifies their host specificity and
narrow host range. However, cross-infection in rust fungi is also observed, which opens
up the need for further investigation of their host range. Among various rust fungi, the
genus Uromyces also exhibited a broad host diversity among different terrestrial regions
of the world. An analysis of data collected in the present study revealed the restriction
of the Uromyces species towards a single host. More than 400 species of Uormyces were
found to be endemic to more than 100 countries, provinces, or islands. The distribution
of endemic species of Uromyces showed that the highest number of endemic species was
found in Brazil (41 species) and South Africa (33 species). Other countries, such as China
(18), Chile (18), Japan (17), India (17), México (15), France (12), Argentina (11), Australia
(11), and Ecuador (11), showed a medium to low endemic distribution of these fungi. If
we compare the continental distributions, it is clear from the geographical heat map that
there is a big difference between the prevalence of endemic species in North and South
America. In fact, South America showed the occurrence of the endemic Uromyces species.
Similarly, Asia also holds a high number of species endemic to the different countries or
regions in the continent. The distribution of the endemic species was found to be scattered
compared to European countries; however, the number of endemic species in this region
was comparatively high. Detailed information on the distribution of endemic or native
Uromyces species is presented in Table 4and Figure 9.
Table 4. World distribution of endemic/native Uromyces species.
Uromyces Species Host Country Reference
Uromyces actinostemonis Actinostemon concolor Brazil [103]
Uromyces acutatus Gageabohemica and G.villosa Germany [105]
Uromyces aegopogonis Aegopogon cenchroides,A.geminiflorus,
A.gracilis, and A.tenellus México [106]
Uromyces aeluropodinus Aeluropus littoralis Ukraine [111]
Uromyces agnatus Jatropha stimulosa Florida [112]
Uromyces agropyri Agropyron sp. India [126]
Uromyces aimeae Cucurbitaceae Ecuador [127]
Uromyces albiziae Albizia procera Indinesia [136]
Uromyces allii-monanthi Allium monanthum Japan [147]
Uromyces allii-sibirici Allium sibiricum Norway [149]
Uromyces allii-victorialis Allium fistulosum,A.macrostemon,
and A.victorialis China [90]
Uromyces alsinis Minuartia hamata and M.meyeri Turkey [80]
Uromyces alyxiae Alyxia oliviformis Hawaii [176]
Uromyces amphilophis-insculptae Amphilophis insculpta India [194]
Uromyces anabasis Anabasis aphylla China [195]
Uromyces anomathecae Anomatheca cruenta South Africa [169]
Uromyces anotidis Anotis richardiana Sri Lanka [214]
Uromyces antioquiensis Rhynchospora polyphylla and R.nervosa Colombia [119,226,227]
Uromyces antipae Rosa lutea Romania [228]
Uromyces aphelandrae Aphelandra pectinata Costa Rica [218]

J. Fungi 2022,8, 633 70 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces aquiriensis Cucurbitaceae Israel [127]
Uromyces araucanus Senecio otites Chile [240]
Uromyces arenariae-grandiflorae Arenaria saponarioides Turkey [80]
Uromyces argutus Spartina alterniflora and S.glabra Florida [106]
Uromyces asperulae Asperula conferta and A.oligantha Australia [268]
Uromyces aspiliellus Aspilia latifolia Ivory Coast [270]
Uromyces astragali-alopecuri Astragalus alopecurus Turkey [276]
Uromyces astragali-atropilosuli Astragalus atropilosulus Kenya [276]
Uromyces
astragali-pseudoutrigeris Astragalus pseudoutriger Turkey [276]
Uromyces atlanticus Hippocrepis scabra Morocco [279]
Uromyces azorellae Pozoa trifoliate and
Schizeilema trifoliolatum New Zealand [284]
Uromyces babianae Babiana disticha Western Cape
Province [285]
Uromyces baccarinii Wedelia sp. Eritrea [287]
Uromyces badius Haemanthus coccineus,H.pumilio,
H.rotundifolius, and H. sanguineus South Africa [167]
Uromyces bahiensis On leaves of Loranthaceae Panama [15]
Uromyces bauhiniicola Bauhinia chlorantha and B.pringlei México [398]
Uromyces beckmanniae Beckmannia eruciformis and B.
syzigachne Oregon [295]
Uromyces belemensis Ormosia nobilis Brazil [219]
Uromyces bermudianus Cyperus paniculatus Bermuda [224]
Uromyces bethelii Silene verecunda California [301]
Uromyces bisbyi Eriogonum parvifolium California [314]
Uromyces boissierae Boissiera pumilio Iran [320]
Uromyces bolusii Aspalathus pachyloba South Africa [322]
Uromyces bomareae Bomarea sp. Brazil [167]
Uromyces bonaerensis Gomphrena elegans Buenos Aires [324]
Uromyces bonae-spei Tritonia scillaris and Acidanthera
pallida Southern Africa [325]
Uromyces bonaveriae Bonaveria securidaca and
Securigera securidaca Greece [327]
Uromyces borreriae Borreria verticillata Rio de Janeiro [331]
Uromyces bosseri Trochomeriopsis diversifolia Madagascar [127]
Uromyces
bothriochloae-intermediae Bothriochloa intermedia China [108,109]
Uromyces bradburyae Bradburya pubescens,B.virginiana,
Centrosema pubescens, and C.
virginianum Brazil [219]
Uromyces brizae Briza media France [339]
Uromyces bromicola Bromus coloratus and B.lithobius Chile [199]
Uromyces buforrestiae Buforrestia imperforata Ghana [341]
Uromyces bulbinicola Bulbine bulbosa Australia [249]
Uromyces bunsteri Sisyrinchium cuspidatum and
S.graminifolium Chile [166]
Uromyces calopogonii Calopogonium galactioides Guatemala [172]

J. Fungi 2022,8, 633 71 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces calotheus Urginea sp. Sierra Leone [351]
Uromyces calycotomes Calycotome spinosa France [220]
Uromyces caricis-brunneae Carex brunnea Japan [94]
Uromyces caricis-schmidtii Carex schmidtii Khabarovsk [297]
Uromyces cassiae-mimosoidis Cassia mimosoides and
Chamaecrista mimosoides South Africa [172]
Uromyces cearensis Ipomoea sp. Brazil [373]
Uromyces cedrelae Toona serrata Indonesia [177]
Uromyces celtidis Celtis sp. Brazil [102]
Uromyces cenisiae Ononis cenisia France [342]
Uromyces chaetobromi Chaetobromus dregeanus and C.
schraderi South Africa [378]
Uromyces chilensis Lathyrus magellanicus and L.multiceps Chile [386]
Uromyces chiovendae Cissus sp. Somalia [287]
Uromyces chorizanthis Chorizanthe pungens California [388]
Uromyces christensenii Muscari parviflorum and
Hordeum bulbosum Israel [390]
Uromyces chubutensis Poa chubutensis Chubut [391]
Uromyces ciceris-soongaricae Cicer songaricum Pakistan [393]
Uromyces circinalis Scilla prasina South Africa [138]
Uromyces cladomanes Vitis sp. Somalia [287]
Uromyces cladrastidis Cladrastis shikokiana Japan [92]
Uromyces clignyioides Monocymbium ceresiiforme Zimbabwe [243]
Uromyces clitoriae Clitoria mexicana México [172]
Uromyces clivalis Argyrolobium flaccidum India [357]
Uromyces cobresiae Carex sp. Uzbekistan [146]
Uromyces collinus Bauhinia sp. México [292]
Uromyces coluteae Colutea arborescens Austria [299]
Uromyces combreti Combretum sp. Myanmar [405]
Uromyces conicus Cleome sp. Bolivia [411]
Uromyces correntinus Rhynchospora tenuis Argentina [415]
Uromyces costesianus Sphaeralcea velutina Chile [386]
Uromyces crepidis-fraasii Crepisfraasii sp. Greece [419]
Uromyces cretensis Coronilla parviflora and C.rostrata Greece [421]
Uromyces crotalariae-nitens Crotalaria nitens Colombia [423]
Uromyces cruchetii Borreria tenella Colombia [226]
Uromyces cucumivorus Cucumis melo Iraq [127]
Uromyces cuenodii Silene eriocalycina Iraq [196]
Uromyces cyanotidis Cyanotis capitata Papua New Guinea [426]
Uromyces cyathulae Cyathula globulifera Eritrea [287]
Uromyces dendroseridis Dendroseris micrantha Chile [240]
Uromyces densus Bidens pilosa Puerto Rico [76,77]
Uromyces desmodiicola Desmodium albiflorum Brazil [102]

J. Fungi 2022,8, 633 72 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces desmodii-leiocarpi Desmodium leiocarpum Brazil [172]
Uromyces dieramatis Dierama spp. South Africa [273]
Uromyces dilucidus Sisyrinchium striatum Argentina [415]
Uromyces diniensis Ononis fruticosa France [460]
Uromyces dipcadi Dipcadi viride Kenya [487]
Uromyces discariae Discaria toumatou New Zealand [313]
Uromyces dispersus Apios fortunei Japan [222]
Uromyces dobremezii Euphorbia stracheyi Nepal [464]
Uromyces doebbeleri Hypericum irazuense Costa Rica [310]
Uromyces dorystaechadis Dorystaechas hastata Turkey [471]
Uromyces drimiopsidis Drimiopsis maculata South Africa [138]
Uromyces dubiosus Lantana sp. Goiás[291]
Uromyces ducellieri Anabasis aphylla China [91]
Uromyces dusenii Gilliesia graminea,G.monophylla,
Miersia chilensis, and Ornithogalum
biflorum Chile [240]
Uromyces echinodes Asclepiadaceae Suriname [473]
Uromyces eclipsis Zygophyllum morgsana South Africa [234]
Uromyces edwardsiae Edwardsia spp. and Sophora spp. New Zealand [284]
Uromyces ehrhartae-giganteae Ehrharta spp. South Africa [429]
Uromyces ellipticus Glycyrrhiza astragalina Chile [240]
Uromyces ellisianus Euphorbia marginata Minnesota [277]
Uromyces emmeorhizae Emmeorhiza umbellata Venezuela [478]
Uromyces eriogoni Eriogonum virgatum California [388]
Uromyces ermelensis Indigofera sp. South Africa [138]
Uromyces erythrinae Erythrina sp. Ecuador [165]
Uromyces euphlebius Phoradendron calyculatus México [15]
Uromyces euphorbiae-javanicae Euphorbia javanica Indonesia [309]
Uromyces euphorbiae-lunulatae Euphorbia esula,E.Kansui, and E.
lunulata China [108,109]
Uromyces evastigatus Phthirusa pyrifolia El Salvador [15]
Uromyces fiebrigii Bauhinia sp. Paraguay [216]
Uromyces fiorianus Peucedanum fraxinifolium and
Peucedanum sp. South Africa [138]
Uromyces flemmingiae Flemingia sp. Uganda [507]
Uromyces fleuryae Fleurya podocarpa Gabon [508]
Uromyces floralis Bauhinia hiemalis,B.cuyabensis,
B.Holophylla, and B.rufa Brazil [102]
Uromyces floscopae Floscopa peruviana Brazil [102]
Uromyces fontii Peplis acutangula Morocco [510]
Uromyces foveolatus Bauhinia hirsuta and B.mirandina Brazil [102]
Uromyces fuscatus Polygonum alpinum Idaho and Utah [281]
Uromyces fusisporus Acacia neriifolia and A.salicina Australia [177]
Uromyces fremonti Oenothera fremontii Kansas [79]

J. Fungi 2022,8, 633 73 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces galactiae Galactia pedunculata Brazil [517]
Uromyces galii Galium aparine and G.spurium Japan [222]
Uromyces galii-californici Galium californicum and Galium sp. California [520]
Uromyces galphimiae Galphimia glauca and G.humboldtiana México [456]
Uromyces garanbiensis Ehretia dicksonii Taiwan [521]
Uromyces gaubae Caltha introloba Australia [522]
Uromyces gausseni Dorycnopsis gerardii France [516]
Uromyces geissorhizae Geissorhiza sp. Western Cape
Province [523]
Uromyces geraniicola Geranium patagonicum Chile [240]
Uromyces gigantiformis Bidens sp. Colombia [423]
Uromyces globosus Sapium spp. México [103]
Uromyces gnaphalii Gnaphalium sp. Colorado [388]
Uromyces goyazensis Bauhinia sp. Brazil [538]
Uromyces grandiotii Ancrumia cuspidata Chile [540]
Uromyces greenstockii Ipomoea greenstockii South Africa [138]
Uromyces guayacuru Statice brasiliensis Buenos Aires [542]
Uromyces habrochloae Habrochloabul lockii Malawi [244]
Uromyces hainanicus Ipomoea sumatrana China [319]
Uromyces handelii Lotus gebelia Iraq [220]
Uromyces hardenbergiae Hardenbergia monophylla Australia [280]
Uromyces hawksworthii Phthirusa stelis Brazil [60]
Uromyces heimii Medicago arborea France [516]
Uromyces hellebori-thibetani Helleborus thibetanus China [195]
Uromyces hessii Zantedeschia angustiloba Angola [558]
Uromyces heterantherae Heteranthera reniformis Brazil [102]
Uromyces heterogeneus Hibiscus syriacus India [126]
Uromyces heteromallus Haloxylon recurvum Pakistan [563]
Uromyces hewittiae Hewittia bicolor Philippines [565]
Uromyces hidakaensis Pisum sativum Japan [222]
Uromyces himalaicus Lilium sp. Nepal [567]
Uromyces holubii Dracaena sp. Gauteng [573]
Uromyces huallagensis Desmodium sp. Peru [575]
Uromyces hyderabadensis Atylosia scarabaeoides India [579]
Uromyces hydrocotylicola Hydrocotyle sp. China [96]
Uromyces hypericinus Hypericum brasiliense Formosa [180]
Uromyces hypsophilus Euphorbia sp. Mendoza [391]
Uromyces indicus Sporobolus indicus Barbados [589]
Uromyces infarctus Cayaponia sp. Costa Rica [127]
Uromyces inflatus Anisotome sp. New Zealand [591]
Uromyces ingicola Inga sp. Amazonas [575]

J. Fungi 2022,8, 633 74 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces ingiphilus Inga edulis Argentina [592]
Uromyces insignis Echinocephalum latifolium and
Melanthera latifolia Brazil [219]
Uromyces insularis Clitoria cajanifolia Puerto Rico [466]
Uromyces invisus Solanum sisymbriifolium Argentina [415]
Uromyces ipatingae Clitoria fairchildiana Brazil [219]
Uromyces isachnes Isathne kunthiana Sri Lanka [594]
Uromyces jatrophicola Cnidoscolus sp. and Jatropha sp. Brazil [103]
Uromyces juncicola Juncus stipulatus Mendoza [180]
Uromyces johowii Vicia macraei,V.nigricans, and Vicia
sp. Chile [282]
Uromyces kentaniensis Antholyza aethiopica and
Chasmanthe aethiopica South Africa [322]
Uromyces kigesianus Pittosporum abyssinicum Uganda [609]
Uromyces kochianus Geranium nodosum Switzerland [611]
Uromyces koeleriae Koeleria caucasica Russia [106]
Uromyces krantzbergensis Anthericum sp. Namibia [138]
Uromyces kurtzii Senecio spp. Argentina [415]
Uromyces kwangensis Justicia sp. Congo [218]
Uromyces langtangensis Anaphalis nepalensis Nepal [463]
Uromyces largus Chamaesyce lata Colorado [87,88]
Uromyces sp. Lasiocorys abyssinica Eritrea [287]
Uromyces latimammatus Ipomoea sumatrana China [621]
Uromyces leonotidis Leonotis nepetifolia India [75]
Uromyces lereddei Colutea arborescens France [220]
Uromyces lespedezae Lespedeza capitata Vermont [583]
Uromyces lespedezae-bicoloris Lespedeza bicolor and L.formosa China [108,109]
Uromyces
lespedezae-macrocarpae Campylotropis macrocarpa,
Lespedeza bicolor, and L.formosa China [108,109]
Uromyces lespedezae-sericeae Lespedeza sericea Pakistan [208]
Uromyces libycus Lotus pusillus Libya [197]
Uromyces lomandracearum Lomandra longifolia Australia [62]
Uromyces loranthi Loranthus sp. Brazil [376]
Uromyces lotononidicola Lotononis cytisoides South Africa [632]
Uromyces lygei Lygeum spartum Sardegna [325]
Uromyces macnabbii Chionochloa spp. and Danthonia spp. New Zealand [591]
Uromyces mangenotii Vicia pubescens France [516]
Uromyces manihoticola Manihot spp. Brazil [103]
Uromyces manihotis-catingae Manihot spp. Brazil [102]
Uromyces marinus Medicago marina Morocco [220]
Uromyces martinii Melanthera spp. and Bidens spp. Florida [262]
Uromyces massoniae Massonia latifolia South Africa [138]
Uromyces megalosporus Tessaria absinthioides Tucumán[371]

J. Fungi 2022,8, 633 75 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces melandrii Melandrium cucubaloides Los Lagos [162]
Uromyces melasphaerulae Melasphaerula graminea Western Cape
Province [424]
Uromyces melothriae Melothria tomentosa Eritrea [287]
Uromyces meygounensis Euphorbia bungei Iran [638]
Uromyces miersiae Miersia chilensis Chile [540]
Uromyces mikaniae Mikania sp. Brazil [219]
Uromyces mimusops Mimusops sp. South Africa [700]
Uromyces moehringiae Moehringia lateriflora Japan [297]
Uromyces mongolicus Euphorbia kozlovii Mongolia [535]
Uromyces montis-ferrati Euphorbia luteola Northern Africa [649]
Uromyces moraeae Moraea spathacea South Africa [144]
Uromyces mussooriensis Stipa sibirica India [106]
Uromyces myosotidis Myosotis sp. Turkey [470]
Uromyces myristicus Euphorbia bicolor Texas [657]
Uromyces namaqualandus Roepera cordifolia Namibia [384]
Uromyces nassauviae Nassauvia lagascae Argentina [415]
Uromyces nassellae Nassell apubiflora Bolivia [106]
Uromyces natricis Ononis rotundifolia France [342]
Uromyces nattrassii Statice spicata Cyprus [350]
Uromyces naucinus Cayaponia sp. Ecuador [127]
Uromyces nevadensis Primula suffrutescens California [78]
Uromyces notabilis Cyperus sp. and Kyllinga sp. Uganda [487]
Uromyces nothoscordi Nothoscordum striatum Texas [660]
Uromyces numidicus Geranium atlanticum Northern Africa [649]
Uromyces nymphoidis Nymphoides peltata Romania [228]
Uromyces oberwinklerianus Acalypha sp. Costa Rica [310]
Uromyces oblectaneus Rhynchospora corymbosa and R.
exaltata Brazil [219]
Uromyces obscurus Phaseolus sp. México [456]
Uromyces ocimi Ocimum menthifolium Uganda [168]
Uromyces oedipus Sophora japonica Japan [665]
Uromyces oenotherae Oenothera linifolia Illinois [582]
Uromyces oliveirae Bellevalia eigii Israel [390]
Uromyces ophiorrhizae Ophiorrhiza longiflora Indonesia [309]
Uromyces orchidearum Chiloglottis spp. Australia [268]
Uromyces ornatipes Phrygilanthus sonorae México [15]
Uromyces ornithopodioides Ornithopus isthmocarpus and
O.compressus Portugal [317]
Uromyces orthosiphonis Orthosiphon glabratus India [152]
Uromyces otakou Poa spp. New Zealand [591]
Uromyces otaviensis Ipomoea verbascoidea Namibia [385]
Uromyces ovalis Leersia oryzoides Japan [671]

J. Fungi 2022,8, 633 76 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces ovirensis Primula wulfeniana Austria [672]
Uromyces pannosus Bauhinia candicans Brazil [216]
Uromyces papillatus Heteromorpha arborescens South Africa [137]
Uromyces parilis Rumex occultans Israel [81]
Uromyces paspalicola Paspalum racemosum Ecuador [294]
Uromyces pavgii Achyranthes aspera India [677]
Uromyces pavoniae Pavonia racemosa Puerto Rico [259]
Uromyces pazschkeanus Vigna sp. Eritrea [287]
Uromyces penniseti Pennisetum lanatum Pakistan [536,679]
Uromyces pentaceae Pentace burmanica India [680]
Uromyces pentaschistidis Pentaschistis airoides South Africa [243]
Uromyces peraffinis Bauhinia sp. Brazil [219]
Uromyces perlebiae Bauhinia spp. Brazil [102]
Uromyces phalaridicola Phalaris minor Turkmenistan [106]
Uromyces phaseolicola Phaseolus prostratus Argentina [126]
Uromyces phlogacanthi Phlogacanthus celebicus Indonesia [309]
Uromyces phtirusae Phthirusa pyrifolia Colombia [15]
Uromyces phyllachoroides Cynosurus elegans Tunisia [684]
Uromyces physanthyllidis Physanthyllis tetraphylla Greece [544]
Uromyces pittospori Pittosporum abyssinicum Eritrea [287]
Uromyces planiusculus Rumex frutescens Tristan da Cunha [688]
Uromyces plantaginis Plantago barbata and P.tubulosa Argentina [415]
Uromyces poiretiae Poiretia scandens Venezuela [431]
Uromyces polemanniae Polemannia spp. South Africa [142]
Uromyces poliotelis Anguria sp., Gurania sp. and
Selysia prunifera Costa Rica [127]
Uromyces politus Muehlenbeckia cunninghamii Australia [343]
Uromyces polygoni-avicularis Polygonum nepalense Nepal [693]
Uromyces polytriadicola Polytrias amaura Philippines [232]
Uromyces poonensis Sesbania aegyptiaca,S. grandiflora, and
S.sesban India [357]
Uromyces porcensis Inga ingoides Colombia [226]
Uromyces porosus Vicia americana and V.sparsifolia Iowa [155]
Uromyces pozoae Pozoa hydrocotylifolia Chile [240]
Uromyces pratensis Poa pratensis,Ranunculus auricomus,
and R.cassubicus Finland [329]
Uromyces prismaticus Secale montanum Iran [320]
Uromyces procerus Festuca procera Chile [240]
Uromyces pseudarthriae Pseudarthria robusta South Africa [700]
Uromyces psychotriae Psychotria sp. Brazil [219]
Uromyces pulvinatus Euphorbia inaequilatera South Africa [702]
Uromyces quaggafonteinus Ehrharta calycina South Africa [429]

J. Fungi 2022,8, 633 77 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces ramacharii Ocimum sp. India [706]
Uromyces rapaneae Rapanea sp. São Paulo [450]
Uromyces ratoides Cayaponia sp. Ecuador [212]
Uromyces ratus Cayaponia spp. Brazil [212]
Uromyces rayssiae Scilla hyacinthoides Israel [390]
Uromyces rebeccae Suaeda californica California [47]
Uromyces regius Bauhinia candicans Brazil [219]
Uromyces reichei Milla bivalvis and Triteleia
gaudichaudiana Chile [415]
Uromyces reichertii Scilla hyacinthoides and Hordeum
bulbosum Israel [81]
Uromyces reynoldsii Modeccabracteata and Trichosanthes
spp. Myanmar [212]
Uromyces riloensis Doronicum cordifolium Bulgaria [82]
Uromyces rostratus Eriosema sp. Rio de Janeiro [331]
Uromyces rubidus Andropogon condensatus Brazil [199]
Uromyces rugosus Lupinus sp. México [398]
Uromyces rugulosus Campylotropis spp. and Lespedeza spp. China [96]
Uromyces ruiz-leali Anarthrophyllum elegans Argentina [415]
Uromyces rzedowskii Ledenbergia macrantha México [292]
Uromyces sakawensis Solidago virgaurea Japan [711]
Uromyces sasaensis Valerianakilimandscharica and V.
volkensii Uganda [712]
Uromyces satarensis Blainville acmella and B.latifolia China and India [319]
Uromyces saulensis Selysia prunifera France [127]
Uromyces scaberulus Lespedeza bicolor,L. cuneate,
L.cyrtobotrya, and L.formosa China [156]
Uromyces scirpinus Scirpus supinus Philippines [232]
Uromyces secamones Secamone platystigma Uganda [168]
Uromyces sedi Sedum anacampseros France [717]
Uromyces seligeri Lathyrus grandiflorus and L.sylvestris Greece and Russia [421]
Uromyces sellierae Selliera radicans New Zealand [67]
Uromyces semnanensis Astragalus fridae Iran [276]
Uromyces senecionicola Cacalia sp. and Senecio roldana México [292]
Uromyces senecionis-gigantis Senecio gigas Ethiopia [719]
Uromyces seselis Seseli tortuosum Portugal [722]
Uromyces sesseae Sessea sp. Ecuador [325]
Uromyces shahrudensis Onobrychis sp. Iran [723]
Uromyces shikokianus Cladrastis platycarpa and C.shikokiana Japan [222]
Uromyces silenes-chloraefoliae Silene chlorifolia Iran [320]
Uromyces silksvleyensis Bartholina burmanniana Western Cape and South
Africa [378]
Uromyces simulans Vilfa sp. Colorado [724]
Uromyces siphocampyli-gigantei Siphocampylus giganteus Ecuador [725]
Uromyces smilacis Smilax sp. Colombia [227]
Uromyces snowdeniae Snowdenia scabra Kenya [140]

J. Fungi 2022,8, 633 78 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces solariae Solaria miersioides Chile [240]
Uromyces solidaginis-caricis Carex varia Indiana [728]
Uromyces solidus Desmodium strictum North Carolina [657]
Uromyces sonorensis Merremia palmeri México [729]
Uromyces sophorae-japonicae Sophora japonica Japan [92]
Uromyces sophorae-viciifoliae Sophora viciifolia China [732]
Uromyces sparaxidis Sparaxis lineata and S.tricolor South Africa [142]
Uromyces sphaericus Perymenium ecuadoricum Ecuador [210]
Uromyces sphaerophysae Swainsona salsula China [198]
Uromyces splendens Astragalus oroboides Norway [735]
Uromyces sporoboloides Sporobolus berteroanus Ecuador [737]
Uromyces standleyanus Gaudichaudia schiedeana El Salvador [301]
Uromyces statices Statice sp. California [598]
Uromyces statices-mucronatae Statice mucronata Morocco [279]
Uromyces steironematis Spartina michauxiana Nebraska [738]
Uromyces stellariae Stellaria kotschyana Iran [739]
Uromyces stellariae-saxatilis Stellaria media,S.saxatilis, and S.
vestita China [96]
Uromyces stenorrhynchi Stenorrhynchus sp. Peru [740]
Uromyces stipinus Stipa rubens Russia [106]
Uromyces strauchii Clutia daphnoides Southern Africa [138]
Uromyces struthanthi Struthanthus sp. Panama [15]
Uromyces substriatus Lupinus argenteus Montana [79]
Uromyces superstomatalis Cayaponia rigida France [127]
Uromyces tarapotensis Camptosema sp. Peru [575]
Uromyces teheranicus Trifolium retense Iran [744]
Uromyces tehuelches Alstroemeria patagonica Argentina [167]
Uromyces tener Manettia gracilis Brazil [219]
Uromyces tepicensis Loeselia amplectens México [292]
Uromyces thelymitrae Thelymitra antennifera and T.flexuosa Australia [81]
Uromyces thermopsidicola Thermopsis chinensis Japan [222]
Uromyces tolerandus Manihot esculenta Brazil [219]
Uromyces tomentellus Leguminosae sp. California [747]
Uromyces tosensis Commelina communis Japan [711]
Uromyces tournefortiae Tournefortia sp. Brazil [600]
Uromyces transcaspicus Astragalus angustidens Turkmenistan [748]
Uromyces traucoensis Selliera radicans Chile [749]
Uromyces triandrae Themeda triandra India [669]
Uromyces trichoclines Trichocline polymorpha Brazil [219]
Uromyces tricholenae Tricholaena rosea Dominican Republic [750]
Uromyces tricorynes Tricoryne elatior Australia [245]
Uromyces trigonellae-occultae Trigonella occulta Egypt [752]

J. Fungi 2022,8, 633 79 of 99
Table 4. Cont.
Uromyces Species Host Country Reference
Uromyces tripogonicola Tripogon lisboae Maharashtra [753]
Uromyces triteleiae Brodiaea porrifolia,Leucocoryne alliacea,
and Triteleia porrifolia Chile [415]
Uromyces trollipii Roepera foetida and Zygophyllum
foetidum South Africa [385]
Uromyces truncatulus Geranium versicolor Greece [421]
Uromyces tulipae Tulipa edulis Japan [756]
Uromyces tungurahuensis Aspilia lanceolata Ecuador [757]
Uromyces uleanus Euphorbia sp. Brazil [219]
Uromyces umiamensis Cucumis sp. and
Momordica cochinchinensis India [127]
Uromyces undulatoparietis Ligularia hookeri China [108,109]
Uromyces unioniensis Desmodium sp. and Meibomia sp. Brazil [394]
Uromyces unitus On living leaves of Calandrinia leeana Washington [758]
Uromyces urariae Uraria lagopodioides China [759]
Uromyces urgineae Urginea indica India [397]
Uromyces
valerianae-microphyllae Valeriana microphylla Ecuador [766]
Uromyces vankyorum Atriplex lampa Argentina [282]
Uromyces venustus Cestrum nitidum México [375]
Uromyces verus Bauhinia rufa Brazil [100]
Uromyces vesiculosus Zygophyllum spp. Australia [245]
Uromyces viciae-unijugae Vicia unijuga Japan [92]
Uromyces vicinus Ipomoea sp. Brazil [219]
Uromyces vicosensis Bauhinia sp. Brazil [219]
Uromyces viegasii Bauhinia forficata and Bauhinia sp. Brazil [219]
Uromyces viennot-bourginii Bellevalia eigii and Hordeum
spontaneum Israel [81]
Uromyces vignae-luteolae Vigna luteola Congo [610]
Uromyces visci Viscum congolense Rwanda [551]
Uromyces volkartii Trisetum flavescens Switzerland [770]
Uromyces vulpiae Vulpia broteri Portugal [722]
Uromyces waipoua Hypericum gramineum and H.
japonicum New Zealand [313]
Uromyces wedeliae-biflorae Wedelia biflora Indonesia [309]
Uromyces wellingtonicus Sporobolus indicus India [629]
Uromyces wolfii Borreria laevis Venezuela [609]
Uromyces wulffiae Wulffia baccata,W.maculata,W.
scandens, and W.stenoglossa Brazil [219]
Uromyces yoshinagae Pisum sativum Honshu [360]
Uromyces zeyheri Ixia scillaris,I. scillaris, and Tritonia
pallida South Africa [123]
Uromyces zigadeni Zigadenus paniculatus Utah [328]
Uromyces zizaniae-latifoliae Zizania latifolia Taiwan [773]

J. Fungi 2022,8, 633 80 of 99
J. Fungi 2022, 8, x FOR PEER REVIEW 59 of 88
5.5. Endemic/Native Uromyces Species
The distribution of rust fungi is governed by various ecological factors that affect
their host range and endemism. As with other biotrophic pathogens, rust fungi are also
found to be co-evolved with their host plants, which possibly justifies their host specificity
and narrow host range. However, cross-infection in rust fungi is also observed, which
opens up the need for further investigation of their host range. Among various rust fungi,
the genus Uromyces also exhibited a broad host diversity among different terrestrial
regions of the world. An analysis of data collected in the present study revealed the
restriction of the Uromyces species towards a single host. More than 400 species of
Uormyces were found to be endemic to more than 100 countries, provinces, or islands. The
distribution of endemic species of Uromyces showed that the highest number of endemic
species was found in Brazil (41 species) and South Africa (33 species). Other countries,
such as China (18), Chile (18), Japan (17), India (17), México (15), France (12), Argentina
(11), Australia (11), and Ecuador (11), showed a medium to low endemic distribution of
these fungi. If we compare the continental distributions, it is clear from the geographical
heat map that there is a big difference between the prevalence of endemic species in North
and South America. In fact, South America showed the occurrence of the endemic
Uromyces species. Similarly, Asia also holds a high number of species endemic to the
different countries or regions in the continent. The distribution of the endemic species was
found to be scattered compared to European countries; however, the number of endemic
species in this region was comparatively high. Detailed information on the distribution of
endemic or native Uromyces species is presented in Table 4 and Figure 9.
Figure 9. Geographical heat map showing distribution of endemic/native Uromyces species.
Figure 9. Geographical heat map showing distribution of endemic/native Uromyces species.
6. Discussion
Uromyces is the second-largest rust genus, the species of which are phytopathogenic to
any category of plants, causing severe damage and reducing growth and yields. The present
study provides literature-based complete information on this rust in a single compilation.
In addition to being distributed worldwide on vascular plants, Uromyces species cause
several damaging diseases on major agricultural crops such as alfalfa (Medicago sativa), bean
(Phaseolus vulgaris), carnation (Dianthus caryophyllus), chickpea (Cicer arietinum), clover (Tri-
folium sp.), and pea (Pisum sativum). This study contributes to a better understanding of the
taxonomy of these rust fungi in terms of their taxonomic placement, biology, pathogenicity,
life cycle, diversity, and distribution. The information presented in this study helps to
better understand all possible aspects of Uromyces in a single document.
The genus Uromyces is distributed globally on around 647 plant genera belonging to
95 plant families. Poaceae and Fabaceae are the most affected families, with the occurrence
of more than 100 species of Uromyces. However, these fungi infect about 95 species; their oc-
currence on Poaceae and Fabaceae reflects the specificity of these rusts to grasses and legumes.
In addition to host diversity, the distribution of species of Uromyces exhibited a wide range
across the globe. The distribution of this genus extends to over 150 countries, territories,
and occupancies of the world. In its continental diversity distribution, North America is
followed by Asia, Europe, South America, Africa, and Oceania, respectively. A large varia-
tion in the geographical distribution along with the vast diversity of hosts demonstrated
the impact of significantly changing climatic zones on rust fungi. Besides this, more than
400 Uromyces species
are endemic to more than 100 countries, provinces, and islands. This
may be due to the climatic conditions and precise distribution of hosts. Available studies on
global diversity and distribution are rare; however, regional descriptions are available. A
checklist of rust fungi of New Zealand provided by Mckenzie [
313
] reported the occurrence
of 31 species of Uromyces. Similarly, Bahcecioglu and Kabaktepe [
80
] reported 74 species
from Turkey, while Afshan and Khalid [
536
] reported 15 species of the grass family Poaceae
from Pakistan. In India, 97 species of Uromyces have been reported on various hosts [
75
].

J. Fungi 2022,8, 633 81 of 99
A total of 61 Uromyces species were reported from Portugal, whereas about 91 were from
Iran [
430
]. The occurrence of species of Uromyces reported from different countries also
supports the broad diversity of these fungi on a wide range of hosts.
Although 988 species of Uromyces investigated in the present study are found all over
the world, only 73 species are known to have DNA sequence data. As in the case of other
rust fungi, the species of Uromyces are also difficult to culture, which may be one of the
major factors behind the reduced availability of molecular data. In addition, the isolation
of DNA directly from rust fungi present on a natural host and then its sequencing is not
simple or easy, which also affects the molecular studies of these fungi. Phylogenetic studies
based on LSU and ITS sequence data revealed that Uromyces species are polyphyletic taxa
and required more DNA-based analyses for a better understanding of their taxonomic
placement. The polyphyletic nature of Uromyces species was also confirmed by Aime and
McTaggart [
5
] in their study to propose a higher ranking classification for rust fungi, with
notes on genera. Similarly, this was also proposed by Gautam et al. [
34
] during their study
on Indian Pucciniales with the description of the taxonomic outline, including important
descriptive notes. Overall, the present study proposes the requirement of fresh collections
of Uromyces species and their molecular characterization to generate molecular data so that
their phylogenetic relationships can be explained more precisely. The development of a
universal digital platform exclusively for global rust fungi should be developed for the
benefit of researchers working on this specific group of fungi.
7. Conclusions
Being the second-largest plant pathogenic rust genus, Uromyces showed a great varia-
tion with respect to its diversity and distribution. After a complete analysis of information
gathered in the present study, it was concluded that the species of rust genus Uromyces are
distributed globally. Their distribution has been reported in over 150 countries and territo-
ries or occupancies of the world. However, the genus Uromyces predominantly showed
its great diversity in North America in comparison to other continents. Approximately
647 plant genera belonging to 95 plant families are reported to be affected by these rust
pathogens. Apart from this, the endemic nature of this genus is also revealed, which
concluded that more than 400 species of Uormyces are found to be endemic in more than
100 countries. The biocontrol nature of some species of Uormyces is also elucidated in this
study. Moreover, analyses of LSU and ITS sequence data revealed the polyphyletic nature
of species of Uromyces. Further DNA-based analyses of rust disease caused by Uromyces are
still required to develop a better understanding of their taxonomic placement.
Author Contributions:
Conceptualization, A.K.G. and R.K.V.; methodology, A.K.G., S.A., and S.;
software, R.K.V., A.K.G., B.D. and M.N.; validation, A.K.G., R.K.V., N.S., and S.C.K.; formal analysis,
A.K.G., N.S., and M.N.; investigation, A.K.G., R.K.V., and S.C.K.; resources, A.K.G., R.K.V., S., and
N.S.; data curation, A.K.G., R.K.V., and N.S.; writing—original draft preparation, A.K.G., R.K.V., N.S.,
S., and S.C.K.; writing—review and editing, A.K.G., N.S., S.C.K., and R.S.J.; visualization, N.S., R.K.V.,
and A.K.G.; supervision, A.K.G. and N.S.; project administration, A.K.G., R.K.V., and N.S.; funding
acquisition, N.S. All authors have read and agreed to the published version of the manuscript.
Funding:
The authors gratefully acknowledge the financial support provided by Chiang Mai Univer-
sity, Thailand.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Acknowledgments:
The authors wish to thank their respective organizations for providing the
necessary laboratory facilities and valuable support during the study. The publication of this article
was funded by Chiang Mai University, Thailand.
Conflicts of Interest: The authors declare no conflict of interest.

J. Fungi 2022,8, 633 82 of 99
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