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A contribution to the rust flora (Uredinales) of southern Africa, with an emphasis on South Africa

February 2005
Mycological Progress 4(1):55-75

February 2005
4(1):55-75

DOI:10.1007/s11557-006-0109-5

Authors:

Wolfgang Maier

Julius Kühn-Institut

Franz Oberwinkler

University of Tuebingen

Five new rust species are described and hitherto unknown spore states for the following seven species are reported: Puccinia desertorum on Evolvulus alsinoides, Uromyces comptus on Merremia bipinnatipartita, Puccinia halsei on Acacia hereroensis, Ravenelia transvaalensis on Acacia mellifera, Puccinia abutili on Abutilon angulatum, on Abutilon cf. austroafricanum, and on Abutilon cf. rehmannii, Puccinia lycii on Lycium sp. and Puccinia turgida on Lycium europaeum and on Lycium cf. oxycarpum. We also examined Uredo combreticola on Combretum cf. engleri, on Combretum hereroense, and on Combretum zeyheri, Puccinia afra on Lycium sp., and Uredopeltis cf. chevalieri on Grewia flavescens. All mentioned rust fungi are described in detail and are shown by line drawings. Selected species are illustrated with SEM-photographs.

Puccinia namibiana on Blepharis obmitrata. Peridium cells, aecio-and teliospores (Holotype NA 216). Scale bar = 10 µm.

…

Puccinia desertorum on Evolvulus alsinoides. Peridium cells and aeciospores (NA 172), uredinio-and teliospores (NA 276). Scale bar = 10 µm.

…

Uromyces otaviensis on cf. Ipomoea verbascoidea. Peridium cells, aecio-and teliospores (Holotype NA 305). Scale bar = 10 µm.

…

Uromyces comptus on Merremia bipinnatipartita. Peridium cells, aecio-, uredinio-, and teliospores (NA 221). Scale bar = 10 µm.

…

Puccinia windhoekensis on Coccinia rehmannii. Peridium cells, aecio-, uredinio-, and teliospores (Holotype NA 152). Scale bar = 10 µm.

…

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ntil now the rust flora of Namibia has not been ex-

amined systematically. As far as could be estab-

lished only 13 rust species from Namibia have been

recorded so far (HENNINGS 1898; DIETEL in SYDOW & SYDOW

1904; SYDOW & SYDOW 1912b, 1915, 1924; POLE EVANS

1915; SYDOW 1928; DOIDGE 1939, 1950; GJÆRUM 1988). If

this number is compared to the circa 4.200 known higher plant

species (C

RAVEN 1999) it is clear that many more rust species

await to be found: for example, Austria has 491 rust species

(ZWETKO

2000) and about 3.300 higher plant taxa (A

DLER et

al. 1994), corresponding to a ratio of circa 15 %, thus, rein-

forcing the observations of H

ENNEN & MCCAIN (1993) that

rust species are about 5–25 % as numerous as plant species.

As a consequence, a total rust flora between 200 to 1.000

species could be expected for Namibia.

A first approach to complete the Namibian rust flora was

made during the pilot phase of the BIOTA (Biodiversity Moni-

toring Transect Analyses) Southern Africa project: rust fungi

were collected extensively in Namibia and in the western parts

of South Africa. In the present study we report observations

made predominantly in Namibia with superior taxonomic in-

terest like new rust species, new spore states of known rust

fungi and records of species from which until now only the

type collections were known. The complete species list from

Namibia with more than 60 species will be soon published as

‘Preliminary flora of rust fungi in Namibia’ in the Namibian

botanical journal ‘Dinteria’.

Material and methods

Most of the specimens were collected in Namibia by the first

author between 1 March 2002 and 3 May 2002. The specimens

collected in the course of this study as well as some additional

specimens from herbaria were studied from freehand sections

and scrape mounts of infected plant material. The samples

were heated in “Hoyer’s fluid” (C

UNNINGHAM 1972) and sub-

sequently examined with a Carl Zeiss microscope with bright

field and phase contrast optics. For scanning electron micro-

scopy, air-dried fragments of rust-infected leaves were put

onto double-sided adhesive tape fixed to an observation stub,

spattered with gold-palladium and observed with a Cambridge

Stereoscan 250 Mk-2.

With the exception of pycnio- and basidiospores, 25 spores

of each occurring spore states were measured. If only one col-

lection of a rust species was available or in the case of type

collections 50 spores of each present spore state were mea-

sured. If a spore state is put in parentheses, measurements were

A contribution to the rust flora (Uredinales) of southern Africa, with

an emphasis on Namibia

Mechthilde

MENNICKEN

1,+

, Wolfgang MAIER

and Franz OBERWINKLER

Five new rust species are described and hitherto unknown spore states for the following seven species are reported: Puc-

cinia desertorum on Evolvulus alsinoides, Uromyces comptus on Merremia bipinnatipartita, Puccinia halsei on Acacia

hereroensis, Ravenelia transvaalensis on Acacia mellifera, Puccinia abutili on Abutilon angulatum, on Abutilon cf. austro-

africanum, and on Abutilon cf. rehmannii, Puccinia lycii on Lycium sp. and Puccinia turgida on Lycium europaeum and

on Lycium cf. oxycarpum. We also examined Uredo combreticola on Combretum cf. engleri, on Combretum hereroense,

and on Combretum zeyheri, Puccinia afra on Lycium sp., and Uredopeltis cf. chevalieri on Grewia flavescens. All men-

tioned rust fungi are described in detail and are shown by line drawings. Selected species are illustrated with SEM-

photographs.

Taxonomical novelties: Puccinia namibiana Mennicken, Maier & Oberw. (Anamorph Aecidium acanthopsidis Henn.) on Blepharis ob-

mitrata, Uromyces otaviensis Mennicken, Maier & Oberw. on cf. Ipomoea verbascoidea, Puccinia windhoekensis Mennicken, Maier

& Oberw. on Coccinia rehmannii, Puccinia ovamboensis Mennicken, Maier & Oberw. on Triaspis hypericoides, and Uredopeltis flavae

Mennicken, Maier & Oberw. on Grewia flava.

Keywords: Biodiversity, rust fungi, South Africa, taxonomy

Mycological Progress 4(1): 55–75, February 2005 55

Lehrstuhl für Spezielle Botanik & Mykologie, Botanisches Institut,

Universität Tübingen, Auf der Morgenstelle 1, D-72076 Tübingen,

Germany; e-mail: info@systbot.uni-tuebingen.de.

Corresponding author.

* Part 220 in the series “Studies in Heterobasidiomycetes” from the

Botanical Institute, University of Tübingen

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

not available; the reasons for that are either that there were too

small a number of spores or that there were strong alterations

of spores which had already germinated.

Peridium cells are presented with the outer wall of the cells

on the left hand side of the line drawing, and the inner wall on

the right hand side. All specifications with the prefix ‘circa’

are based on measurements of less than 25 cells.

The morphological types of anamorphic states follow

UMMINS & HIRATSUKA (2003).

The terms to describe the teliospores of the genus Rave-

nelia follow H

ERNÁNDEZ & HENNEN (2002). They “consider

the ‘spore head’ to be a single teliospore composed of many

pigmented probasidial cells. Hyaline, hygroscopic cysts occur

on the abaxial surface of the teliospore and sterile intercalary

cells are sometimes present between the probasidial cells and

the cysts.” The description of the shape of the teliospores, like

‘convex above, even below’, is restricted to the pigmented

probasidial cells.

For the determination of the rust fungi the encyclopaedic

publications of D

OIDGE (1927, 1928, 1939, 1941, 1948a,

1948b, 1950) were the main source of information. In addi-

tion, we checked many other publications concerning the rust

flora of Africa, which are available via internet under http://

www.mycology.uni-tuebingen.de/databases/rust-literature/.

Results and discussion

The rust species are listed in alphabetical order referring to the

host family and the host genus, respectively. The following

abbreviations are used: 0 = pycnia = spermogonia; I = aecia;

II = uredinia; III = telia, IV = basidia.

Acanthaceae – Blepharis

Puccinia namibiana Mennicken, Maier & Oberw. sp.

nov.

Fig. 1

Anamorph. Aecidium acanthopsidis Syd. & P. Syd. (SYDOW

YDOW 1915: 36). Type on cf. Blepharis obmitrata C.B. Clarke (as

Acanthopsis sp. juv.). Namibia, Okahandja, 10. 2. 1911, leg. Kurt

Dinter No. 2530 (B 70 0007226).

Pycnia rara, amphigena in foliis, typo 4. Aecia aecidioidea, amphi-

gena, laxe aggregata. Cupulae conicae, usque ad 600 µm longae,

circa 300–400 µm diam. Peridia cremeo-alba, recurvata, irregulari-

ter lacerata, cellulis peridii firme conjunctis, pariete exteriore striato,

circa 9–13 µm crasso, pariete interiore verruculoso, circa 3–5 µm

crasso. Aeciosporae angulato-globoideae vel ellipsoideae, 17–24

(26) × 13–19 µm, pariete irregulariter verruculoso, hyalino, 0.5–1.5 µm

crasso. Uredinia ignota. Telia amphigena, subepidermalia, atro-brun-

nea vel nigra, usque ad 4 mm diam., epidermide tecta, loculata pa-

raphysibus castaneis. Teliosporae bicellulares, cylindraceae, clavatae

vel fusiformes, apice obtuso vel subacuminato, base attenuata, medio

leviter constrictae, (38) 43–61 (67) × 13–20 (23) µm, pariete laevi-

gato, lateraliter 1–2 µm crasso, apicaliter 4–8 µm crasso, pedicello

fusco vel brunneo, usque ad 95 µm longo. Mesosporae et teleuto-

sporae tricellulares adsunt.

In foliis Blepharidis obmitratae (Acanthaceae).

Pycnia rarely present, amphigenous on leaves, subepidermal,

type 4.

Aecia Aecidium-type, amphigenous on leaves, densely scat-

tered in roundish or irregular groups up to 4 mm diameter on

slightly brightened spots, which tend to be hypertrophy in the

range of leaf veins, groups separate or admixed with the telia,

aecial cups conical, long remaining closed, erumpent, circa

300–400 µm diameter, up to 600 µm tall, spore mass yellow-

ish-white or creamish-white, surrounded by finely recurved,

irregularly lacerated, creamish-white peridium, cells of the

peridium firmly connected, outer wall finely striate, circa

9–13 µm, inner wall verrucous, circa 3–5 µm. Aeciospores

angular globoid to ellipsoid, 17–24 (26) × 13–19 µm, verru-

cous, warts irregularly developed, from delicate and even to

coarse and irregular, spore wall 0.5–1.5 µm, hyaline, germ

pores inconspicuous.

Uredinia unknown.

Telia amphigenous on leaves, subepidermal, dark chocolate-

brown to blackish pustules up to 4 mm, roundish or irregular

in outline, separate or admixed with the aecia, covered by the

epidermis, clearly margined on the host leaves, strongly locu-

late with chestnut-brown, thin paraphyses, interspaces up to

120 µm wide and circa 160 µm high, small yellowish-brown

spots surrounding the telia, not clearly margined on the host

Fig. 1: Puccinia namibiana on Blepharis obmitrata. Peridium

cells, aecio- and teliospores (Holotype NA 216). Scale bar =

10 µm.

Mycological Progress 4(1) / 2005

leaves. Teliospores in general bicellular, cylindrical, clavate,

or ± fusiforme, obtuse or subacuminate at the apex, attenu-

ate at the base, slightly constricted at the septum, (38) 43–61

(67) × 13–20 (23) µm, wall smooth, yellowish-brown, about

1–2 µm at the sides, about 4–8 µm thick at the apex, germ

pores obscure, pedicel up to 95 µm long, persistent, partly col-

lapsing, yellowish-brown. Mesospores and tricellular spores

existing.

On the leaves of Blepharis obmitrata (Acanthaceae).

Etymology. Named after the country of occurrence, Namibia.

Specimens examined:

On Blepharis obmitrata. Namibia, B 2 Okahandja - Wilhelmstal,

21°56’44.6’’S, 16°43’18.5’’E, 1.484 m asl., 6. 4. 2002, leg. M. Men-

nicken No. NA 216, I III. Holotype (PREM), Isotype (WIND). – On

cf. Blepharis obmitrata. Namibia, Oshikoto, D 3031, C 39 - Lake

Guinas, 19°25’45.2’’S, 17°24’16.8’’E, 1.362 m asl., 10. 4. 2002, leg.

M. Mennicken No. NA 241, I III. Paratypes (PREM, WIND). – On

cf. Blepharis obmitrata (as Acanthopsis sp. juv.). Namibia, Oka-

handja, Akazienbuschsteppe, 10. 2. 1911, leg. Kurt Dinter No. 2530,

I. Type of Aecidium acanthopsidis (B 70 0007226). – On Blepharis

obmitrata. Namibia, B 2 Karibib - 100 - Okahandja, 21°57’22.4’’S,

15°57’31.5’’E, 1.319 m asl., 6. 4. 2002, leg. M. Mennicken No. NA

144, 0 I. (PREM, WIND). – On Blepharis obmitrata. Namibia,

Windhoek, Love Street, 22°33’48.9’’S, 17°05’16.8’’E, 1.736 m asl.,

5. 4. 2002, leg. M. Mennicken No. NA 161, 0 I. (PREM, WIND).

After LAUNDON (1963a) the aecia differ from all other rust

fungi on Acanthaceae in having peridial cells with very thick

outer walls.

Combretaceae – Combretum

Uredo combreticola Doidge

(DOIDGE 1939: 506). Fig. 2

Type on Combretum zeyheri Sond. South Africa, Transvaal, Bar-

berton, Nelspruit, Research Station, 11. 9. 1931, leg. Liebenberg No.

2777 (PREM 26038).

Pycnia, aecia, and telia unknown.

Uredinia Uredo-type, amphigenous on leaves, subepidermal,

scattered, occasionally covering the whole leaf surface, irregu-

larly rupturing pustules up to 0.5 mm diameter, generally

smaller, not clearly margined on the host leaves, roundish or

irregular in outline, elongated in a tip, brown, powdered by

the pale brownish urediniospores, spots surrounding pustules

inconspicuous, slightly brightened, not clearly margined on

the host leaves, with an undefined layer of presumably fungal

cells between epidermis and urediniospores. Paraphyse-like

structures hyaline, short, uniformly thin-walled. Uredinio-

spores borne singly on pedicels, subgloboid, ovoid to elliptic,

15–25 (29) × 13–21 µm, spore wall uniformly about 1–1.5

(2) µm wide, pale golden to hyaline, echinulate, germ pores

rather obscure, occasionally indicated in optical sections, pro-

bably numerous and scattered, without papillae.

Specimens examined:

On Combretum zeyheri Sond. South Africa, Transvaal, Barberton,

Nelspruit, Research Station, 11. 9. 1931, leg. Liebenberg No. 2777,

II. Type (PREM 26038). – On Combretum cf. engleri Schinz.

Namibia, Koukuas, 18°54’16.7’’S, 18°17’14.6’’E, 1.197 m asl.,

14. 4. 2002, leg. M. Mennicken No. NA 265, II. (PREM, WIND).

– On Combretum hereroense Schinz. Namibia, D 3001, B 1 -

Tsintsabis, 18°43’55.1’’S, 17°34’54.1’’ E, 1.167 m asl., 13. 4. 2002,

leg. M. Mennicken No. NA 256, II. (PREM, WIND). – On Com-

bretum hereroense. Namibia, D 2860, B 8 - Hoba Meteorite,

19°37’43.7’’S, 17°56’39.0’’E, 1.518 m asl., 16. 4. 2002, leg. M.

Mennicken No. NA 289, II. (PREM, WIND).

CROUS et al. (2000) have not cited this rust for South Africa.

Uredo combreticola is new to the rust flora of Namibia, Com-

bretum hereroense and Combretum cf. engleri seem to be new

host plants.

Before examining the type collection we were in doubt

about our collections belonging to Uredo combreticola, be-

cause D

OIDGE (1939) stated thinner urediniospores (17–23 ×

10–15 µm) with verrucous spore walls, and did not mention

paraphyses. Our collections conform well with the type col-

lection. It could not be determined whether the uniformly thin-

walled ‘paraphyse-like structures’ are paraphyses or deformed

pedicels after losing the urediniospores.

Following three other species of rust fungi are known with

urediniospores on Combretum species:

Uredo longaensis Henn. (H

ENNINGS 1903), which is de-

scribed on Combretum baumii Engl. & Gilg. in Angola. It dif-

fers in having catenulate urediniospores with apically thickened

spore walls (V

IENNOT-BOURGIN 1958).

Cerotelium combreti Cummins (CUMMINS 1952) is de-

scribed on Combretum sp. from Uganda and differs in having

paraphyses with externally thickened walls.

Uredo combreti F. Kern & Thurst. (K

ERN & THURSTON

1944) on Combretum fruticosum (Loefl.) Stuntz from Vene-

zuela has much bigger urediniospores (18–23 × 26–35 µm).

Convolvulaceae - Evolvulus

Puccinia desertorum Syd. & P. Syd. (S

YDOW & SYDOW

1911: 259). Fig. 3

Type on Evolvulus alsinoides (L.) L. Namibia, Okahandja, 1.200 m

asl., 26. 5. 1907, leg. Kurt Dinter No. 570 (B 70 0007270).

Pycnia rarely developed, adaxial on leaves, opposite to or ad-

mixed with the aecia, type 4.

Aecia Aecidium-type, amphigenous on leaves, densely scat-

tered in small and roundish, partially confluent, groups up to

3 mm diameter on slightly brightened and yellowed leaf spots,

Fig. 2: Uredo combreticola on Combretum hereroense. Pa-

raphyse-like structures and urediniospores (NA 256). Scale

bar = 10 µm.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

groups often admixed with the uredinia, aecial cups small con-

ical to cylindrical, erumpent, circa 100–200 µm diameter, up

to circa 300 µm tall, spore mass yellowish-white or white, sur-

rounded by recurved, irregularly lacerated, whitish peridium,

cells of the peridium firmly connected, outer wall finely striate,

3–8 µm, inner wall verrucous, 3–5 µm. Aeciospores angular

globoid to ellipsoid, 15–21 × 12–19 µm, delicately verrucous,

spore wall uniformely 0.5–1 µm, hyaline, germ pores incon-

spicuous.

Uredinia Uredo-type, amphigenous on leaves, rarely cauli-

colous, subepidermal, small pustules, ± roundish, up to 0.5 mm

wide, separate, scattered, or in irregular or concentric groups,

cinnamon-brown, exposed and surrounded by the torn epi-

dermis. Spots surrounding partially the uredinia up to 2 mm

wide, yellow, brown to pale yellowish-green, not clearly mar-

gined on the host leaves. The whole leaves are yellowing,

later browning in the case of heavy infection. Urediniospores

obovoid, subglobose to ellipsoid, 19–28 × 15–23 µm, echinu-

late, spore wall uniformly about (1) 1.5–2.5(3) µm thick, yel-

lowish-brown, germ pores conspicuous, 2 (to 3), equatorial,

with medium developed hyaline papillae.

Telia amphigenous on leaves, rarely caulicolous, subepider-

mal, small pustules up to 0.5 mm, roundish or irregular in out-

line, separate, scattered, or in irregular or concentrical groups,

partially replacing the uredinia, black, exposed and surrounded

by the torn epidermis. Spots surrounding partially the telia up

to 2 mm wide, yellow to pale yellowish-green, not clearly mar-

gined on the host leaves. Teliospores bicellular, ellipsoid,

clavate to oblong, flattened, rounded or acuminate at the apex,

attenuate to subattenuate at the base, medium constricted at

the septum, 38–54 (56) × 18–25 µm, spore wall smooth, about

1–2.5 µm thick at the sides, up to 13 (15) µm thick at the apex,

brown, apical brightened, germ pores apical and just below

the septum, pedicel up to 55 µm, thick-walled, not collapsing

in fresh material, yellowish to hyaline.

Specimens examined:

On Evolvulus alsinoides. Namibia, Okahandja, 1.200 m asl., 26. 5. 1907,

leg. Kurt Dinter No. 570, II III. Type (B 70 0007270). – On Evolvulus

alsinoides. Namibia, BIOTA-observatory at Sonop 903 Research Sta-

tion, 19°04’51.9’’S, 18°54’47.4’’E, 5. 3. 2002, leg. M. Mennicken No.

NA 119, II. (PREM, WIND). – On Evolvulus alsinoides. Nami-bia, D

3016, 18°51’22.5”S, 18°28’49.2’’E, 1.204 m asl., 25. 4. 2002, leg. M.

Mennicken No. NA 276, II III. (PREM, WIND). – On Evolvulus alsi-

noides. Namibia, Ombeameiata, house camp, 21°25’50.1’’S,

18°01’22.6’’E, 1.606 m asl., 2. 4. 2002, leg. M. Mennicken No. NA

185, 0 I II. (PREM, WIND). – On Evolvulus alsinoides. Namibia, Stein-

hausen, camp WNM of the farmhouse, 21°48’50.3’’S, 18°13’07.1’’E,

1.661 m asl., 1. 4. 2002, leg. M. Mennicken No. NA 172, 0 I II III.

(PREM, WIND). – On Evolvulus alsinoides. Namibia, Windhoek, wa-

ter tower mountain, 22°33’58.2’’S, 17°05’39.5’’E, 1.759 m asl.,

5. 4. 2002, leg. M. Mennicken No. NA 157, II. (PREM, WIND).

Puccinia desertorum seems to be only known from the Nami-

bian type collection, from Madras, India (R

AMAKRISHNAN

al. 1953), and from Kenya (G

JÆRUM

1986). Until now only

uredinia and telia have been found. Therefore, pycnia and ae-

cia of Puccinia desertorum are described as new.

Convolvulaceae – Ipomoea

Uromyces otaviensis Mennicken, Maier & Oberw. sp.

nov.

Figs. 4 (Plate 1), 5

Pycnia et uredinia ignota. Aecia aecidioidea, amphigena, cupulata.

Cupulae circa 150–550 µm diam., usque ad 750 µm altae. Peridia

alba, recurvata, irregulariter lacerata, cellulis peridii firme conjuncta,

pariete exteriore laevi, circa 4–8 µm crasso, pariete interiore verru-

culoso, circa 8–12 µm crasso. Aeciosporae angulatae globoideae vel

ellipsoideae, 23–34 × 17–27 µm, pariete verruculoso, hyalino,

1.5–3 µm crasso. Telia amphigena, subepidermalia, nigra, mox nuda,

pulverulenta. Teliosporae unicellulares, dimorphae, juniores ap-

planato-globoideae, castaneae, 21–29 × 27–33 µm, vetustiores pilea-

tae, atro-castaneae vel nigrae, 14–18 × 24–28 µm, pariete irregulari-

ter corrugato, lateraliter 2–3.5 µm crasso, apicaliter usque ad 10 µm

crasso, bilaminato, vetustiorum sporarum strato exteriore incon-

spicuo, pedicello hyalino, usque ad 50 µm longo, inflato.

In foliis cf. Ipomoeae verbascoideae Choisy (Convolvulaceae).

Pycnia unknown.

Aecia Aecidium-type, amphigenous on leaves, subepidermal,

scattered in small and roundish, partly confluent groups up to

Fig. 3: Puccinia desertorum on Evolvulus alsinoides. Peridium

cells and aeciospores (NA 172), uredinio- and teliospores (NA

276). Scale bar = 10 µm.

Mycological Progress 4(1) / 2005

3 mm diameter, on slightly brightened and yellowed leaf spots,

groups often admixed with the telia, aecial cups erumpent, cir-

ca 150–550 µm diameter, up to circa 700 µm tall, spore mass

yellowish-white or white, surrounded by recurved, irregularly

lacerated, whitish peridium, cells of the peridium firmly con-

nected, outer wall smooth, circa 4–8 µm thick, inner wall ver-

rucous, circa 8–12 µm thick. Aeciospores angular globoid to

ellipsoid, 23–34 × 17–27 µm, regularly verrucous, spore wall

1.5–3 µm, hyaline, germ pores inconspicuous.

Uredinia and urediniospores unknown.

Telia amphigenous on leaves, subepidermal, forming small

patches up to 1 mm wide, exposed, pulverulent, roundish, se-

parate or scattered, often associated with the aecia, black, spots

surrounding the telia up to 2 mm wide, yellow to yellowish-

green, not clearly margined on the host leaves. Teliospores

unicellular, tending to be dimorphic depending on maturity,

younger spores flattened globoid, chestnut-brown, 21–29 ×

27–33 µm, full-grown spores collapsed on the bottom side and

pileolus-shaped, 14–18 × 24–28 µm, dark chestnut-brown to

blackish, spore wall bilaminate, about 2–3.5 µm thick at the

sides, up to 10 µm thick at the apex, outer layer irregularly

wrinkled, forming a broad and flattened papilla over the api-

cal germ pore, in young spores recognisable as a separate lay-

er and swellable in heated lactic acid up to 1–3 µm, golden, in

full-grown spores not or uncontinuous recognisable as a sepa-

rate layer and not swellable, dark chestnut-brown to blackish,

inner layer chestnut-brown to blackish, probably smooth, con-

tinuously thickened from the base to the apex, pedicel up to

50 µm long, not collapsing, usually with spherical swelling

short below the attachment, hyaline.

On the leaves of cf. Ipomoea verbascoidea (Convolvulaceae).

Etymology. Named after the habitat in Namibia, Otavi Moun-

tains.

Specimen examined:

On cf. Ipomoea verbascoidea. Namibia, Otavi Mountains, D 2863,

19°33’30.1’’S, 17°44’37.6’’E, 1.757 m asl., 17. 4. 2002, leg. M.

Mennicken No. NA 305, I III. Holotype (PREM), Isotype (WIND).

A similar rust fungus Uromyces ipomoeae (Thüm.) Berk. was

found on three Ipomoea species in Eastern Cape, Mpumalan-

ga, and North-West Province in South Africa (C

ROUS et al.

2000). It differs from Uromyces otaviensis in having an “epi-

spore … with regular longitudinal striae which radiate from

the apex and converge towards the base” as well as in having

smaller (20–25 µm), and thinner-walled (about 1 µm thick)

aeciospores (D

OIDGE 1927).

Uromyces pieningii Cummins (C

UMMINS, 1960: 45), which

is described on Ipomoea argentaurea Hall. f. from Ghana, dif-

fers from Uromyces otaviensis mainly in having thin-walled

peridial cells (inner wall 3 µm thick, outer wall 3–4 µm thick)

and in having smaller (23–27 × 15–20 µm), thin-walled

(1 µm) aeciospores.

Convulvulaceae – Merremia

Uromyces comptus Syd. & P. Syd.

(SYDOW & SYDOW

1911: 259). Fig. 6

Type on Merremia bipinnatipartita (Engl.) Hallier f. (= Ipomoea bi-

pinnatipartita Engl.) Namibia, Okahandja, Wilhelmstal, 1.300 m asl.,

19. 5. 1907, leg. Kurt Dinter No. 562. (B 70 0007272).

Pycnia (type 4) admixed with aecia.

Aecia Aecidium-type, abaxial on leaves, beginning in the area

of veins, causing slightly hypertrophy deformity, densely scat-

tered in longitudinal groups up to 1.5 cm, aecial cups cylin-

drical, erumpent, circa 200–300 µm diameter, up to 500 µm

tall, spore mass white, surrounded by recurved, irregularly and

deeply lacerated, white peridium, cells of the peridium loosely

connected, outer wall smooth, 2–3 µm, inner wall coarsely

verrucous, 3–5 µm. Aeciospores angular subgloboid, ovoid

to ellipsoid, 18–25 × 15–21 µm, delicately verrucous, spore

wall 0.5–1.5 µm, hyaline, germ pores inconspicuous.

Uredinia Uredo-type, amphigenous on leaves, separate or

scattered, subepidermal, irregularly rupturing pustules small,

up to 0.8 mm wide, roundish, cinnamon-brown, early ex-

posed, pulverulent, surrounded by the torn epidermis, with-

out spots surrounding pustules. Urediniospores globoid,

ovoid, or ellipsoid, partly angular, 22–32 × 18–24 µm, echi-

nulate, spore wall uniformly 1.5–2.5 µm thick, brown, germ

Fig. 5. Uromyces otaviensis on cf. Ipomoea verbascoidea.

Peridium cells, aecio- and teliospores (Holotype NA 305).

Scale bar = 10 µm.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

pores conspicuous, 2 to 3, ± equatorial, papillae very weakly

developed.

Telia amphigenous on leaves, developing from the uredinia,

separate or scattered, subepidermal, irregularly rupturing pus-

tules small, up to 0.8 mm wide, pulvinate, roundish, black,

early exposed, surrounded by the torn epidermis, without spots

surrounding pustules. Teliospores unicellular, globoid to ovoid,

chestnut-brown, 28–38 × 25–30 µm, spore wall smooth, about

4–6 µm thick at the sides, up to 12 µm thick at the apex, germ

pore apical, with a broad brownish-yellow papilla, pedicel up

to 120 µm long, thick-walled, not collapsing, hyaline.

Specimens examined:

On Merremia bipinnatipartita. Namibia, Okahandja, Wilhelmstal,

auf Lateritflächen, 1.300 m asl., 19. 5. 1907, leg. Kurt Dinter No. 562,

II III. Type (B 70 0007272). – On Merremia bipinnatipartita. Namib-

ia, C 36 Wilhelmstal - Omaruru, behind D 2110, 21°34’44.9’’S,

16°08’00.0’’E, 1.314 m asl., 6. 4. 2002, leg. M. Mennicken No. NA

221, 0 I II III. (PREM, WIND).

In the Namibian type collection of Uromyces comptus only

uredinia and telia could be found. The only other record of

Uromyces comptus is known from the same host plant from

Tanzania (W

ATSON 1971): “Dark-brown to black rust pustules

on leaves of Ipomoea bipinnati-partita”. Therefore, pycnia

and aecia are newly described here.

CUCURBITACEAE – COCCINIA

Puccinia windhoekensis Mennicken, Maier & Oberw.

sp. nov.

Figs. 7 (Plate 1), 8

Pycnia rara, amphigena, typo 4. Aecia caeomatoidea, in foliis am-

phigena vel caulem deformantia, late aggregata, circa 0.3–0.8 mm

diam., sine peridia, ochracea, pulverulenta. Aeciosporae catenulatae,

subgloboideae, ovoideae, ellipsoideae, oblongae vel pyriformes, par-

tim angulatae, 21–36 × 17–27 µm, pariete verruculoso, lateraliter

1–1.5 µm crasso, apicaliter 2–6 (8) µm crasso, hyalino, 2–4 (5) poris

germinationis dispersis. Uredinia caulicola vel in foliis amphigena,

subepidermalia, usque ad 1 mm diam., cinnamomea, mox nuda, pul-

verulenta. Urediniosporae globoideae, ovoideae vel subellipsoideae,

23–35 × 17–24 µm, pariete echinulato, lateraliter 1–2 µm crasso, api-

caliter 1.5–2 (2.5) µm crasso, castaneo, 2 poris germinationis ae-

quatorialibus. Telia caulicola vel in foliis amphigena, subepidermalia,

atro-brunnea vel nigra, usque ad 1 mm diam., mox nuda, pulveru-

lenta. Teliosporae bicellulares, subgloboideae, ovoideae vel ellipsoi-

deae, cylindraceae, subclavatae vel irregulares, apice rotundato, ap-

planato vel mucronato, base rotundata vel leviter attenuata, medio

haud vel leviter constrictae, 35–48 × 26–36 µm, pariete irregulariter

corrugato vel rugoso, lateraliter 3–7 µm crasso, apicaliter 3 to 10 µm

crasso (apiculo incl.), bilaminato, strato exteriore aureo-fulvo, strato

interiore atro-castaneo, poris germinationis in quaque cellula 2, juxta

septum positis, pedicello aureo vel hyalino, usque ad 85 µm longo.

Mesosporae adsunt.

In foliis caulisque Cocciniae rehmannii Cogn. (Cucurbitaceae).

Pycnia rarely present, amphigenous on leaves, subepidermal,

type 4.

Aecia Caeoma-type, caulicolous, causing slight hypertrophy

of the stem similar to witches’ brooms, amphigenous on leaves,

subepidermal, widely scattered in groups, groups about a few

millimetres to several centimetres wide, without surrounding

leaf spots, erumpent, without peridium, circa 0.3–0.8 mm dia-

meter, roundish to ellipsoid, spore mass ochraceous, pulve-

rulent, surrounded by the convex epidermis, partly admixed

with uredinia and telia. Aeciospores catenulate, variable in

shape, subgloboid, ovoid, ellipsoid, oblong, pear- or lemon-

shaped, partly angular, 21–36 × 17–27 µm, delicately verru-

cous, spore wall variably thick, about 1–1.5 µm thick at the

sides, about 2–6 (8) µm thick at the apex, about 1.5–3 µm thick

at the base, hyaline, germ pores inconspicuous, 2 to 4 (to 5),

scattered.

Fig. 6: Uromyces comptus on Merremia bipinnatipartita.

Peridium cells, aecio-, uredinio-, and teliospores (NA 221).

Scale bar = 10 µm.

Fig. 8: Puccinia windhoekensis on Coccinia rehmannii. Peri-

dium cells, aecio-, uredinio-, and teliospores (Holotype NA

152). Scale bar = 10 µm.

Mycological Progress 4(1) / 2005

Plate 1. Fig. 4. Uromyces otaviensis on cf. Ipomoea verbascoidea. Teliospores (Holotype NA 305). SEM. Scale bar = 10 µm.

Fig. 7. Puccinia windhoekensis on Coccinia rehmannii. Teliospores (Holotype NA 152). SEM. Scale bar = 10 µm. Fig. 11.

Puccinia ovamboensis on Triaspis hypericoides ssp. nelsonii. Teliospores (Holotype NA 292). SEM. Scale bar = 10 µm. Fig.

13. Puccinia abutili on Abutilon cf. austro-africanum. Teliospores (NA 195). SEM. Scale bar = 10 µm.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

Uredinia Uredo-type, caulicolous or amphigenous on leaves,

separate, scattered, and in irregular or ± concentric groups,

subepidermal, irregularly rupturing pustules small, about

0.1–1 mm wide, roundish, cinnamon-brown, early exposed,

pulverulent, surrounded by the torn epidermis, without spots

surrounding pustules, partly admixed with aecia and telia. Ure-

diniospores borne singly on pedicels, globoid, ovoid, or sub-

ellipsoid, 23–35 × 17–24 µm, finely echinulate, spore wall

about 1–2 µm thick at the sides and about 1.5–2 (2.5) µm thick

at the apex, medium chestnut-brown, basally paler, germ pores

conspicuous, 2, equatorial, with hyaline papillae that are weak-

ly developed.

Telia caulicolous or amphigenous on leaves, (partly) deve-

loping from the uredinia, separate, scattered, and in irregular

or ± concentric groups, partly confluent, subepidermal, irregu-

larly rupturing pustules small, about 0.1–1 mm wide, roundish,

ellipsoid, oblong or irregular in outline, dark chocolate-brown

to black, early exposed, pulverulent, surrounded by the torn

epidermis, without spots surrounding pustules, partly admixed

with aecia and uredinia. Teliospores in general bicellular, sub-

globoid, ovoid to ellipsoid, rounded, flattened, or apiculate

through an apicule at the apex, rounded or slightly attenuate

at the base, not or very slightly constricted at the septum,

chestnut-brown, 35–48 × 26–36 µm, spore wall about 3–7 µm

thick at the sides, about 3–10 µm thick at the apex (with

apicule), bilaminate, outer layer thin, golden-brown, irregular-

ly corrugated-wrinkled, forming a broad and flattened papilla

over the lateral germ pores, inner layer thick, dark chestnut-

brown, probably smooth, 2 germ pores per cell short above

and below the septum, inconspicuous, pedicel up to 85 µm

long, thick-walled, generally not collapsing, yellowish to hya-

line. Mesospores rarely existing.

On the leaves of Coccinia rehmannii (Cucurbitaceae).

Etymology. Named after the habitat, the capital city of Nami-

bia, Windhoek.

Specimens examined:

On Coccinia rehmannii. Namibia, Windhoek, Sinclairstreet,

22°33’38.9’’S, 17°05’19.6’’E, 1.731 m asl., 5. 4. 2002, leg. M. Men-

nicken No. NA 152, 0 I II III. Holotype (PREM), Isotype (WIND).

– On Coccinia rehmannii. Namibia, Windhoek, Botanical Garden,

22°34’17.5’’S, 17°05’36.7’’E, 1.770 m asl., 5. 4. 2002, leg. M. Men-

nicken No. NA 163, 0 I II III. Paratypes (PREM, WIND).

The only known autoecious and macrocyclic rust fungus on

Cucurbitaceae is Puccinia cephalandrae Thüm. (T

HÜMEN 1876:

425). After S

YDOW & S

YDOW (1904) Puccinia cephalandrae

is synonymous with Uredo cephalandrae Thüm. (THÜMEN

1878: 355), Uredo dolichospora Kalchbr., and Aecidium

cephalandrae Cooke (C

OOKE 1884: 6). DOIDGE (1927) has

given a detailed description of Puccinia cephalandrae, which

differs from Puccinia windhoekensis in having aecidioid aecia

with a recurved peridium and in the much longer urediniospores

(33–53 × 16–20 µm). D

OIDGE (1927), additionally listed Puc-

cinia momordicae Kalchbr. & Cooke in K

ALCHBRENNER (1882:

24), and Puccinia trochomeriae Cooke (C

OOKE 1882: 125) as

synonyms of Puccinia cephalandrae, and Cephalandra, Coc-

cinia, Cucumis, Kedrostis, Momordica, Trochomeria, and

Zehneria as host genera. We examined teliospores from a col-

lection of Puccinia cephalandrae (B 70 0008488) and from a

collection of Puccinia momordicae (B 70 0008485). They

were paler than the teliospores of Puccinia windhoekensis and

had a coarser relief, which consisted of irregular, bent or flexu-

ous ridges. Similar to the teliospores of Puccinia windhoe-

kensis, Puccinia cephalandrae and Puccinia momordicae

have also 2 gems pores in each cell. We have not seen aecia

of Puccinia cephalandrae or Puccinia momordicae.

Another rust fungus with autoecious life cycle on Cucur-

bitaceae is the demicyclic rust fungus Puccinia cucumeris

Henn. (H

ENNINGS 1892: 371) (Type on Cucumis ficifolius,

Abyssinia, Colon. Eritrea, near Keren on the river Dari,

14. 3. 1891, leg. G. Schweinfurth). We examined the holotype

(B 70 0008495) but could not find uredinia or urediniospores.

Puccinia cucumeris differs from Puccinia windhoekensis

in having smaller, ± hyaline aeciospores (20–27 × 16–22 µm)

with obscure germ pores and with a maximum thickness of

the spore wall of up to 3 µm in one edge, and in having telio-

spores (35–47 × 26–36 µm) with only one germ pore in each

cell and pedicels which break short below the attachment.

For Puccinia cephalandrae-indicae Syd. & P. Syd. in S

Y-

DOW et al. (1906: 433) only teliospores are known. Puccinia

cephalandrae-indicae differs from Puccinia windhoekensis

in having smaller and thinner teliospores (32–42 × 18–27 µm)

with a thinner spore wall (epispore circa 2 µm) (S

YDOW et al.

1906).

In the genus definition of Puccinia only one germ pore

in each teliospore cell is quoted (C

UMMINS

& HIRATSUKA

2003). Nevertheless, some species of ‘Puccinia’ exist having

a second germ pore occasionally in the lower teliospore cell

like Puccinia abutili (see below) or having a second germ pore

in each teliospore cell like Puccinia cephalandrae (see above).

The same is true for Cumminsiella which, accordingly, falls

well into a Puccinia/Uromyces cluster in a molecular phylo-

genetic study (M

AIER et al. 2003).

Other characteristics of our specimens are Caeoma-type

aecia with catenulate, verrucous aeciospores. After C

UMMINS

& HIRATSUKA (2003), the aecia of Puccinia “are Aecidium-

type with peridium and catenulate and verrucose spores or

Uredo-type with mostly echinulate spores borne singly on

pedicels”. According to the construction similarities between

the telio- and urediniospores of Puccinia windhoekensis and

Puccinia cephalandrae (D

OIDGE 1927) we suppose that both

rusts are closely related in spite of the absence of a peridium

in the case of Puccinia windhoekensis. Nevertheless, both rust

species should be included in molecular analyses to clarify

their phylogenetic relationship.

One collection of ‘Puccinia cucumeris’ on Coccinia bar-

teri (Hook.) Keay in Nigeria (PUR F18688), which is pub-

lished by E

BOH (1981), may also belong to Puccinia windhoe-

kensis. E

BOH (1981) found only urediniospores with the size

24–28.8 × 19.2–21.6 µm.

Mycological Progress 4(1) / 2005

FABACEAE – ACACIA

Ravenelia halsei Doidge

OIDGE 1939: 504). Fig. 9

Type on Acacia ataxacantha DC. South Africa, Natal, Ndwedwe,

26. 7. 1938, leg. R. H. Halse (PREM 30117).

Pycnia and aecia unknown.

Uredinia Uredo-type, on pods, amphigenous on leaves and on

the petioles of the leaves, chiefly on the lower part of the leaf-

lets and the secondary rachis, separate, scattered, confluent in

the range of the secondary rachis, subepidermal, irregularly

rupturing pustules up to 4 mm wide, roundish or irregular in

outline, dark cinnamon-brown, early exposed, pulverulent,

surrounded by the torn epidermis, without spots surrounding

pustules. Urediniospores borne singly on pedicels, oblong el-

lipsoid or pyriforme, sometimes irregularly shaped, 22–33 ×

12–16 µm, finely echinulate, spore wall about 1–1.5 µm thick

at the sides and about 1.5–2 (2.5) µm thick at the apex, golden-

brown, germ pores conspicuous, 6 to 8 (to 12), when 6 equa-

torial, when more than 6 in a broader zone eguatorial, with

hyaline papillae that are weakly developed.

Telia rarely developed on the petioles of the leaves, chiefly on

the lower part of the leaflets and the secondary rachis, subepi-

dermal, blackish-brown, admixed with the uredinia, scattered,

later confluent, surrounded by the torn epidermis, without

spots surrounding telia. Teliospores slightly convex above and

slightly concave below, circular or subcircular, 105–125 µm

diameter, chestnut-brown, with 7 to 10 probasidial cells on

every diameter, outer layer of the cells smooth, cell wall about

6–7 µm thick at the apex, with a thin, hyaline epispore and a

thick, dark chestnut-brown endospore, with hyaline cysts on

the abaxial surface of the teliospore, margins of the cysts dif-

fusing in Hoyer’s fluid, pedicel deciduous.

Specimens examined:

On Acacia ataxacantha. South Africa, Natal, Ndwedwe, 26. 7. 1938,

leg. R. H. Halse, III. Type (PREM 30117). – On Acacia hereroensis

Engl. Namibia, C 26, 22°49’10.8’’S, 16°52’12.4’’E, 1.964 m asl.,

27. 4. 2002, leg. M. Mennicken No. NA 336, II. (PREM, WIND).

– On Acacia hereroensis. Namibia, C 28 Khomas Hochland,

22°35’41.4’’S, 16°52’27.4’’E, 1.938 m asl., 21. 4. 2002, leg. M.

Mennicken No. NA 321, II. (PREM, WIND). – On Acacia hereroen-

sis. Namibia, C 30 Steinhausen - Gobabis, 21°48’33.9’’S,

18°18’45.3’’E, 1.676 m asl., 3. 4. 2002, leg. M. Mennicken No. NA

194, II III. (PREM, WIND). – On Acacia hereroensis. Namibia, D

2860, B 8 - Hoba Meteorite, 19°40’09.6’’S, 17°52’25.9’’E, 1.586 m

asl., 16. 4. 2002, leg. M. Mennicken No. NA 288, II. (PREM,

WIND).

Ravenelia halsei was only known from the South African type

collection. Our collections agree well with the diagnosis of

Ravenelia halsei, from which only teliospores were so far de-

scribed (D

OIDGE 1939). Therefore, urediniospores are described

as new. Acacia hereroensis seems to be a new host plant and

Ravenelia halsei is new to the rust flora of Namibia.

Ravenelia transvaalensis Doidge (DOIDGE 1939: 505).

Fig. 10

Type on Acacia mellifera (Vahl) Benth subsp. detinens (Burch.) Bre-

nan (as Acacia detinens Burch.) South Africa, Gauteng Province,

near Pienaar’s river, leg. Mogg (PREM 27382).

Pycnia in general abaxial on leaves, vis-à-vis of the telia, form-

ing small, ± roundish groups up to 1 mm diameter, type 7.

Aecia and uredinia unknown.

Telia in general adaxial on leaves, vis-à-vis of the pycnia,

subepidermal, dark reddish-brown to blackish-brown, sepa-

Fig. 9: Ravenelia halsei on Acacia hereroensis. Uredinio- and

teliospores (NA 194). Scale bar = 10 µm.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

rate or scattered in confluent groups up to 2 mm diameter,

roundish or irregular in outline, pulverulent, surrounded by

the torn epidermis, without leaf spots surrounding telia, oc-

casionally with inconspicuous paler green leaf spots. Teliospo-

res convex above and slightly concave to even below, circu-

lar, subcircular, or irregular in outline, 60–115 µm diameter,

45–60 µm high, chestnut-brown, with 4 to 8 probasidial cells

on every diameter, outer layer smooth, cell wall about 5–8 µm

thick at the apex, with a thin, hyaline to yellowish epispore

and a thick, dark chestnut-brown endospore, with hyaline cysts

on the abaxial surface of the teliospores, with sterile intercalary

cells between the probasidial cells and the cysts, cysts equal

in number to the marginal cells, margins of the cysts diffus-

ing in Hoyer’s fluid, pedicel deciduous.

Specimens examined:

On Acacia mellifera (Vahl) Benth. Namibia, C 36 Wilhelmstal -

Omaruru, 21°31’02.4’’S, 16°02’28.0’’E, 1.287 m asl., 6. 4. 2002,

leg. M. Mennicken No. NA 217, 0 III. (PREM, WIND). – On Aca-

cia mellifera. Namibia, D 2337, short before C 33, riverbank,

21°04’56.3’’S, 16°02’43.7’’E, 1.537 m asl., 7. 4. 2002, leg. M. Men-

nicken No. NA 219, 0 III. (PREM, WIND). – On Acacia mellifera.

Namibia, D 2337, 21°05’12.1’’S, 15°58’42.4’’E, 1.527 m asl.,

7. 4. 2002, leg. M. Mennicken No. NA 226, 0 III. (PREM, WIND).

– On Acacia mellifera. Namibia, C 39 Outjo - Otavi, 19°39’11.8’’S,

16°59’54.8’’E, 1.422 m asl., 9. 4. 2002, leg. M. Mennicken No. NA

240, 0 III. (PREM, WIND). – On Acacia mellifera. Namibia, C 39

near Otavi, 19°38’02.0’’S, 17°19’23.4’’E, 1.421 m asl., 9. 4. 2002,

leg. M. Mennicken No. NA 249, 0 III. (PREM, WIND). – On Aca-

cia mellifera. Namibia, C 73, near D 2848, 19°27’20.1’’S,

18°07’42.7’’E, 1.455 m asl., 15. 4. 2002, leg. M. Mennicken No. NA

267, 0 III. (PREM, WIND). – On Acacia mellifera. Namibia, D 2512,

near Waterberg, 20°31’07.9’’S, 17°15’33.5’’E, 1.428 m asl.,

19. 4. 2002, leg. M. Mennicken No. NA 313, 0 III. (PREM, WIND).

– On Acacia mellifera. Namibia, C 26 near Windhoek, 22°40’40.3’’S,

16°59’17.9’’S, 1.814 m asl., 27. 4. 2002, leg. M. Mennicken No. NA

330, 0 III. (PREM, WIND).

Ravenelia transvaalensis is only known from the type col-

lection (D

OIDGE 1939) and two herbarium specimens on Aca-

cia nigrescens, Botswana, Mahalapye, leg. C.G. Hansford,

4. 1959, (BPI 191920, BPI 1110459) (FARR et al. n.d.). Our

collections conform well with the diagnosis of Ravenelia

transvaalensis, from which only teliospores were so far de-

scribed (DOIDGE 1939). Pycnia are newly described. Ravenelia

transvaalensis seems to be new to the rust flora of Namibia.

MALPIGHIACEAE – TRIASPIS

Puccinia ovamboensis Mennicken, Maier & Oberw. sp.

nov.

Figs. 11 (Plate 1), 12

Pycnia et aecia ignota. Uredinia caulicola vel amphigena, subepi-

dermalia, cinnamomea, usque ad 1 mm diam., mox nuda, pulveru-

lenta. Urediniosporae subgloboideae, obovoideae, pyriformes vel

reniformes, 27–36 × 19–27 µm, pariete echinulato, aureo-fusco,

1.5–2.5 µm crasso, 4–6 poris germinationis ± aequatorialibus vel su-

peraequatorialibus. Telia caulicola vel amphigena, subepidermalia,

nigra, usque ad 1 mm diam., mox nuda, pulverulenta. Teliosporae

bicellulares, ovoideae vel ellipsoideae, medio haud constrictae, apice

rotundato vel subacuto, base rotundata vel attenuata, raro diorchidi-

oideae, sanguineo-brunneae vel atro-fuscae, (38) 43–58 × (23)

25–37 µm, pariete grosse verrucoso, lateraliter 3–6 µm crasso, api-

caliter 4–12 µm crasso, poris germinationis inconspicuis, pedicello

hyalino, usque ad 60 µm longo, base discoideo-dilatata.

In caulis foliisque Triaspidis hypericoidis (DC.) Burch. subsp. nel-

sonii (Oliv.) Immelmann (Malpighiaceae).

Pycnia and aecia unknown.

Uredinia Uredo-type, caulicolous or amphigenous on leaves,

separate or scattered, subepidermal, small pustules, about

0.1–1 mm wide, roundish in outline, at first covered by the

blistered epidermis, early exposed, cinnamon-brown, pulve-

rulent, surrounded by the torn epidermis, without spots sur-

rounding pustules, early displaced by the telia. Urediniospores

borne singly on pedicels, variably in shape, subgloboid, ob-

ovoid, pyriform or reniform, often somewhat obliquely an-

gular, 27–36 × 19–27 µm, finely echinulate, echinulation

coarsely meshed, spore wall uniformly about 1.5–2.5 µm thick,

golden-brown, basal paler, germ pores conspicuous, 4 to 6,

± equatorial to superequatorial, with hyaline papillae that are

weakly developed.

Telia caulicolous and amphigenous on leaves, (partly) devel-

oping from the uredinia, separate or scattered in irregular

groups which can cover the whole leaf surface, partly conflu-

ent, subepidermal, small pustules, about 0.1–1 mm wide,

roundish or irregular in outline, at first covered by the blis-

tered epidermis, early exposed, black, pulverulent, surround-

ed by the torn epidermis, without spots surrounding pustules.

Teliospores bicellular, ovoid to ellipsoid, not constricted at the

septum, rounded or slightly apiculate at the apex, rounded or

attenuate at the base, occasionally diorchidioid, mahogany-

Fig. 10. Ravenelia transvaalensis on Acacia mellifera.

Teliospores (NA 240). Scale bar = 10 µm.

Mycological Progress 4(1) / 2005

coloured to deep mahogany-coloured, (38) 43–58 × (23)

25–37 µm, spore wall about 3–6 µm thick at the sides, about

4–12 µm thick at the apex, gibbous, warts respectively tuberc-

les coarse, closely crowded, germ pores obscure, pedicel per-

sistent, up to 60 µm long, thick-walled, not collapsing, hya-

line, sometimes obliquely inserted, consisting of a cylindrical

to conical upper portion and a roundish to napiform swelling

on the lower portion, margins of the swelling diffusing in Hoy-

er’s fluid.

On the leaves of Triaspis hypericoides subsp. nelsonii (Mal-

pighiaceae).

Etymology. Named after the biggest population group in

Namibia, the Ovambo.

Specimens examined:

On Triaspis hypericoides subsp. nelsonii. Namibia, Otavi Mountains,

D 3022, 19°25’04.8’’S, 17°53’28.3’’E, 1.478 m asl., 17. 4. 2002, leg.

M. Mennicken No. NA 292, II III. Holotype (PREM), Isotype

(WIND). – On Triaspis hypericoides subsp. nelsonii (= Triaspis nel-

sonii Oliv. var. austro-occidentalis Schinz). Namibia, Distr.

Kaokoveld, Dolomite outcrop on mountainside 17.3 miles S of

Kaoko-Otavi on road to Sesfontein, 21. 4. 1957, leg. de Winter &

Leistner No. 5606, II III. Paratype (B 10 9009305).

The only known rust fungus on the host genus Triaspis is the

rather similar Puccinia haematites Syd. & P. Syd. (S

YDOW &

YDOW 1911: 260) (Type on Triaspis auriculata Radlk.,

Uganda, near Kibwezi, 23. 6. 1906, leg. G. Scheffler No. 4 on

Herb. No. 46, Holotype B 70 0007785). Puccinia haematites,

from which only teliospores are known, differs from Puccinia

ovamboensis in having telia predominantly adaxial on leaves

and in having smaller teliospores (32–42 × 22–28 µm) with-

out an apical thickening (S

YDOW & SYDOW 1911). We have

examined the type material of Puccinia haematites and scaled

up the measurement of the teliospores to 36–48 (50) ×

22–33 µm. The teliospores of Puccinia haematites are either

not at all or are only very slightly constricted at the septum

and are rounded above and below. The colouration is pale ma-

hogany-coloured to mahogany-coloured, and the spore wall

is ± uniformly about 2.5–6 µm thick. Using light microscopy

the spore wall ornamentation of Puccinia haematites seems

to be very similar to those of Puccinia ovamboensis. In addi-

tion to the type collection from Uganda, Puccinia haematites

is known from Ghana (CUMMINS 1960), Kenya (DUKE 1926),

(NATTRASS 1961), and Tanzania (EBBELS & ALLEN 1979).

None of the authors gave a description of the rust fungus, par-

ticularly E

BBELS & ALLEN (1979) mentioned two collections,

one with pycnia and uredinia and the other with pycnia, ure-

dinia, and telia. No descriptions of the pycnia and uredinia

of Puccinia haematites are available.

Because of the telia which are developed amphigenous on

leaves and the bigger size of the teliospores with an usually

apically thickened spore wall Puccinia ovamboensis is de-

scribed as new.

MALVACEAE – ABUTILON

Puccinia abutili Berk. & Broome

(BERKELEY & BROOME

1875: 91). Figs. 13 (Plate 1), 14

Type on Abutilon graveolens. Ceylon, Kandy, 2. 1868, No. 523.

Syn. Puccinia carbonacea Kalchbr. & Cooke in K

ALCHBRENNER

(1882: 24). Type on Abutilon sp. South Africa, P. Natal, leg MacOwan

No. 1275.

Pycnia rarely? present, admixed with the telia, type 4.

Aecia and uredinia unknown.

Telia amphigenous on leaves, usually abaxial, on petiole, on

twigs, and on calyx, subepidermal, occasionally separate, usu-

ally scattered, arranged in concentrical, confluent blotches

of up to 6 mm diameter, forming more elongated pustules and

streaks along the veins of the leaves and along the twigs, or

more or less scattered over the whole leaf surface by heavy

infection, dark chocolate-brown, early exposed, pulverulent,

surrounded by the torn epidermis, spots surrounding telia

roundish or irregular in outline, yellowish, sometimes absent.

Teliospores bicellular, ellipsoid to oblong, occasionally dior-

chidioid, not or slightly constricted at the septum, 31–44 ×

19–30 µm, rounded or slightly attenuate at the apex and at the

base, spore wall about 2–5 µm thick, not or slightly up to 5 µm

thickened at the apex, brown, verrucous, upper germ pore api-

Fig. 12: Puccinia ovamboensis on Triaspis hypericoides ssp.

nelsonii. Uredinio- and teliospores (Holotype NA 292). Scale

bar = 10 µm.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

cal, sometimes obliquely inserted, lower germ pore just to one

side of the pedicel, occasionally a second germ pore just to the

other side of the pedicel, without papillae, pedicel normally

up to 10 µm long, occasionally up to 60 µm long, thin-walled,

persistent or not, not collapsing, hyaline, often obliquely in-

serted.

Specimens examined:

On Abutilon angulatum (Guill. & Perr.) Mast. Namibia, C 3016,

18°51’22.4’’S, 18°28’48.5’’E, 1.199 m asl., 13. 4. 2002, leg. M.

Mennicken No. NA 274, III. (PREM, WIND). – On Abutilon cf.

austro-africanum Hochr. Namibia, C 30 Steinhausen - Gobabis,

riverbank of the Black Nossop, 22°22’53.7’’S, 18°56’35.5’’E,

1.479 m asl., 3. 4. 2002, leg. M. Mennicken No. NA 195, 0 III.

(PREM, WIND). – On Abutilon cf. austro-africanum. Namibia,

Ombeameiata, housecamp, 21°35’27.1’’S, 18°01’14.0’’E, 1.591 m

asl., 2. 4. 2002, leg. M. Mennicken No. NA 183, III. (PREM, WIND).

– On Abutilon cf. austro-africanum. Namibia, BIOTA-observatory

at Toggekry 250 (Omatako-Ranch), 21°30’22.3’’S, 16°44’05.7’’E,

9. 3. 2002, leg. M. Mennicken No. NA 136, III. (PREM, WIND). –

On Abutilon cf. rehmannii Baker f. Namibia, Steinhausen, camp W

farmhouse, 21°48’37.5’’S, 18°12’54.6’’E, 1.666 m asl., 2. 4. 2002,

leg. M. Mennicken No. NA 170, III. (PREM, WIND).

The microcyclic rust fungus Puccinia abutili is widespread in

Africa (e.g. C

ROUS et al. 2000; DOIDGE 1927, 1950; EBOH

1984; GJÆRUM 1985, 1995; HENNINGS 1907; HOPKINS 1950;

ØRSTAD 1956; KRANZ 1964; MAJEWSKI & NOWAK 1982;

ATTRASS

1961; R

OTHWELL 1983; SACCARDO 1910; SYDOW

& SYDOW 1909; WAKEFIELD 1920; WAKEFIELD & HANSFORD

1949). Neither pycnia nor a second germ pore in the lower cell

of the teliospores (see discussion under Puccinia windhoe-

kensis) have been mentioned so far. Puccinia abutili is new to

the rust flora of Namibia. Abutilon austro-africanum and Abu-

tilon rehmannii seem to be new host plants.

SOLANACEAE – LYCIUM

Puccinia afra G. Winter

(WINTER 1887: 26).

Figs. 15a, b (Plate 2), 16

Type on Lycium afrum L. South Africa, near Cape Town, 1886, leg.

P. MacOwan.

Pycnia unknown.

Aecia not seen.

Uredinia Uredo-type, amphigenous on leaves, subepidermal,

small pustules up to 1 mm wide, ± roundish, separate or scat-

tered, without spots surrounding uredinia, cinnamon-brown,

early exposed, pulverulent and surrounded by the torn epi-

dermis. Urediniospores ellipsoid to oblong, (48) 51–66 (72)

× 20–33 µm, slightly attenuate at the apex, rounded at the base,

smooth in the lower one sixth, echinulate in the upper five

sixths of the spore, spines becoming continuously larger from

the lower parts to the apex, spore wall about 2–3 µm thick at

the sides, about 3–5 µm thick at the apex, yellowish-brown,

germ pores 4 to 5, supraequatorial, with hyaline papillae that

are medium developed.

Telia amphigenous on leaves, subepidermal, small pustules

up to 1 mm wide, roundish, separate or scattered, partially re-

placing the uredinia, blackish-brown, early exposed, pulve-

rulent, without spots surrounding telia, surrounded by the torn

epidermis. Teliospores bicellular, ellipsoid, not or slightly con-

stricted at the septum, 46–56 × 26–31 µm, acuminate at the

apex, apex drawn out into a yellowish cone-shaped apicule,

Fig. 14: Puccinia abutili on Abutilon cf. austro-africanum.

Teliospores (NA 195). Scale bar = 10 µm.

Fig. 16: Puccinia afra on Lycium sp. Uredinio- and teliospores

(RSA 167). Scale bar = 10 µm.

Mycological Progress 4(1) / 2005

Plate 2. Fig. 15. Puccinia afra on Lycium sp. (RSA 167). SEM. Scale bars = 10 µm. a. Urediniospores. b. Teliospore. Fig.

17. Puccinia lycii on Lycium sp. (RSA 177). SEM. Scale bars = 10 µm. a. Uredinio- and teliospores. b. Teliospores and ure-

diniospore.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

rounded at the base, spore wall about 2–3.5 µm thick at the

sides, about 5–10 µm thick at the apex, dark chestnut-brown,

densely and roughly verrucous, upper germ pore centrically

between the apex and the septum, lower germ pore centrical-

ly between the septum and the base, with yellowish papillae,

pedicel up to 80 µm long, thick-walled, persistent, consist-

ing of a cylindrical upper portion and a vesiculous inflation

with irregular crenate margins on the lower portion, margins

of the swelling diffusing in Hoyer’s fluid, yellowish in the up-

per portion, hyaline in the lower portion.

Specimens examined:

On Lycium sp. South Africa, Western Cape Province, BIOTA-ob-

servatory at Moedverloren 208, 31°27’40.1’’S, 18°26’31.3’’E,

18. 9. 2002, leg. M. Mennicken No. RSA 167, II III. (PREM,

WIND). – On Lycium sp. South Africa, Western Cape Province,

BIOTA-observatory at Moedverloren 208, 31°27’43.2’’S,

18°26’27.3’’E, 119 m asl., 18. 9. 2002, leg. M. Mennicken No. RSA

175, II III. (PREM, WIND).

Our collections conform well with the description of Puccinia

afra given by DOIDGE (1927).

Puccinia lycii Kalchbr. (KALCHBRENNER 1882: 21).

Figs. 17a, b (Plate 2), 18

Type on Lycium tubulosum Nees. South Africa, Somerset East, leg.

MacOwan No. 1410 (B 70 0007290).

Pycnia unknown.

Aecia rarely developed, Aecidium-type, amphigenous? on

leaves, densely scattered in small and roundish groups up to

2 mm diameter, without spots surrounding aecia, aecial cups

small, spore mass creamish-white or white, surrounded by ir-

regularly and finely lacerated, white to light brownish peri-

dium, cells of the peridium medium connected, outer wall

finely striate, inner wall coarsely verrucous. Aeciospores an-

gular globoid, subgloboid to ellipsoid, 13–20 × 11–15 µm,

spore wall 0.5–1 µm, hyaline, germ pores inconspicuous.

Uredinia Uredo-type, amphigenous on leaves, subepidermal,

small pustules, ± roundish, up to 1 mm wide, separate or scat-

tered, without spots surrounding uredinia, foxy red, early ex-

posed, pulverulent and surrounded by the torn epidermis. Ure-

diniospores ovoid, ellipsoid to oblong, 45–53 (65) × (21)

23–27 (30) µm, smooth in the lower one tenth, echinulate in

the upper nine tenths of the spore, spines becoming continu-

ously larger from the lower parts to the apex, spore wall about

(1.5) 2–2.5 µm thick at the sides, about 3–4 µm thick at the

apex, yellowish-brown, germ pores 4 to 6 and slightly supra-

equatorial, with hyaline papillae that are medium developed.

Telia amphigenous on leaves and on twigs, subepidermal,

small pustules up to 1.5 mm, roundish, separate or scattered,

partially replacing the uredinia, blackish-brown to black, early

exposed, without spots surrounding telia, surrounded by the

torn epidermis. Teliospores bicellular, ellipsoid to oblong, not

or slightly constricted at the septum, 43–52 × 27–33 µm,

rounded or acuminate at the apex, rounded at the base, spore

wall about 2–4 µm thick at the sides, about 3–6 µm thick at

the apex, (dark) chestnut-brown, finely verrucous, warts ir-

regularly, closely meshed, brownish-yellow, upper germ pore

centrically between the apex and the septum, occasionally at

the apex, lower germ pore ± centrically between the septum and

the base, with brownish-yellow papillae, pedicel persistent,

up to 110 µm long, thick-walled, consisting of a short cylind-

rical upper portion and a long napiform inflation on the lower

portion, margins of the swelling diffusing in Hoyer’s fluid,

yellowish in the upper portion, hyaline in the lower portion.

Specimens examined:

On Lycium tubulosum Nees. South Africa, Somerset East, leg. Mac

Owan No. 1410, ex Herbar Winter, III. (Type B 70 0007290). – On

Lycium sp. South Africa, Western Cape Province, BIOTA-observato-

ry at Moedverloren 208, 31°27’52.1’’S, 18°26’31.1’’E, 136 m asl.,

18. 9. 2002, leg. M. Mennicken No. RSA 177, I II III. (PREM, WIND).

Our collection agrees well with the description of Puccinia

lycii given by D

OIDGE (1927). In South Africa Puccinia lycii

is known from four Lycium species in the provinces Free-State

and Eastern Cape (C

ROUS et al. 2000). Until now only uredinia

and telia were known. The aecia are newly described.

Fig. 18: Puccinia lycii on Lycium sp. Peridium cells, aecio-,

uredinio-, and teliospores (RSA 177). Scale bar = 10 µm.

Puccinia turgida P. Syd. & Syd. (SYDOW & SYDOW 1904:

266). Figs. 19a, b (Plate 3), 20

Type on Lycium europaeum L. Palaestina, near Jericho, 300 m asl.,

29. 3. 1897, leg J. Bornmüller, Iter Syriacum 1897 No. 1021. (B 70

0007287).

Pycnia rarely developed, amphigenous? in the centre of the

aecia, type 4.

Aecia Aecidium-type, amphigenous on leaves, densely scat-

tered in small and roundish groups up to 3 mm diameter,

without spots surrounding aecia, aecial cups small, erumpent,

circa 300 µm diameter, up to 1 mm tall, spore mass whitish-

yellow or white, surrounded by irregularly and finely lacerat-

ed, white to light brownish peridium, cells of the peridium

medium connected, outer wall smooth, inner wall coarsely

verrucous. Aeciospores angular subgloboid, cuboid to ellip-

soid, 27–36 × 20–28 µm, delicately verrucous, spore wall

1–2 µm, hyaline, germ pores inconspicuous.

Uredinia Uredo-type, amphigenous on leaves, subepidermal,

small pustules, ± roundish, up to 1 mm wide, separate or scat-

tered, without spots surrounding uredinia, cinnamon-brown,

early exposed, pulverulent and surrounded by the torn epi-

dermis. Urediniospores obovoid, ellipsoid to oblong, (32)

36–51 × (17) 19–25 µm, smooth in the lower one fifth, ech-

inulate in the upper four fifths of the spore, spines becoming

continuously larger from the lower parts to the apex, spore

wall about 1.5–2.5 µm thick at the sides, about 2–2.5 µm thick

at the apex, yellowish-brown, germ pores generally 4 to 5 and

equatorial to slightly supraequatorial, occasionally up to 8 and

tending to be bizonate with 4 to 5 germ spores in a lower zone

and 1 to 3 germ pores in an upper zone, with hyaline papil-

lae that are medium developed.

Telia amphigenous on leaves, subepidermal, small pustules

up to 1 mm, roundish, separate or scattered, partially replac-

ing the uredinia, black, early exposed, without spots sur-

rounding telia, surrounded by the torn epidermis. Teliospores

bicellular, ellipsoid to oblong, not or slightly constricted at the

septum, 39–55 (60) × 27–38 µm, rounded or acuminate at the

apex, rounded at the base, spore wall uniformly about 2–4 µm

thick, not or slightly up to 5 µm thickened at the apex, chest-

nut-brown, roughly verrucous, warts and tubercle irregularly,

sparsely to coarsely meshed, brownish-yellow, germ pore of

the upper cell centrical between the apex and the septum, germ

pore of the lower cell centrical between the septum and the

base, with brownish-yellow papillae, spore wall with papilla

up to 6 µm thick, pedicel up to 55 µm long, thick-walled, per-

sistent, consisting of a cylindrical upper portion and a napi-

form inflation on the lower portion, margins of the swelling

diffusing in Hoyer’s fluid, yellowish in the upper portion, hya-

line in the lower portion, occasionally obliquely inserted.

Specimens examined:

On Lycium europaeum. Palaestina, near Jericho, 300 m asl.,

29. 3. 1897, leg J. Bornmüller, II III. Iter Syriacum 1897 No. 1021.

Type, but labelled as type of Puccinia lycii Kalchbr. nov. var. asia-

tica Sydow (B 70 0007287). – On Lycium europaeum. Palaestina,

near Jericho, II III. Kryptogamae exsiccatae No. 802 (B 70 0007281).

– On Lycium cf. oxycarpum Dunal. Namibia, D 1852 Witvlei - Omi-

tara, short behind B 6, 22°21’20.6’’S, 18°04’35.6’’E, 1.595 m asl.,

4. 4. 2002, leg. M. Mennicken No. NA 206, 0 I. (PREM, WIND). –

On Lycium cf. oxycarpum. Namibia, D 1852 Nossop - Seeis,

22°19’29.7’’S, 17°43’11.3’’E, 1.608 m asl., 4. 4. 2002, leg. M. Men-

nicken No. NA 207, II III. (PREM, WIND). – On Lycium cf. oxy-

carpum. Namibia, B 1 Windhoek - Okahandja, riverbank of the

Swakop, 22°02’03.2’’S, 16°56’08.9’’E, 1.346 m asl., 6. 4. 2002, leg.

M. Mennicken No. NA 214, 0 I II III. (PREM, WIND). – On Lycium

cf. oxycarpum. Namibia, D 2337, short before C 33, riverbank,

21°04’57.6’’S, 16°02’44.3’’E, 1.539 m asl, 7. 4. 2002, leg. M. Men-

nicken No. NA 223, II III. (PREM, WIND). – On Lycium sp. Nami-

bia, Okahandja, 1.200 m asl., 1. 7. 1907, leg Kurt Dinter No. 597, III.

(B 70 0007276).

The type collection is labelled as a type of Puccinia lycii

Kalchbr. nov. var. asiatica Sydow. H. & P. Sydow did not

publish this name. MAGNUS (1900: 437) cited the same col-

lection as Puccinia lycii Kalchbr. We propose that H. & P.

Sydow first wanted to describe a new variant of Puccinia lycii.

Later, they described a new species without changing the in-

scription of the herbarium specimen. Until now only uredinia

and telia were known. The aecia are described as new.

This rust fungus differs clearly from all other known

species of Puccinia with verrucous teliospores on Lycium:

Fig. 20: Puccinia turgida on Lycium cf. oxycarpum. Peridi-

um cells and aeciospores (NA 206), uredinio- and teliospores

(NA 223). Scale bar = 10 µm.

Mycological Progress 4(1) / 2005

Plate 3. Fig. 19. Puccinia turgida on Lycium cf. oxycarpum. (NA 207). SEM. Scale bars = 10 µm. a. Uredinio- and teliospores.

b. Telio- and urediniospores. Fig. 21. Uredopeltis flavae on Grewia flava. Uredinium with peripheral paraphyses (Holotype

NA 293). SEM. Scale bar = 100 µm.

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

Puccinia afra (see above) has longer uredinio- and teliospores

with close-by warts. Puccinia spinulosa Jørst. (J

ØRSTAD 1956:

592), from which only teliospores are known, differs in hav-

ing smaller teliospores with “upper germ pore usually some-

what depressed, lower germ pore in the lower part of basal

cell, often near pedicel”. Puccinia lycii (see above) has very

finely verrucous teliospores. Puccinia globosipes Peck (P

ECK

1885: 34) lacks the cylindrical upper portion of the pedicel,

and the teliospores are smaller. Puccinia verwoerdiana Van

der Byl (V

AN DER B

YL 1927: 226) has longer urediniospores.

Puccinia turgida is new to the rust flora of Namibia. Lycium

oxycarpum seems to be a new host plant.

TILIACEAE – GREWIA

Uredopeltis flavae Mennicken, Maier & Oberw. sp. nov.

Figs. 21 (Plate 3), 22

Pycnia et aecia ignota. Uredinia abaxialia, corbuliformia, in lacunis

foliorum nata, sparsa vel laxe aggregata, usque ad 350 µm diam., pa-

raphysibus peripheralibus numerosis, incurvatis, brunneis, usque ad

circa 120 µm longis, 5–12 µm latis, deorsum conjunctis, pariete ir-

regulari, 1–5 µm crasso, apicaliter incrassato usque ad 15 µm. Ure-

diniosporae subgloboideae, ellipsoideae, guttiformes vel pyriformes,

18–29 × 15–18 µm, pariete echinulato, hyalino vel dilute aureo,

1–1.5 µm crasso, 2–6 poris germinationis inconspicuis, approx. ae-

quatorialibus. Telia abaxialia, subepidermalia, erumpentia, sparsa

vel laxe aggregata, nigro-brunnea vel nigra, usque ad 250 mm diam.,

paraphysibus peripheralibus. Teliosporarum massa lateraliter circa

17–23 sporis, verticaliter circa 12–25 sporis, circa 140–220 µm

diam., circa 130–240 µm alta aggregata, sporis singularibus unicel-

lularibus, 12–27 × 9–15 µm, pariete castaneo.

In foliis Grewiae flavae DC. (Tiliaceae).

Pycnia and aecia unknown.

Uredinia abaxial on leaves, separate or scattered in irregular

groups, partly admixed with the telia, forming roundish sori

which are bordered by numerous, conspicuous, brown para-

physes, sori superepidermal, with an elongate, basal stipe-like

structure which is deeply seated in the leaf tissue and arises

from cavities or sunken areas which are typically for leaves

of the host plant, up to 350 µm diameter, pulverulent, covered

by the pale brownish urediniospores, without leaf spots sur-

rounding uredinia. Stipe-like structure consisting of several

lateral united, septate, hyaline hyphae. Paraphyses surround-

ing uredinia in great quantities arise from the stripe-like struc-

ture, incurved, cylindrical, occasionally clavate or capitate, of-

ten irregular in outline, septate, basally united and/or coad-

unate, up to circa 120 µm long and about 5–12 µm wide. The

wall of the paraphyses irregularly about 1–5 µm thick, up to

15 µm thick at the apex, brown to chestnut-brown. Uredinio-

spores being produced from the apical layers of the stipe-like,

cellular base, sessile or nearly so, subgloboid, ellipsoid, drop-

shaped or pyriform, 18–29 × 15–18 µm, echinulate, echinu-

lation generally coarsely meshed, spore wall uniformly about

1–1.5 µm thick, yellowish to hyaline, germ pores inconspicu-

ous, 2 to 6, generally ± equatorial, without papillae.

Telia abaxial on leaves, separate or scattered in irregular

groups, often admixed with the uredinia, developing partly

from the uredinia, subepidermal in origin but becoming

erumpent, chocolate-brown to blackish pustules, up to 250 mm

wide, roundish, ellipsoid, or irregular in outline, compact to

cushionlike, early exposed, without leaf spots surrounding

telia. Paraphyses partly surrounding telia in great quantities

(attributes see uredinia) probably relicts from the uredinia af-

ter transmutation from uredinia to telia. Teliospores jointed

together in catenulate or irregular arranged crusts, which con-

sist of circa 17 to 23 lateral spores and of circa 12 to 25 vertical

spores. Crusts circa 140–220 µm diameter, circa 130–240 µm

high, compact and generally without spaces between single

spores. Spores unicellular, variable in shape, angular globoid,

subgloboid, ellipsoid, ovoid, oblong or irregular in outline,

12–27 × 9–15 µm, spore wall brown to chestnut brown in the

upper spores, hyaline in the lower spores, becoming progres-

sively darker from the base towards the apex, smooth, in the

lower spores uniformly about 1–1.5 µm thick, in the upper

spores about 1–1.5 µm thick at the sides, up to 3 µm thick at

the apex, germ pore inconspicuous in the upper spores, ob-

scure in the lower spores, apical, with hyaline papilla that is

weakly developed.

Fig. 22. Uredopeltis flavae on Grewia flava. Paraphyses, ure-

diniospores, and a teliospore crust (Holotype NA 293). Scale

bar = 10 µm.

Mycological Progress 4(1) / 2005

On the leaves of Grewia flava DC. (Tiliaceae).

Etymology. Named after the host plant, Grewia flava.

Specimen examined:

On Grewia flava DC. Namibia, Otavi Mountains, D 3022,

19°25’04.0’’S, 17°53’28.4’’E, 1.479 m asl., 17. 4. 2002, leg. M.

Mennicken No. NA 293, II III. Holotype (PREM), Isotype (WIND).

As far as could be established there are eight known rust fungi

on Grewia:

Aecidium warneckeanum Henn. (HENNINGS 1907: 105) (Type

on Grewia carpinifolia, Ost-Usambara, Amani, 09. 1903, leg.

Warnecke & Eichelbaum No. 106) forms “an Zweigen große,

geweihartig verzweigte Hexenbesen”.

Cumminsina clavispora Petr. (P

ETRAK 1955: 474) (Type on

Grewia nenensis, Angola, Huila Banks of River Nene, near

Humpata) forms teliospores “consisting of chains of laterally

adherent cells forming a club-shaped, pigmented head, pedi-

cel simple basally but with apical cells equal in number of the

chains of cells” (C

UMMINS & HIRATSUKA 2003).

Kuehneola grewiae (Thirum. & Mundk.) Thirum. (T

HIRU

MALACHAR 1960: 692) (Syn. Catenulopsora grewiae Thirum.

& Mundk. (T

HIRUMALACHAR & M

UNDKUR 1951: 13/14), type

on Grewia populifolia Vahl, India, Punjab) differs from Ure-

dopeltis flavae inter alia in having teliospores in separated,

permanently united chains, 16–20 spores per chains (T

HIRU-

MALACHAR & M

UNDKUR 1951).

Phakopsora microspora Cummins (C

UMMINS 1960: 37) (Type

on Grewia carpinifolia Fuss. Ghana, Biriwa, 3. 11. 1957, leg.

L. Piening CB 2620, IMI, PUR) differs from Uredopeltis

flavae in having smaller urediniospores (16–19 × 14–17 µm)

and shorter and narrower paraphyses (20–35 µm long and

6–8 µm wide) (C

UMMINS 1960). The teliospores are smaller:

(6) 8–11 (13) × (8) 11–17 (20) µm (C

UMMINS 1960).

Puccinia tiliifoliae T.S. Ramakr. & Sundaram (as Puccinia

tiliaefoliae) (RAMAKRISHNAN & SUNDARAM 1955: 194) (Type

on Grewia tiliifolia Vahl. (as Grewia tiliaefolia). India,

Madras, Ambalavayal (Malabar), 3. 9 1954, leg. N.V. Sun-

daram) has bicellular teliospores with persistent pedicels.

Mesospores are commonly, tricellular teliospores rarely de-

veloped.

Ravenelia atrides Syd. & P. Syd. (S

YDOW & SYDOW 1912a:

438) (Type on Grewia caffra. South Africa, Natal, Durban,

7. 7. 1911, leg. Pole Evans No. 1670) differs from Uredopeltis

flavae in having dark and dull brown, opaque teliospore heads,

110–190 µm diameter, with 15–22 spores in each diameter

and with an epispore about 4–10 µm thick (D

OIDGE 1927). We

have examined the type collection of Ravenelia atrides (S

F29702). The teliospore heads differentiate Ravenelia atrides

undoubtedly from Uredopeltis flavae. The urediniospores of

the type collection of Ravenelia atrides measure15–22 (24) ×

13–16 µm.

Uredo corbiculoides Cummins (C

UMMINS 1945: 219) (Type

on Grewia sp. Uganda, Serere, Teso, 3. 1933, leg. Hansford

No. 1630), from which Cummins published a LM-micro-

graph, has uredinia which are constructed similar to those of

Uredopeltis flavae. It differs from Uredopeltis flavae in hav-

ing shorter urediniospores (16–20 × 13–16 µm) and thinner-

walled paraphyses (about 1–2 µm thick, up to 4 µm thick)

UMMINS 1945).

Uredopeltis chevalieri (see below) differs from Uredopeltis

flavae in the construction of uredinia with shorter, paler para-

physes. The teliospore heads of Uredopeltis chevalieri are

smaller and consist of fewer cells. C

UMMINS (1945) published

a LM-micrograph of a telium.

Our collection conforms well with the characteristics of the

genus Uredopeltis (H

ENNINGS 1908: 223). L

AUNDON (1963b)

re-examined the genus and published two LM-micrographs

of telia. C

UMMINS & H

IRATSUKA (2003) stated that “Uredo-

peltis appears to be similar to Phakopsora and Physopella, but

the latter have non-erumpent telia”.

Uredopeltis cf. chevalieri (Pat. & Har.) J. Walker & R.G.

Shivas

(WALKER & SHIVAS 2004: 43). Fig. 23

Type on Grewia breviflora Benth. Australia, Western Australia, Kim-

berley region, Beverley Springs Station, 9. 5. 1995, leg. A. A.

Mitchell (Holotype PERTH 3970574, Isotype DAR 74838).

Anamorph. Uredo grewiae Pat. & Har. (P

ATOUILLARD & HARIOT

1900: 237). Type on Grewia ferruginea Hochst. Senegal, Cayor, near

Thies.

Syn. Phakopsora grewiae (Pat. & Har.) Cummins (C

UMMINS 1945:

206). Type on Grewia sp. Senegal, Thies (Cayor), leg. Chevalier.

Syn. Dasturella grewiae (Pat. & Har.) Thirum. (T

HIRUMALACHAR

1946: 348). Type on Grewia monticola Sond. South Africa, Nel-

spruit? (as Kelspruit), Tvl., Experimental Station, 2. 8. 1940 (PREM?

No. 32401).

Syn. Uredopeltis grewiae (Cummins) Sathe (S

ATHE 1969: 176). Type

on Grewia asiatica L. India, Poona.

Pycnia and aecia unknown.

Uredinia amphigenous on leaves, subepidermal, separate or

scattered in irregular groups, partly admixed with the telia,

brown to pale brown pustules, small, up to 150 µm wide,

roundish, ellipsoid or irregular in outline, long remaining co-

vered by the epidermis, opening through an apical pore, later

gapping, surrounded by the torn epidermis, pulverulent, co-

vered by the pale brownish urediniospores, generally sur-

rounded by brownish leaf spots, spots often margined by the

nerves of the host leaves. Paraphyses surrounding uredinia in

great quantities are incurved, cylindrical, often irregular in out-

line, about 25–75 µm long and about 5–12 µm wide, wall of

the paraphyses about 1 µm thick at the base, up to 4 µm thick

near the apex, yellowish-brown to hyaline. Urediniospores

subgloboid, ellipsoid, drop-shaped or pyriform, 16–29 ×

12–17 µm, slightly echinulate, spore wall uniformly between

0.5–1.5 µm thick, yellowish to hyaline, germ pores incon-

spicuous, 3 to 5, ± equatorial, without papillae.

Telia amphigenous on leaves, separate or scattered in irregu-

lar groups, often admixed with the uredinia, partly develop-

MENNICKEN, MAIER

& OBERWINKLER:A contribution to the rust flora (Uredinales) of southern Africa

ing from the uredinia, subepidermal in origin but becoming

erumpent, chocolate-brown pustules small, up to 250 mm

wide, roundish, ellipsoid, oblong, or irregular in outline, com-

pact to cushion-like, long remaining covered by the epider-

mis, later exposed, generally surrounded by brownish leaf

spots, spots often margined by the nerves of the host leaves.

Paraphyses partly surrounding telia in great quantities (attribu-

tes see uredinia), probably relicts from the uredinia after trans-

mutation from uredinia to telia. Teliospores jointed together

in catenulate or irregularly arranged crusts which consist of

circa 10 to 18 lateral spores and of circa 5 to 11 vertical spores.

Crusts measure circa 80–180 µm diameter, circa 60–120 µm

high, at the beginning compact and without spaces between

single spores, later becoming disintegrated and releasing the

spores. Spores unicellular, variable in shape and size, globoid,

subgloboid, ellipsoid, ovoid, oblong, amygdaline, drop shap-

ed, or irregular in outline, often angular, 10–29 × 6–16 µm,

spore wall brown to chestnut brown, smooth, about 0.5–2 µm

thick at the sides, up to 3 µm thick at the apex, germ pore in-

conspicuous in the upper spores, obscure in the lower spores,

apical, with a weakly developed hyaline papilla.

Specimens examined:

On Grewia flavescens Juss. Namibia, Koukuas, 18°54’17.4’’S,

18°17’16.0’’E, 1.210 m asl., 14. 4. 2002, leg. M. Mennicken No. NA

264, II III. (PREM, WIND). – On Grewia flavescens. Namibia, Otavi

Mountains, D 2863, 19°30’50.1’’S, 17°44’42.5’’E, 1.659 m asl.,

17. 4. 2002, leg. M. Mennicken No. NA 298, II III. (PREM, WIND).

– On Grewia flavescens. Namibia, C 39 near Otavi, 19°38’00.2’’S,

17°19’24.2’’E, 1.416 m asl., 9. 4. 2002, leg. M. Mennicken No. NA

252, II III. (PREM, WIND).

Our collections do not conform in all details with the charac-

teristics of Uredopeltis chevalieri and its synonymous species.

Therefore, we use the prefix cf. before the epithet. The ure-

diniospores reported so far are slightly broader than in our

collections: 21–28 × 16–21 µm (Grewia ferruginea:PA-

TOUILLARD & HARIOT 1900), 20–28 × 15–21 µm (Grewia sp.:

CUMMINS

1945), 20–26 × 15–20 µm (Grewia monticola:

HIRUMALACHAR

1946), 18–26 × 15–19 µm (Grewia pubes-

cens Beauv.: V

IENNOT-BOURGIN 1953), and (22) 24–31 (37)

× (17) 18–21 (22) µm (Grewia breviflora: W

ALKER & SHIVAS

2004). The urediniospores of the type collection of Uredo-

peltis chevalieri are longer compared to any other cited col-

lection (see above). The spectrum of sizes of teliospore crusts

is as follows: 64–165 µm in diameter and 100–140 µm high

(CUMMINS 1945: type collection of Phakopsora grewiae),

130–154 µm in diameter and 130–167 µm high (THIRU-

MALACHAR 1946: type collection of Dasturella grewiae), and

up to 180 µm in diameter as well as in height (WALKER &

SHIVAS 2004: type collection of Uredopeltis chevalieri). Gi-

ven these size differences in both urediniospores and telia, the

occurrence of the rust on different species of Grewia and its

geographic distribution on three continents, we consider it pos-

sible, that Uredopeltis chevalieri has to be divided into dif-

ferent species. To address this question, all known specimens

have to be included in morphological analyses to clarify their

taxonomic relationships. Grewia flavescens seems to be a new

host plant of Uredopeltis cf. chevalieri.

Acknowledgements

The authors thank two unknown referees for peer reviewing

the manuscript, the curators of B, PREM, and S for the loan

of specimens, Matthias Lutz and Michael Weiß for critical

comments on the manuscript, Karl-Heinz Hellmer & Friedhelm

Albrecht for performing the scanning electron microscopy,

Michael Weiß for help with the Latin diagnosis, and Sheila

and Brian Longman for comments on the English text. The

study was made possible by the Federal Ministry of Educa-

tion and Research in Germany through the sourcing of the

BIOTA Southern Africa project.

The Namibian Ministry of Environment and Tourism, the

Western Cape Nature Conservation Board, and the Conser-

vation Services of the South African National Parks are grate-

Fig. 23. Uredopeltis cf. chevalieri on Grewia flavescens. Pa-

raphyses, urediniospores, and a teliospore crust (NA 264).

Scale bar = 10 µm.

Mycological Progress 4(1) / 2005

fully acknowledged for issuing research permits. The authors

thank all land owners and decision makers for the permission

to collect rust-infected plants. Renate Austermühle and Ben

Strohbach are gratefully acknowledged for help with the de-

termination of the host plants.

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Mycological Progress 4(1) / 2005

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Full-text available

Jan 2013

Some collections of Puccinia pelargonii-zonalis made in Belgium are described and illustrated. A synopsis of the three species of Puccinia growing on Pelargonium is proposed, as well as a key to identify them. These three Rust fungi are P. granularis, P. morrisonii and P. pelargonii-zonalis. This last species is the only one to be present in Belgium. Some additional data are given about its presence in the country.

Host range and potential distribution of the rust fungus, Miyagia pseudosphaeria, a biological control agent for Sonchus species

Article

Aug 2017

Sonchus species (Sowthistles) have a world-wide distribution and are serious weeds of crops and the environment. We assessed the suitability of the rust, Miyagia pseudosphaeria, for use in biological control of Sonchus species in Australia and elsewhere. Testing of a limited range of plant species showed the host-range of the rust to be restricted to Sonchus species. We measured spore germination and development of the rust on the host plant in relation to temperature. These measurements, the biology and distribution of the rust, and that of the host S. oleraceus, were used to model the potential world distribution of the rust in CLIMEX. The model indicated that there was limited potential for further spread in Australia; the rust already being widespread. The rust is not present in Canada, but the CLIMEX model indicated that cold temperature will be the main limiting factor for the rust should it be introduced, compromising its suitability as a biological control agent. In South America the rust could potentially be introduced as a biological control agents in areas with Mediterranean and subtropical climate. Molecular studies in combination with more comprehensive inoculation and temperature studies are needed to establish if there are pathotypes of the rust that might be more suitable for use in biological control.

The rusts of geraniaceae in Australia

Article

Jan 2010

J. Walker

Two native and two introduced rusts are present on hosts in Geraniaceae in Australia. The rare Puccinia geranii-pilosi McAlpine occurs on native species of Geranium, and P. morrisonii McAlpine is found on native Pelargonium spp. The introduced P. pelargonii-zonalis Doidge is common, widespread and often damaging on cultivated zonal pelargonium (Pelargonium x hortorum L. H. Bailey). Uromyces geranii (DC.) Fr. was first recognised in Australia in 2003 on Geranium sp. in New South Wales but herbarium specimens show that it has been present since at least 1911. All species are fully described and illustrated from Australian material, with comments on host ranges and geographic distributions. Lectotypes are selected for P. geranii-pilosi and P. morrisonii. Differences between species are discussed, an identification key given and comparisons made with exotic rusts of Geraniaceae. Evidence suggests that P. pelargonii-zonalis was introduced to Australia between 1906 and 1912; its subsequent early spread is discussed. A smooth wall patch free of spines (a tonsure) is present on urediniospores of the four rusts. Erroneous Australian reports of rusts on Erodium are clarified, Puccinia polygoni-amphibii is newly recorded from uredinial collections on Fallopia and Persicaria, and its potential connection with Geranium hosts discussed.

Checklist of Namibian plants protected by law

Data

Full-text available

May 2020

Working list of plants protected by law in Namibia. Input and comments welcome

Uredinales of Maharashtra State, India

Article

May 2024

Arun V. Sathe

This paper deals with 140 species of rust fungi, which are so far reported from the Maharashtra State and are distributed amongst 34 rust genera. The deposition number of an authentic state material is indicated, wherever available. The critical remarks by the author, wherever necessary are included, which might help in clarifying the taxonomic position of the particular rust. The nomenclature of some rusts which needed rectification is achieved in this work. The host index is also provided to facilitate ready reference.

Additions to the fungi of Madras—XIII

Article

Aug 1952

A Taxonomic Survey of Nigerian Rust Fungi: Uredinales Nigerianensis—II

Article

May 1981

Dan O. Eboh

The second taxonomic survey of rust fungi in Nigeria is presented. Twenty fungus taxa distributed among seven genera on 20 host genera in 15 families are reported. Seventeen taxa are new for Nigeria. Most of the collections are from Nsukka environs.

A Taxonomic Survey of Nigerian Rust Fungi: Uredinales Nigerianensis—III

Article

Mar 1984

Dan O. Eboh

Results of the third taxonomic survey of rust fungi in Nigeria are presented. Thirty-one taxa distributed among thirteen genera on thirty-one species of host genera in 15 families are reported. Twenty-two taxa are new for Nigeria. Most of the collections are from Nsukka environs.

Critical Notes on Some Plant Rusts III

Article

Sep 1960

M J Thirumalachar

A Taxonomic Survey of Nigerian Rust Fungi—Uredinales Nigerianensis—I

Article

Sep 1978

Dan O. Eboh

The first exclusive taxonomic survey of the Uredinales in Nigeria is carried out. Thirty-two taxa of rust fungi are reported on ten host families collected from the Nsukka area.

A Miracle Mounting Fluid for Permanent Whole-Mounts of Microfungi

Article

Jul 1972

John L. Cunningham

New Species and Records of Uredinales from the Neotropics

Article

Nov 1993

Four new species of rust fungi are described (Puccinia ibrae, Uredo clavo, U. crusa, and U. tuitei) and five others are reduced to synonymy (Uredo macella, U. nigropuncta, U. panamensis, Uromyces cissampelidis, and U. rzedowskii). Among the 35 new host and geographic range extensions are the first North American records of Aecidium opuntiae, P. duthiae, P. mutisiae, P. puta, and U. occultus. Olivea petitiae, formerly known only from one Puerto Rican specimen, is now reported from 13 locations in six Caribbean nations. Uromyces sabineae, formerly known only from one Puerto Rican specimen, is now reported on four new host taxa from five locations in three Caribbean nations. Spermogonia and uredinoid aecia are described for the first time for Ravenelia pithecolobii, and the first systemic aecia are reported for P. dietelii.

Fungi exotici. III

Article

G.E. Massee

A contribution to the rust flora (Uredinales) of southern Africa, with an emphasis on South Africa

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