Mycol Progress (2007) 6:81-91
DOI 10.1007/s11557-007-0526-0

ORIGINAL ARTICLE

A new species, Dicheirinia panamensis, and new
records of rust fungi from Panama

Jose R. Hernandez • Meike Piepenbring •
Maritza Betzaida Vega Rios

Received: 13 July 2006/Revised: 15 November 2006/Accepted: 19 January 2007/Published online: 11 April 2007
© German Mycological Society and Springer-Verlag 2007

Abstract Based on a recent fieldwork in Panama, 25 species
of rust fungi and several new hosts are reported for the first
time from this country. Among the new records is one new
species, Dicheirinia panamensis on Cojoba rufescens
(Fabaceae). It differs from known species in the genus
Dicheirinia by the presence of uredinia and telia without
paraphyses, irregularly tuberculate urediniospores with two
germ pores on the flattened sides, and tuberculate telio-
spores formed by three probasidial cells, subtended by a
pedicel with three hyaline, apical cells.

Introduction

Among the first fungi reported from Panama were rusts
collected by N.G. Lagerheim (Dietel 1894). Since then,
data on rust fungi (Uredinales, Basidiomycota) from Panama
have been published by numerous authors: Arthur (1918),

Jackson (1926), Standley (1927), Kern and Chardon
(1927), Kern (1938), Hennen and Cummins (1956), J0rstad
(1957), Toler et al. (1959), Ramachar and Cummins (1965),
Cummins (1978), Buritica and Hennen (1980), Ono and
Hennen (1983), Hennen and McCain (1993), Buritica

(1999a, b), Berndt (2002), Hernandez and Hennen (2003),

Hernandez et al. (n.d.), and Piepenbring (2005). In total,
only about 67 species of rust fungi are known from Panama
(Piepenbring 2006), although diversity of plants is very
high and rusts are common in this country.

Several days of intensive field work in Panama yielded
numerous new records of rust species, new host records,
and a species of Dicheirinia on Fabaceae that is different
from all known species and therefore described as new.

Materials and methods

Fresh specimens were collected from 24-29 November
2004 mainly in the province of Chiriqui, not only in agri-
cultural fields and natural topical forest but also on road-
sides between Panama City and Chiriqui. Collections were
pressed in a conventional plant press and slowly dried in an
electric drier. For microscopic examination, material was
mounted in aqueous lactic acid and examined using a Zeiss
Axioplan 2 microscope with bright-field optics. Size ranges
in species descriptions are based on at least 20 measure-
ments of each structure. Digital images were taken using a
Nikon Coolpix 995 and a Nikon DXM 1200. Drawings
were done with a drawing mirror or free hand on scaled
paper, redrawn with ink, and scanned. SEM pictures were
taken from spores dusted onto double-adhesive tape,
covered during 45 s at 5 kV with gold, and observed with
a Hitachi S4500. Plates were made using Adobe Photoshop
Elements 2.0 and Corel Draw 12. Specimens from BPI were

Taxonomical novelty: Dicheirinia panamensis J.R. Hern.,
M. Piepenbr. and Vega Rios.

J. R. Hernandez (*)

Systematic Botany and Mycology Lab, USDA/ARS,

10300 Baltimore Ave.,
Beltsville, MD 20705-2350, USA

e-mail: jose@nt.ars-grin.gov
M. Piepenbring

Institute for Ecology, Evolution and Diversity,
J.W. Goethe-Universität Frankfurt am Main,
60054 Frankfurt am Main, Germany
e-mail: piepenbring@em.uni-frankfurt.de

M. B. Vega Rios

Universidad Autonoma de Chiriqui,
David, Chiriqui, Panama
e-mail: maritzabvr@hotmail.com

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Mycol Progress (2007) 6:81 -91

examined for comparison with different species of Dicheir-
inia and also with other rusts reported in this paper. Authority
names are based on recommendations given in Authors of
Fungal Names (CABI): http://www.indexfungorum.org/
FungalNameAuthors.pdf. Host plants were identified by
the authors as well as Ralph Mangelsdorff (University of
Frankfurt) and Joe Kirkbride and John Wiersema (USDA-
ARS, SBML, Beltsville). Collections were deposited at the
Herbario Nacional of the Universidad de Panama (PMA)
and at the US National Fungus Collection (BPI).

DNA was extracted from leaves of Desmodium sp. and
Phaseolus multiflorus var. coccineus containing sori using
the PureLink Plant DNA Reagent (Invitrogen), and the
identity of P. meibomiae was confirmed using the polymer-
ase chain reaction (PCR) protocol (Frederick et al. 2002)
with Ppm1/Ppa2 primers.

Specimens of Dicheirinia examined

Dicheirinia archeri Cummins on Lonchocarpus nicou DC.
Surinam, Scotelweg, 2 Nov. 1934, leg. W.A. Archer H-256,
0-III (BPI 143084), isotype.

Dicheirinia binata (Berkeley and Curtis) Arthur on Ery-
thrina sp. Guatemala, Livingston, Dept. Izabal, 18 Jan.
1905, leg. W.A. Kellerman 5465, II (BPI 143091), isotype
of Uredo cabreriana Kern. and Kell. On Erythrina glauca
Willd, Venezuela, near Los Caobos, 28 Apr. 1934, leg. F.D.
Kern and R.A. Toro 1713, II (BPI 853921). Venezuela, road
from Caracas to El Valle, 950 m, 9 July 1932, leg.
Chardon and Toro, II (BPI 143115). On Erythrina crista-
galli, Puerto Rico, Agr. Exp. Sta. Mayaguez, 7 July 1924,

leg. H.H. Whetzel, F.D. Kern and R.A. Toro 2415, II (BPI
853922).

Dicheirinia guianensis Cummins on L. nicou DC, Guyana,
Bonisiki Landing, Arawau River, North West District, 16
July 1934, leg. W.A. Archer H-248, 0-I-II-III (BPI 143120),
Isotype, intercepted at New York. On Lonchocarpus sp., 3

June 1938, leg. Insp. Hodson, 0-I-II-III (BPI 143119).

Dicheirinia manaosensis (Henn.) Cummins (=Diorchidium
manaosensis Henn.) on Lonchocarpus rariflorus Martius.

Brazil, Manaos, Rio Negro, Aug 1900, leg. E. Ule, III
(BPI 143122, Mycotheca brasiliensis No. 11), isotype. On
Lonchocarpus sp. Colombia, Letecia, 15 Oct. 1943, leg. C.

O. Grassl 10098, III (BPI 853923).

Dicheirinia ormosiae (Arthur) Cummins (=Puccinia ormo-
siae Arthur) on Ormosia krugii Urban, Puerto Rico, El

Yunque, 14 Apr. 1916, leg. H.H. Whetzel and E.W. Olive

276, II-III (BPI143124), isotype.

Dicheirinia solenioides Cummins on Ormosia sp. (as
Nectandra aff. rigida). Brazil, Sao Francisco, Apr. 1885,
leg. E. Ule, II-III (BPI 143136), isotype.

Dicheirinia spinulosa (J. W. Baxter) J.F. Hennen and
Cummins on Leucaena sp. Mexico (locality unknown),
intercepted at the Nogales Plant Quarantine Station (Ari-
zona, USA) by J.M. Kaiser and D. Noel, III (BPI 115592),
isotype of Diorchidium spinulosum J.W. Baxter.

Dicheirinia superba H. S. Jacks. and Holw. on Piptadenia
sp. (originally erroneously reported as Inga sp.). Brazil, Rio

de Janeiro, Petropolis, 20 Oct 1921, leg. E.W.D. and M.M.

Holway 1234, 0-III (BPI 143138), isotype.

Results

Dicheirinia panamensis J.R. Hern., M. Piepenbr. and Vega
Rios, sp. nov. (Figures 1, 2, and 3).

Etymology The name of the species is based on the country
where it was collected.

Spermogonia et aecia ignota. Uredinia amphigena, prae-
cipue abaxialia, dispersa vel aggregata, ca. 0.1 -1 mm longa,
0.5-1 (2) lata, cinnamomea, subepidermalia, erumpentia;
urediniosporae (30) 37-45 (52)x(15) 19-25 (27) µm, leviter
complanatae, visae de parte lato obovoideae, ellipsoideae,
sphaeroideae vel stellatae, cinnamomeae, episporio tuber-
culato praecipue marginis sporae, pori germinationis 2,
aequatoriis, pedicello hyaline, fragili, 4-5 µm lato. Telia
amphigena, praecipue abaxialia, atro-rufo-brunnea, rotunda
vel elongata, ad 1 mm diam., subepidermalia, erumpentia;
teliosporae (30) 36-40 µm longae, (31) 38-45 (48) µm latae,
atro-cinnamomea, 3-cellulares, cellula 19-24x24-26 µm,
pariete 2-4 µm crassa, tuberculata, tuberculis ad 3 µm
longae; pedicellus hyalinus, 5-7 µm latus, 3 cellulis
apicalibus hyalinis, una pro singulis cellulis probasidialibus.

Spermogonia and aecia unknown. Uredinia amphigenous,
mainly hypophyllous, scattered or more or less grouped,
subepidermal, becoming erumpent, 0.1 -1 mm diameter,
cinnamon brown, sometimes on round chlorotic areas of
the leaf up to 0.5 cm, old lesions brown, up to 1 cm diameter;
urediniospores (30) 37-45 (52)x(15) 19-25 (27) µm, some-
what flattened, in side view ovoid, ellipsoid, spheroid,
star-shaped, cinnamon brown, cell wall 2-4 Hm thick,
tuberculate, tubercles 2-10 Hm long, 2-5 Hm wide and
tapering to a blunt apex, arranged mainly on margin of
spores, sometimes at apex as a crown, sometimes tubercles
very conspicuous giving spore a divided appearance, germ
pores two, equatorial, in center of flattened side of spores,
pedicel hyaline, fragile, 4-5 Hm wide. Telia amphigenous,

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Mycol Progress (2007) 6:81-91

Fig. 1 Dicheirinia panamensis on Cojoba rufescens. a Leaf with
groups of uredinia and telia (from Piepenbring and Vega 3518). b
Group of uredinia and telia, as seen with the stereo microscope (from

Piepenbring and Vega 3518), scale bar= 2 mm. c Teliospores, as seen
by LM (from BPI 871146, type), scale bar=20 µm. d Urediniospores
as seen by LM (from BPI 871146, type), scale bar=20 µm

mainly hypophyllous, close to uredinia, dark chestnut
brown, round, sometimes elongated, up to 1 mm diameter,
subepidermal becoming erumpent; teliospores sometimes in
uredinia, (30) 36-40 µm long, (31) 38-45 (48) µm wide,
yellowish when young, dark cinnamon brown when old,
three probasidial cells delimited by vertical septa, probasidial
cells 19-24x24-26 |j.m, wall 2-4 µm, outer wall tubercu-
late, tubercles up to 3 long, tapering to a blunt apex;
pedicel hyaline, 5-7 µm wide, with three apical hyaline cells
(12x5 (µm), one subtending each probasidial cell.

Holotypus On C. rufescens (Benth.) Britton (Fabaceae,
subfamily Mimosoideae). Panama, Chiriqui, Boquete,
08°46.691'N, 82°25.968'W, 26 Nov. 2004, leg. J.R.
Hernandez and M. Vega Rios 2004-120, II-III (BPI
871146). Isotype in PMA.

Other specimens examined: at the same place on the same
host plant, 27 Aug. 2005, leg. M. Piepenbring and M. Vega
Rios 3518 (PMA); 6 Nov. 2005, M. Piepenbring 3680

(PMA).

On Fabaceae indet. Panama, Chiriqui, Boquete,
08°46.691'N, 82°25.968'W, 26 Nov. 2004, leg. J.R. Hernandez
and M. Vega Rios 2004-122, II-III (BPI 871147, PMA).

Discussion

Species of Raveneliaceae (Uredinales) are particularly
diverse on Fabaceae in the tropics (Hernandez and Hennen
2002). This family includes species in numerous genera,
among which species of Dicheirinia, Diorchidium, Dior-
chidiella, Hapalophragmium, and Ravenelia are morpho-

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Mycol Progress (2007) 6:81-91

Fig. 2 Uredinium, uredinio-
spores, and teliospores of
Dicheirinia panamensis, as seen
by LM. a-c Urediniospores
(from type). d Longitudinal
section of part of an uredinium
(from Piepenbring 3680).
e-h Teliospores (from type)

logically somewhat similar. Species of Dicheirinia differ
from species of the other genera by vertically septate,
tuberculate teliospores, subtended by a hyaline pedicel with
one apical cell for each probasidial cell. The type species of
the genus is D. binata (Berk. and M.A. Curtis) Arthur (=
Triphragmium binatum Berk. and M.A. Curtis) on Eryth-
rina sp. from Nicaragua. Twelve species of Dicheirinia
have been described on the three subfamilies of Fabaceae
(Cummins and Hiratsuka 2003): Caesalpinioideae (Cassia),
Faboideae (Abrus, Chamaecytisus, Cytisus, Erythrina, Lon-
chocarpus, Ormosia, Piptadenia,and Teline), and Mimosoi-

deae (Albizia, Archidendropsis,andLeucaena). Up to now,
no other rusts have been reported on the host species C.
rufescens. Dicheirinia panamensis differs from other species
in the genus by the presence of flattened, tuberculate
urediniospores of irregular shape with two germ pores and
teliospores with three probasidial cells.

Eight of the Dicheirinia species are known only from the
Americas. D. archeri Cummins was described on L. nicou
(Aubl.) DC. from Peru (Spaulding 1961) and Surinam
(Cummins 1937) and on L. urucu Killip and A.C. Sm. and
L. utilis A.C. Sm. also from Peru (Spaulding 1961). It is a

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Mycol Progress (2007) 6:81-91

Fig. 3 Dicheirinia panamensis,
as seen by SEM (from type),
scale bars =10 Hm. a, b
Urediniospores, with central
germpore. c, d Three-celled
teliospores

microcyclic rust with spermogonia and telia and teliospores
with two probasidial cells. D. binata (Berk. and M.A.
Curtis) Arthur was reported on species of Erythrina and an
unknown host (type specimen) from Central America, the
Caribbean, and tropical South America (Farr et al. 2004). It
has paraphysate uredinia, strongly echinulate uredinio-
spores with three to four germ pores, and teliospores with
two probasidial cells. D. guianensis Cummins was reported
on L. nicou (Aubl.) DC. from French Guiana and Guyana
(Cummins 1937; Hernández et al. 2005a, b; Spaulding
1961) and is a macrocyclic rust. The aeciospores and
urediniospores are similar to each other and have an
irregular triangular shape with one germ pore in each angle
of the spore. The teliospores of both species have three
probasidial cells, but those of D. guianensis are less
prominently tuberculate than in D. panamensis. Dicheirinia
manaoensis (Henn.) Cummins was reported on L. rariflorus
Mart. ex Benth. from Brazil (Hennen et al. 2005) and on
Lonchocarpus sp. from Brazil and Colombia (Buriticá and
Pardo-Cardona 1996;Hennenetal.2005;Kernand
Thurston 1954). Spermogonia and telia were reported for
this species: teliospores mostly with three probasidial cells,
(38) 44-55 (65) µm high and (36) 40-55 (59) µm wide,
with tubercles or plaques up to 8 µm thick at the apex of

the probasidial cells. D. ormosiae (Arthur) Cummins was
reported on O. krugii Urb. from Puerto Rico, Santo
Domingo, and Virgin Islands (Cummins 1935; Stevenson
1975) and on Ormosia nobilis Tul. from Brazil (Hennen et
al. 1982). It differs from all other species of the genus in
having a single intercalary cell bearing teliospores with two
probasidial cells (Hennen et al. 2005). Its uredinia and telia
include characteristically branched paraphyses. D. sole-
nioides (Henn.) Cummins was reported on Ormosia
monosperma (Sw.) Urb. (=Ormosia dasycarpa Jacks.) and
Ormosia sp. from Brazil (Cummins 1940; J0rstad 1959). It
produces uredinia and telia with cylindrical hyaline paraph-
yses, triangular, echinulate urediniospores, and teliospores
with two prominently echinulate probasidial cells. D.
spinulosa (J.W. Baxter) J.F. Hennen and Cummins was
reported on Cassia sp. and Leucaena sp. from Mexico
(Cummins 1978; Gallegos and Cummins 1981; Hennen and
Cummins 1973). It produces only telia, which have
teliospores with two to eight probasidial cells. D. superba
H.S. Jacks. and Holw., reported on Piptadenia sp. from
Brazil (Cummins 1935; Hennen et al. 2005; Jackson 1931),
produces spermogonia and telia; teliospores have two
(rarely three) verrucose probasidial cells with cubical
projections.

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Four species of Dicheirinia were reported from outside
the Americas. Dicheirinia canariensis Urries was reported
on Chamaecytisus proliferus (L. f.) Link (as Cytisus
proliferus L. f.), Cytisus congestus Ball, Cytisus perezii
Hutch., and Teline canariensis (L.) Webb and Berth. (as
Genista canariensis L.) from Canary Islands (Gjaerum 1987;
J0rstad 1958; Urries 1954, 1957). D. canariensis produced
only telia, which are paraphysate, and the teliospores have
two to three probasidial cells. Dicheirinia maderensis
Gjaerum was reported on leaves of cf. Teline maderensis
Webb and Berth. from Madeira (Portugal). We did not
examine the type specimen of this species, but, according
to Gjaerum (1982), it produces uredinia with uredinio-
spores subgloboid to ellipsoid, often attenuated at the apex,
27-31 x 18-21 Hm, echinulate, and telia with teliospores
20-29x 11 -13.5 Hm with few warts, while D. panamensis

has larger urediniospores that are variable in shape and
strongly tuberculate and larger teliospores with block-like
tubercles. Dicheirinia trispora Cummins was reported on
Abrus precatorius L. from Mauritius (Cummins 1960).
According to Cummins (1960), it is microcyclic producing
spermogonia and telia with teliospores 26-34x30-40 µm,
which are smaller than those of D. panamensis. Dicheir-
inia viennotii B. Huguenin was reported on Archidendrop-
sis granulosa (Labill.) I.C. Nielsen [=Albizia granulosa
(Labill.) Benth.] from New Caledonia (Huguenin 1966). It
is a macrocyclic rust that produces urediniospores with five
equatorial germ pores, 26.3-36.8x19.3-33.3 µm, and
teliospores with two probasidial cells, 29.8-38.5x
18.9-33.3 µm, with smooth walls.

Hennen et al. (1998) considered Allotelium mirabile Syd.
to belong to the genus Dicheirinia. Cummins and Hiratsuka

Key to species of Dicheirinia

1. Teliospores composed of one or two probasidial cells
2. Telia aparaphysate

3. Teliospores verrucose ........................................................... D. maderensis

3. Teliospores smooth ............................................................... D. viennotii

2. Telia paraphysate

4. Paraphyses cauliflower-like ................................................... D. ormosiae

4. Paraphyses not cauliflower-like

5. Teliospore ornamentation digitate, projections up to 12 µm ........ D. binata

5. Teliospore ornamentation bead-like

6. Paraphyses cylindrical ................................................ D. superba

6. Paraphyses otherwise

7. Paraphyses clavate-capitate ......................................... D. archeri

7. Paraphyses cylindrical to slightly clavate

8. Paraphyses peripheral ........................................ D. solenoides

8. Paraphyses intermingled with teliospores .................. D. canariensis

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1. Teliospores mostly composed of three or four probasidial cells

9. Teliospores mostly composed of four probasidial cells ................... D. spinulosa

9. Teliospores mostly composed of three probasidial cells
10. Telia aparaphysate

11. Urediniospores present, flattened, irregularly tuberculated ...... D. panamensis

11. Urediniospores absent ................................................... D. trispora

10. Telia paraphysate

12. Abundant peripheral branched paraphyses, uredinia produced .. D. guianensis
12. Few peripheral paraphyses, uredinia not produced .................. D. manaoensis

(2003) recognized Allotelium as a separate genus with two
species, A. mirabile Syd. and A. calliandrae (R. Berndt) Y.
Hirats., based on the lack of uredinia and the telia that are
covered by a peridium for a long time.

New records

Twenty-five new records of rust fungi for Panama are listed
below, which bring the total of rusts known from Panama to
92. The considerable number of new records obtained,
based on specimens collected in less than 1 week, suggests
that we are still far from knowing the complete diversity of
rusts in Panama. In Costa Rica, a neighboring country with
similar plant diversity where rusts have been collected
much more intensively than in Panama, 292 species are

known (Berndt 2004).
New records of rusts in Panama

Coleosporium ipomoeae (Schwein.) Burrill on Ipomoea sp.
Panama, Chiriqui Province, Distrito de Alanje, La Barqueta,
8°18.258'N, 82°34.822'W, 27 Nov. 2004, leg. J.R. Hernandez
2004-134, II-III (BPI 864101). Panama, Chiriqui Province,
Quebrada Cermeno, Cascada, 8°30.598TN, 82°26.193'W, 28

Nov. 2004, leg. J.R. Hernandez and R. Mangelsdorff 2004-
149, II (BPI 864102).

C. ipomoeae is widespread in Central, North, South
America and the Caribbean and is reported from several
countries in Africa and Asia on species in the Convolvulaceae
and Pinaceae (Farr et al. 2004; Hernandez et al. 2005a, b).
This is the first report of C. ipomoeae from Panama.

Coleosporium plumeriae Pat. on Plumería sp. Panama,
Chiriquí Province, Quebrada Cermeño, Cascada, 8°30.598N,
82°26.193'W, 28 Nov. 2004, leg. J.R. Hernández and R.
Mangelsdorff 2004-148, II-III (BPI 864104).

C. plumeriae was reported from the Caribbean, Central
America, Mexico, northern South America and the USA
(Florida and Hawaii). It has also been reported from
Micronesia, a green house in Canada (Farr et al. 2004),
and Nigeria (Hernández et al. 2005a, b). This is the first
report of C. plumeriae from Panama.

Coleosporium vernoniae Berk. and M.A. Curtis on Ele-
phantopus mollis Kunth. Panama, Chiriquí Province, Volcán,
8°47.398TN, 82°40.565'W, 25 Nov. 2004, leg. J.R. Hernández
2004-101, II (BPI 864106). Panama, Chiriquí Province,
Boquete, 8°46.691N, 82°25.968'W, 26 Nov. 2004, leg. J.R.
Hernández and M. Vega Rios 2004-125, II (BPI 864107).

C. vernoniae was reported from the Caribbean, Central,
North and South America, and China on species in the
Asteraceae and Pinaceae (Farr et al. 2004; Hernández et al.
2005a, b). This is the first report of C. vernoniae from
Panama.

Desmella aneimiae Syd. and P. Syd. on Thelypteris sp.
Panama, Chiriquí Province, San Felix, 8°16.387'N,
81°51.71'W, 29 Nov. 2004, leg. J.R. Hernández 2004-153,

II-III (BPI 864108).

D. aneimiae was reported from Mexico to Argentina
and the Caribbean on species in the Blechnaceae,
Dennstaedtiaceae, Dryopteridaceae, Nephroleptidaceae,
Schizaeaceae, Pteridaceae, and Thelypteridaceae (Farr et

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al. 2004; Hernandez et al. 2005a, b). This is the first report
of D. aneimiae from Panama.

Gerwasia pittieriana (Henn.) Leon-Gall. and Cummins on
Rubus sp. Panama, Chiriqui Province, Boquete, 8°46.691N,
82°25.968'W, 26 Nov. 2004, leg. J.R. Hernandez and
M. Vega Rios 2004-123, II (BPI 864110).

G. pittieriana was reported from Costa Rica and Mexico
on species of Rubus (Farr et al. 2004). This is the first
report of G. pittieriana from Panama.

Kuehneola malvicola (Speg.) Arthur on Malvaviscus
arboreus Cav. Panama, Chiriqui Province, Caizan,
8°44.406N, 82°49.348'W, 25 Nov. 2004, leg. J.R. Hernandez
2004-109, II (BPI 864198).

K. malvicola was reported from the Americas and Asia
on species in the Malvaceae (Farr et al. 2004). This is the
first report of K. malvicola from Panama.

Phakopsora cupheae Buritica on Cuphea ignea A. DC.
Panama, Chiriqui Province, road to Caizan, 8°47.382'N,
82°42.547'W, 25 Nov. 2004, leg. J.R. Hernandez 2004-103,

II (BPI 871907).

P. cupheae was reported from the Caribbean, Central
America, and northern South America (Farr et al. 2004).
This is the first report of P. cupheae from Panama.

Phakopsora meibomiae (Arthur) Arthur on Desmodium
sp. Panama, Cocle Province, Aguadulce, 8°13.227'N,
80°36.558'W, 24 Nov. 2004, leg. J.R. Hernandez 2004-
084, II (BPI 864113). On P multiflorus var. coccineus (L.)
DC. Panama, Chiriqui Province, Dominical, 8°40.455'N,
82°48.013'W, 25 Nov. 2004, leg. J.R. Hernandez 2004-114,
II (BPI 864114). Identification was confirmed using real-
time PCR (Frederick et al. 2002).

P. meibomiae was reported from Mexico to Argentina
and the Caribbean; in addition, there are several uncon-
firmed reports from out of the Americas (Farr et al. 2004).
This is the first report of P. meibomiae from Panama.

Puccinia arechavaletae Speg. on Serjania decapleuria
Croat. Panama, Chiriqui Province, David, 8°25.947'N,
82°27.045'W, 26 Nov. 2004, leg. J.R. Hernandez 2004-
128, II-M (BPI 864115).

P. arechavaletae is widespread in the Americas on species
in the Sapindaceae (Farretal. 2004; Hernandezetal. 2005a, b).
This is the first report of P. arechavaletae from Panama.

Puccinia hemerocallidis Thum. on Hemerocallis sp. Pana-
ma, Chiriqui Province, Boquete, central plaza, 8°46.691'N,
82°25.968'W, 26 Nov. 2004, leg. J.R. Hernandez and M.
Vega Rios 2004-116, II (BPI 864127).

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P. hemerocallidis was reported from Asia, Australia,
Costa Rica, and the USA (Farr et al. 2004; Hernandez et al.
2002). This is the first report of P. hemerocallidis from
Panama.

Puccinia heterospora Berk. and M.A. Curtis on Hibiscus
sp. Panama, Chiriqui Province, David, El Cabrero,
UNACHI, 8°25.947TN, 82°27.045'W, 26 Nov. 2004, leg.
J.R. Hernandez 2004-130, III (BPI 864206). On Hibiscus
sp. Panama, Chiriqui Province, San Felix, side of San Felix
river, 8°16.387'N, 81°51.71'W, 29 Nov. 2004, leg. J.R.
Hernandez 2004-151, III (BPI 864210).

P. heterospora is cosmopolitan, reported from the
Americas, Africa, Asia, and Oceania on species in the
Malvaceae (Farr et al. 2004). This is the first report of P.
heterospora from Panama.

Puccinia hydrocotyles Cooke on Hydrocotyle bonariensis
Comm. ex Lam. Panama, Chiriqui Province, Boquete,
8°46.691'N, 82°25.968'W, 26 Nov. 2004, leg. J.R. Hernandez

and M. Vega Rios 2004-124, III (BPI 864128).

P. hydrocotyles was reported from the Americas, Africa,
Asia,andOceaniaonspeciesofHydrocotyle (Farr et al. 2004).
This is the first report of P. hydrocotyles from Panama and
confirms the tentative record in Piepenbring et al. (2002).

Puccinia lantanae Farl. on Lantana sp. Panama, Chiriqui
Province, Quebrada Cermeno, 8°30.531'N, 82°26.203'W,
28 Nov. 2004, leg. J.R. Hernandez and R. Mangelsdorff
2004-147, III (BPI 864209).

P. lantanae was reported as tropical and subtropical in
the western hemisphere, Africa, and Asia on species in the
Verbenaceae and Acanthaceae (Farr et al. 2004). This is the
first report of P. lantanae from Panama.

Puccinia lateritia Berk. and M.A. Curtis on Borreria
ocymoides (Burm. f.) DC. Panama, Chiriqui Province,
Volcan, 8°47.398TN, 82°40.565'W, 25 Nov. 2004, leg.
J.R. Hernandez 2004-102, III (BPI 864194). This specimen
confirms the tentative record in Piepenbring et al. (2002).

P. lateritia was reported as tropical and subtropical in the
western hemisphere, Africa, and Asia on species in the
Rubiaceae (Farr et al. 2004). This is the first report of
P. lateritia from Panama.

Puccinia oxalidis Dietel and Ellis on Oxalis sp. Panama,
Chiriqui Province, Caizan, 8°44.406TN, 82°49.348'W, leg. J.
R. Hernandez 2004-107, II (BPI 864197). Panama, Chiriqui
Province, Boquete, 8°46.691N, 82°25.968'W, 26 Nov. 2004,
leg. J.R. Hernandez and M. Vega Rios 2004-117, II (BPI
842600). Panama, Chiriqui Province, Boquete, 8°46.691'N,
82°25.968'W, 26 Nov. 2004, leg. J.R. Hernandez and M. Vega
Rios 2004-118, II (BPI 864201). Panama, Chiriqui Province,

Springer

Mycol Progress (2007) 6:81-91

Boquete, 8°46.691TN, 82°25.968'W, 26 Nov. 2004, leg. J.R.
Hernandez and M. Vega Rios 2004-119, II (BPI 864202).

P. oxalidis was reported as cosmopolitan (Farr et al.
2004). This is the first report of P. oxalidis from Panama.

Puccinia pelargonii-zonalis Doidge on Pelargonium hor-
torum L.H. Bailey. Panama, Chiriqui Province, Volcan, N
8° 46.421', W 82° 38.271', 25 Nov. 2004, leg. J.R. Hernandez

2004-095, II (BPI 864130).

P. pelargonii-zonalis was reported as cosmopolitan (Farr
et al. 2004). This is the first report of P. pelargonii-zonalis
from Panama.

Puccinia urbaniana Henn on Stachytarpheta cayennensis
(Rich.) Vahl. Panama, Chiriqui Province, Los Algarrobos,
8°29.164'N, 82°25.951'W, 27 Nov. 2004, leg. J.R. Hernandez

2004-136, III (BPI 864133).

P. urbaniana was reported from a number of countries in
the Americas (Farr et al. 2004). This is the first report of
P. urbaniana from Panama.

Pucciniastrum epilobii G.H. Otth on Fuchsia sp. Panama,
Chiriqui Province, Volcan, 8°46.421'N, 82°38.271'W, 25

Nov. 2004, leg. J.R. Hernandez 2004-096, II (BPI
864134).

P. epilobii was reported from the Americas, Asia, Europe,
and Oceania (Farr et al. 2004). This is the first report of P.
epilobii from Panama.

Uromyces commelinae Cooke on Commelina sp. Panama,
Chiriqui Province, Boquete, 8°46.691'N, 82°25.968'W, 26

Nov. 2004, leg. J.R. Hernandez and M. Vega Rios 2004-
121, II-III (BPI 864203).

U. commelinae was reported as cosmopolitan from the
Americas, Africa, Asia, and Oceania (Farr et al. 2004). This
is the first report of U. commelinae from Panama.

Uromyces dolicholi Arthur on Cajanus cajan (L.)
Millsp. Panama, Cocle Province, Aguadulce, 8°13.227N,

80°36.558'W, 24 Nov. 2004, leg. J.R. Hernandez 2004-081,

II (BPI 864136). Panama, Chiriqui Province, Caizan,

8°44.406'N, 82°49.348'W, 25 Nov. 2004, leg. J.R. Hernandez
2004-104, II (BPI 864137). On Rhynchosia sp. Panama,

Chiriqui Province, Siogui, 8°13.241'N, 81°44.758'W, 29 Nov.
2004, leg. J.R. Hernandez 2004-155, II (BPI 864140).

U. dolicholi was reported from the Caribbean, Central,
North and South America, Africa, and Asia (Farretal. 2004).
This is the first report of U. dolicholi from Panama.

Uromyces euphorbiae Cooke and Peck on Chamaesyce
sp. Panama, Cocle Province, Aguadulce, 8°13.227'N,

80°36.558'W, 24 Nov. 2004, leg. J.R. Hernandez 2004-083,
II-III(BPI864141).On Poinsettia heterophylla (L.) Klotzsch

and Garcke. Panama, Chiriqui Province, Distrito de Alanje,
La Barqueta, 8°18.258'N, 82°34.822'W, 27 Nov. 2004, leg. J.
R. Hernandez 2004-135, II (BPI 864142).

U. euphorbiae is cosmopolitan, reported as widespread in
the Americas, Africa, Asia, Europe, and Oceania on species of
Euphorbiaceae (Farr et al. 2004; Hernandez et al. 2005a, b).
This is the first report of U. euphorbiae from Panama.

Uromyces hedysari-paniculati (Schwein.) Farl. apud Ellis
on Desmodium sp. Panama, Chiriqui Province, Distrito de
Alanje, La Barqueta, 8°18.258'N, 82°34.822'W, 27 Nov.
2004, leg. J.R. Hernandez 2004-133, II (BPI 864143).

U. hedysari-paniculati has been reported as widespread
in the Caribbean, Central, North, and South America on
species of Desmodium and Meibomia (Fabaceae; Farr et al.
2004; Hernandez et al. 2005a, b). This is the first report of
U. hedysari-paniculati from Panama.

Uromyces novissimus Speg. on Cayaponia sp. Panama,

Chiriqui Province, Caizan, 8°44.406'N, 82°49.348'W, 25 Nov.

2004, leg. J.R. Hernandez 2004-106, II-III (BPI 864196).

U. novissimus was reported from the Caribbean, Central,
North, and South America (Farr et al. 2004). This is the
first report of U. novissimus from Panama.

Uropyxis diphysae (Arthur) Cummins on Diphysa amer-
icana (Mill.) M. Sousa. Panama, Chiriqui Province, Plaza

de Caizan, 8°43.528'N, 82°48.944'W, 25 Nov. 2004, leg. J.

R. Hernandez 2004-110, III (BPI 864148).

U. diphysae was reported on species of Diphysa from
Central America and Mexico (Farr et al. 2004). This is the
first report of U. diphysae from Panama.

New host records from Panama

Dietelia portoricensis (Whetzel and Olive) Buritica and
J.F. Hennen on Mikania micrantha Kunth. Panama, Chiriqui
Province, David, El Cabrero, UNACHI, 8°25.947'N,

82°27.045'W, 26 Nov. 2004, leg. J.R. Hernandez 2004-
127, III (BPI 864204). Panama, Quebrada Cermeno,
8°30.263'N, 82°26.342'W, 28 Nov. 2004, leg. J.R. Hernandez

and R. Mangelsdorff 2004-144, III (BPI 864208).

D. portoricensis was reported from Brazil, Costa Rica,
Colombia, Guatemala, Guyana, Honduras, El Salvador,
Puerto Rico, Trinidad and Tobago, and Venezuela (Buritica
and Hennen 1980; Evans and Ellison 2005; Farr et al. 2004;
Hernandezetal. 2005a, b). Ithas been reported from Panama
on Mikania scandens (L.) Willd. and Mikania sp. (Buritica
and Hennen 1980). This is the first report of D. porto-
ricensis on M. micrantha from Panama.

Springer

Mycol Progress (2007) 6:81 -91

Puccinia cnici-oleracei Pers. on Emilia fosbergii Nicolson.
Panama, Coclé Province, Aguadulce, 8°13.227'N,
80°36.558'W, 24 Nov. 2004, leg. J.R. Hernández 2004-
082, III (BPI 864118). Panama, Veraguas Province, La
Peña, 8°9.913'N, 81°5.069'W, 24 Nov. 2004, leg. J.R.
Hernández 2004-087, III (BPI 864119). On Melampo-

dium divaricatum (Rich.) DC. Panama, Coclé Province,

Aguadulce, 8°13.227'N, 80°36.558'W, 24 Nov. 2004, leg.

J.R. Hernández 2004-085, III (BPI864124). On Synedrella
nodiflora (L.) Gaertn. Panama, Chiriquí Province, David, El

Cabrero, UNACHI, 8°25.947'N, 82°27.045'W, 26 Nov.

2004, leg. J.R. Hernández 2004-129, III (BPI 864205).

P. cnici-oleracei is widespread in Africa, the Americas,
Asia, and Europe on several species in the Asteraceae (Farr
et al. 2004; Hernández et al. 2005a, b). It has been reported
from Panama on Neurolaena lobata (L.) R. Br. ex Cass. as
Puccinia synendrellae Henn. (Arthur 1916), as P. emiliae
Henn. (Standley 1927, 1933), and as P. melampodii Arthur
(Jackson 1922). This is the first report of P. cnici-oleracei
on E. fosbergii, M. divaricatum, and S. nodiflora from
Panama.

Puccinia thaliae Dietel on Canna sp. Panama, Chiriquí
Province, Volcán, 8°46.421'N, 82°38.271'W, 25 Nov. 2004,

leg. J.R. Hernández 2004-097, II-III (BPI 864132). On

Calathea sp. Panama, Veraguas Province, Los Ruices,
08°12.152'N, 81°30.521'W, 24 Nov. 2004, leg. J.R.
Hernández 2004-091, III. Panama, Chiriquí Province,

Quebrada Cermeño, 08°29.997'N, 82°26.266'W, 28 Nov.
2004, leg. J.R. Hernández 2004-141, II-III.

P. thaliae is widespread in the Americas as well as in
Africa and Asia, reported on species in the Marantaceae
and Cannaceae (Farr et al. 2004; Hernández et al. 2005a, b).
It has been reported from Panama on Calathea sp. (Kern
1938)asPuccinia cannae Henn. This is the firstreportof P.
thaliae on Canna sp. from Panama.

Uromyces appendiculatus (Pers.) Unger on Vigna ungui-
culata (L.) Walp. Panama, Chiriquí Province, Dominical,

8°39.825'N, 82°47.77'W, 25 Nov. 2004, leg J.R. Hernández

2004-115, II (BPI 864138).

U. appendiculatus is cosmopolitan, reported on a number
of species in the Fabaceae (Farr et al. 2004). It has been
reported from Panama on Macroptilium atropurpureum
(Sessé and Moc. ex DC.) Urb. (Lenné 1990, the firstspecific
mention of Panama, although her references only refer to
Central America). This is the first report of U. appendi-
culatus on V. unguiculata from Panama.

Uromyces neurocarpi Dietel on Clitoria falcata Lam.
Panama, Veraguas Province, El Piro, 8°12.516'N,

81°37.735'W, 24 Nov. 2004, leg. J.R. Hernández 2004-
093, II-III (BPI 864145).

U. neurocarpi is a neotropical rust reported on species of
Centrocema, Clitoria, Martiusia, and Neurocarpum from
the Caribbean, Central America, SouthAmerica, and Mexico
(Farr et al. 2004). It has been reported from Panama on
Clitoria sp. (Cummins 1978). This is the first report of
U. neurocarpi on C. falcata from Panama.

Uromyces setariae-italicae Yoshino on Melinis minutiflora

P. Beauv. Panama, Chiriqui Province, Volcan, N 8° 47.398',W
82° 40.565', 25 Nov. 2004, leg. J.R. Hernandez 2004-099,

II (BPI 864147). On Paspalum sp., same data, leg. J.R.

Hernandez 2004-099A, II.

U. setariae-italicae is cosmopolitan, reported as wide-
spread in Africa, the Americas, Asia, Europe, and Oceania
(Farr et al. 2004; Hernandez et al. 2005a, b). It has been
reported from Panama on Panicum maximum Jacq.
(McGuire and Crandall 1967) as Uromyces leptodermus
Syd. and P. Syd. This is the firstreportofU. setariae-italicae
on M. minutiflora and on Paspalum sp. from Panama.

Acknowledgment We thank Laurene Levy (USDA/APHIS) for the
molecular diagnosis of Phakopsora meibomiae. We also appreciate
the collaboration by Ralph Mangelsdorff (University of Frankfurt)
and Joe Kirkbride and John Wiersema (USDA/ARS) for identifying
many host species, Manfred Ruppel for assistance with the SEM, and
Roland Kirschner (University of Frankfurt) for help with the Latin
description. Carl Williams and Omar Monteros (Universidad de
Panama, Chiriqui) helped to visit fields of legume crops and Naira
Camacho (Direccion Salud Vegetal del Ministerio de Desarrollo
Agropecuario de Panama) assisted in obtaining permits. The trip by
JRH was made possible by Doug Luster (USDA/ARS) as part of the
survey for Phakopsora pachyrhizi in Central America and the
Caribbean. The activities by Meike Piepenbring and Maritza Vega Rios
were supported by the German Academic Exchange Service (DAAD), in
the context of a university partnership between the University of
Frankfurt and the Universidad Autonoma de Chiriqui, and by the
German Research Foundation (DFG).

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