Mycol. Res.
101 (7) : 863–866 (1997) Printed in the United Kingdom
Some species of Phaeoramularia causing leaf spots in north-
eastern Uttar Pradesh, India
S A N J A Y K. S I N G H, R. K. C H A U D H A R Y A N D M E E N U
Mycology Laboratory, Department of Botany, University of Gorakhpur, Gorakhpur – 273 009 (U.P.), India
Three species of Phaeoramularia, P. indica sp. nov., P. moracearum sp. nov. and P. psidii-guajavae sp. nov. on Acalypha indica, Ficus
religiosa and Psidium guajava, respectively, are described and illustrated from material collected in Gorakhpur and the adjoining area.
The foliicolous hyphomycete genus Phaeoramularia remained
relatively small for well over a decade, after it was established
by Muntan4 ola (1960), until 21 additional species were included
in it by Deighton (Ellis, 1976). Since then, the number of taxa
included has increased and it now includes 82 species,
including eight recently described (Singh, Chaudhary &
Morgan Jones, 1995, 1996 ; Singh & Singh, 1997).
Species in the genus were previously treated in Cercospora
but on account of certain distinctive features, e.g. pigment,
non-acicular and catenate conidia, it was established as an
independent segregate at generic rank within the Cercospora
complex. However, some doubts on its distinctness exist
perhaps on the stability of catenation which has already been
discussed by Sutton (pers. comm.) for Phaeoramularia calotro-
pidis (Eliis & Everh.) Kamal, A. S. Moses & R. Chaudhary
(1990) and also by Deighton (1976) for P. occidentalis (Cooke)
Deighton. The emergence of conidiophores through stomata
does not appear to be a consistent feature as in one of the
species described below the stromata are not present in sub-
stomatal cavities. This is in agreement with the observations
of Mehrotra & Verma (1991) for Phaeoramularia ougeiniae
M. D. Mehrotra & R. K. Verma.
This paper describes and illustrates Phaeoramularia indica
sp. nov., P. moracearum sp. nov. and P. psidii-guajavae sp. nov.
MATERIALS AND METHODS
Materials were collected from the forests of Gorakhpur and
the adjoining area during Nov.–Dec. 1994. Scrape mounts and
hand-cut sections were made from the fresh collections and
mounted in lactophenol and cotton blue mixture for
morphotaxonomic characterization. Illustrations were pre-
pared with the help of a camera lucida and taxonomic
determinations were made with the help of available literature.
The holotype collections were deposited in IMI.
863
RESULTS AND DISCUSSION
Phaeoramularia indica S. K. Singh, R. K. Chaudhary &
Meenu sp. nov. (Figs 1–5)
Maculae amphigenae, circulares vel subcirculares, usque ad 10 mm
diam., luteae, interdum cinereae. Caespituli hypophylli, pallide grisei.
Mycelium immersum. Stromata substomatalia, subcuticularia vel
superficialia posita, pseudoparenchymatica, parva, brunnea, 10–15 µm
diam. Conidiophora fasciculata, 4–8 in fasciculo, per stomata
emergentia (fasciculi parvi vel modici), interdum densa, 0–2
geniculata, 0–4 septata, ramosa, recta vel flexuosa, laevia, sicca,
pallide brunnea, cylindrica vel subcylindrica, usque ad
5±0–44±5¬2±5–5±5 µm. Cellulae conidiogenae in conidiophoris incor-
poratae, terminales vel intercalarales, sympodiales, polyblasticae,
palliodores versus regionem conidiogenam, cicatricatae ; cicatricibus
conidialibus conspicuis. Conidia acropleurogena, interdum acrogena,
solitaria vel catenata, laevia, sicca, ramosa, leniter recta vel curvata,
subcylindrica vel obclavata, ad 1–7 septata interdum constricta ad
septa, pallide brunnea, hilo incrassato et conspicuo, usque
9±5–136¬3±0–5±5 µm.
In foliis vivis Acalyphae indicae (Euphorbiaceae), Gorakhpur, U.P.,
India, Nov.–Dec. 1994, Kamal, IMI 366370 holotypus, GPU 4015
isotypus.
Leaf spots amphigenous, almost circular to sub-circular, up to
10 mm diam., yellowish, sometimes light grey, surrounded by
a somewhat yellowish ring. Caespituli hypophyllous, light
grey. Mycelium internal, stromata substomatal, subcuticular to
superficial, pseudoparenchymatous, brown, 10–15 µm diam.
Conidiophores fasciculate, arising from stomata, fascicles small
to medium and divergent, somewhat dense, 0–2 geniculate,
0–4 septate, branched, straight to slightly flexuous, smooth,
light brown, cylindrical to sub-cylindrical,
5±0–44±5¬2±5–5±5 µm. Conidiogenous cells integrated, terminal
to intercalary, sympodial, polyblastic (mostly bi-blastic), paler
near the conidiogenous region, cicatrized, scars thickened
and conspicuous. Conidia acropleurogenous, sometimes acro-
genous, solitary to catenate, smooth, dry, branched, somewhat
straight to curved, sub-cylindrical to obclavate, slightly
Species of Phaeoramularia from India 864

and smaller conidia. Although the conidiophores of P. indica

bear some resemblance to those of, for example, P. euphorbiae,

20 mm
other details indicate that they are distinct and different from
P. indica.
Six species of Cercospora and a species of Ramularia were
5
earlier described on the host genus. Four of the names in
Cercospora have been recombined into other genera, namely,
Pseudocercospora (Yen et al., 1980 ; Deighton, 1976) and two
are retained in Cercospora sensu lato (Pavgi et al., 1964 ; Pollack,
1987). Ramularia acalyphae has been recombined as Myco-
4 3 vellosiella acalyphae (Tharp) U. Braun (1993).

20 lm
Phaeoramularia moracearum S. K. Singh, R. K. Chaudhary
& Meenu sp. nov. (Figs 6–9)
2
Maculae amphigenae, angulares, saepe a venis foliorum limitatae,
griseae vel albidae. Caespituli hypophylli, densi, punctiformes, griseo-
albi. Mycelium immersum, subhyalinum, septatum, ramosum. Stromata
parce evoluta, ex hyphis inflatis, subhyalinis composita, elongata vel
1 ovoidea, medio brunnea, 1±5–4±0 µm lata. Conidiophora 6–8 in
fasciculo, non ramosa pallide brunnea, obscure septata (1–2), laevia,
non ramosa, recta vel flexuosa, 1–2 geniculata,
42±5–54±0¬2±5–5±5 µm. Cellulae conidiogenae in conidiophoris incor-
poratae, terminales, polyblasticae, sympodiales, cicatricatae, cica-
tricibus incrassatis. Conidia acropleurogena, solitaria vel raro catenata,
sicca, recta vel leniter curvata, subcylindrica vel obconico truncata,
basi obconica, ad apicem subacuta vel leniter rotundata, subhyalina
Figs 1–5. Phaeoramularia indica sp. nov. Fig. 1. Stromata. Fig. 2.
vel pallide brunnea, 0–4 septata, ad hilum leniter incrassata, breviter
Conidiophores. Fig. 3. Conidia. Fig. 4. Ramoconidium. Fig. 5. Leaf
attenuata, 10–98±5¬2±0–4±5 µm.
spots.
In foliis vivis Fici religiosae (Moraceae), Gorakhpur U.P., India,
Nov.–Dec. 1994, Kamal, IMI 366371 holotypus, GPU 4016 isotypus.

narrower towards the base, apex sub-obtuse to somewhat
rounded, 1–7 septate, occasionally constricted at the septa,
light brown, bearing thickened hila, 9±5–136¬3±0–5±5 µm.
No species of Phaeoramularia has yet been reported either
on the host genus or on the host species. Only three species
of Phaeoramularia, P. ajerikari (Syd.) Deighton (1976),
P. euphorbiae Q. X. Ge, X. Liu, T. Xu & Y. L. Guo (Liu & Guo,
1982) and P. manihotis (F. Stevens & Solheim) M. B. Ellis
(1976) are reported on the host family. The present collection,
therefore, is compared with these in Table 1.
It is evident from the table that all of these species are quite
different from P. indica in having much longer conidiophores
Leaf spots amphigenous, angular, often vein limited, greyish-
white. Caespituli hypophyllous, dense, punctiform, greyish-
white. Mycelium internal, sub-hyaline, septate, branched.
Stromata subepidermal, loosely prosenchymatous, elongated
to sub-spherical or somewhat ovoid, mid-brown, cells
1±5–4±0 µm wide. Conidiophores in loose to dense fascicles
(mostly 6–8 per fascicle), arising from stromata through
ruptured epidermis, unbranched, pale brown, sparsely septate
(1–2) but not distinctly so, smooth, unbranched, straight to
flexuous, 1–2 geniculate, 42±5–54±0¬2±5–5±5 µm. Conidio-
genous cells integrated, terminal, polyblastic, sympodial, slightly
paler in conidiogenous region, cicatrized, bearing usually 3–4

Table 1. Comparison between species of Phaeoramularia reported on Euphorbiaceae and Phaeoramularia indica

Infection}fruiting Stromata Conidiophores Conidia

P. ajerikari Hypophyllous Almost prosenchymatous Compact fasciculate, up to 1–4 septate, up to

60¬4–7 µm, unbranched 30–65¬4–7 µm
P. manihotis Amphigenous, mostly Feebly developed Fasciculate, 50–200¬4–6 µm, Frequently in branched chains,
hypophyllous occasionally branched 1–3 septate, up to
15–45¬6–8 µm
P. euphorbiae Amphigenous, fruiting almost 25±6–65±0 µm wide ca 5–44, fasciculate, 0–2 septate, 1–6 septate,
hypophyllous 11±3–50¬3±8–6±3 µm, 18±8–91±3¬2±8–7±5 µm
unbranched
P. indica Amphigenous 10–15 µm wide ca 4–8, fasciculate, 0–4 septate, Very rarely in branched chains,
5–44±5¬2±5–5±5 µm, 1–7 septate,
branched 9±5–136¬3–5±5 µm
S. K. Singh, R. K. Chaudhary and Meenu 865

20 mm
20 mm
8
15

13
9

20 lm
20 lm
14 12
7

6 11

10

Figs 10–15. Phaeoramularia psidii-guajavae sp. nov. Fig. 10. Stromata.

Figs 6–9. Phaeoramularia moracearum sp. nov. Fig. 6. Stromatic cells.
Fig. 11. Conidiophores. Fig. 12. Branched conidiophore. Fig. 13.
Fig. 7. Conidiophores. Fig. 8. Conidia. Fig. 9. Leaf spots.
Conidia. Fig. 14. Ramoconidium. Fig. 15. Leaf spots.

thickened scars, up to 1 µm diam. Conidia acropleurogenous, Phaeoramularia psidii-guajavae S. K. Singh, R. K.

mostly solitary, rarely catenate, dry, straight to slightly
curved, sub-cylindrical to irregularly obconicotruncate, apex
sub-acute or sub-obtuse to blunt, sub-hyaline to pale brown,
0–4 septate, sometimes slightly attenuating at both ends,
bearing mostly one (rarely 2) thickened hila,
10–98±5¬2±0–4±5 µm.
There is no published record of a Phaeoramularia on the
host family. However, Phaeoramularia ficina S. K. Singh &
R. K. Chaudhary recorded from Ficus heterophylla, has been
described elsewhere (Singh et al., 1996). The present species is
compared with it in Table 2.
Conidiophores of Phaeoramularia ficina are distinct in being
unbranched. Conidia of P. ficina are much shorter than those
of P. moracearum and have a larger number of septa (1–14).
Furthermore, P. ficina is characterized by forming ramoconidia
which are absent in P. moracearum.
Chaudhary & Meenu sp. nov. (Figs 10–15)
Maculae amphigenae, fere circulares vel irregulares interdum
coalescentes, plerumque 20 mm diam., atrogriseo-brunneae. Caespituli
hypophylli. Mycelium immersum ex hyphis septatis, ramosis,
subhyalinis compositum. Stromata parva, bene evoluta, pseudo-
parenchymatosa, substomatalia, ad 15–18±5 µm diam., ex hyphis
inflatis, pallide brunnea, circulares vel leniter elongata composita.
Conidiophora 5–10 in fasciculo, densa, recta vel leniter flexuosa,
divergentia, obscure septata 0–2, laevia, raro ramosa, subhyalina,
plerumque 1–2 geniculata, cylindrica vel subcylindrica,
11±0–27±5¬2±0–4±5 µm. Cellulae conidiogenae in conidiophoris incor-
poratae, terminales, sympodiales, polyblasticae, cicatricatae, cica-
tricibus conidialibus incrassatis, 1±5 µm diam. Conidia acropleurogena,
interdum acrogena, solitaria, raro catenata, sicca, laevia, recta vel
leniter curvata, raro ramosa, 1–4 septata, constricta ad septa,
subcylindrica vel obconicotruncata, ad apicem subobtusa vel leniter
rotundata, subhyalina, hilo leniter incrassata, 20–87±5¬2±5–3±5 µm.

Table 2. Comparison between Phaeoramularia ficina and P. moracearum

Infection}fruiting Stroma Conidiophores Conidia

P. ficina Hypogenous, light brown 14–16 µm wide Fasciculate (3–5), branched, 0–2 Branched, 1–14 septate,
septate, 13±5–50±0¬2±5–5±0 µm 18±5–58±5¬1±5–3±5 µm
P. moracearum Amphigenous, greyish white Loosely arranged In dense fascicles (6–8), sparsely Unbranched, (0–) 2–3 (–4) septate,
(1–2) septate, 10–98±5¬2±0–4±5 µm
42±5–54±0¬2±5–5±5 µm
Species of Phaeoramularia from India
Table 3. Comparison of Phaeoramularia barringtoniae with Phaeoramularia psidii-guajavae
Infection}fruiting Stroma
Phaeoramularia Not distinct, amphigenous, 15–55 µm diam.
barringtoniae abundant on the upper surface,
brown
Phaeoramularia Distinct, amphigenous, forming 15–18 µm diam.
psidii-guajavae greyish brown blotches
In foliis vivis Psidii guajavae (Myrtaceae), Gorakhpur, U.P., India,
Nov.–Dec. 1994, Kamal, IMI 366372 holotypus, GPU 4017 isotypus.
Leaf spots amphigenous, almost circular to irregular, coalescing
to form blotches up to 20 mm diam., dark greyish-brown.
Caespituli hypophyllous. Mycelium internal, septate, branched,
sub-hyaline. Stromata small, well developed, pseudo-
parenchymatous, sub-stomatal, 15–18±5 µm diam., composed
of somewhat swollen hyphae, pale brown, bearing sub-circular
to somewhat elongated or ovoid cells. Conidiophores densely
fasciculate, 5–10 in each fascicle, straight to somewhat
flexuous, divergent, indistinctly 0–2 septate, smooth, rarely
branched, sub-hyaline, 1–2 geniculate, cylindrical to sub-
cylindrical, 11±0–27±5¬2±0–4±5 µm. Conidiogenous cells inte-
grated, terminal, sympodial, polyblastic, cicatrized, bearing
1–2 thickened scars up to 1±5 µm diam. Conidia acrop-
leurogenous, sometimes acrogenous, mostly solitary, rarely
catenate, dry, smooth, straight to slightly curved, unbranched
to rarely branched, 1–4 septate, occasionally constricted at the
septa, sub-cylindrical to somewhat obconico-truncate, apex
sub-obtuse to more or less rounded, sub-hyaline to slightly
pigmented, the base bearing thickened hila,
20–87±5¬2±5–3±5 µm.
A survey of the literature indicates that there is no record
of a Phaeoramularia on this host. One species, Phaeoramularia
barringtoniae (Syd. & P.Syd.) X. J. Liu & Y. L. Guo (Liu & Guo,
1982), has been reported on the host family and the present
fungus is compared with this species in Table 3.
Although there are similarities between P. barringtoniae and
P. psidii-guajavae in dimensions of conidia, the present
collection differs sufficiently in having much shorter, rarely
branched, 0–2 septate, conidiophores and poorly developed,
smaller stromata compared with P. barringtoniae which forms
longer, unbranched conidiophores with more septa (2–8
septate).
Obviously, the present collection cannot be accommodated
with the species described on Myrtaceae and its description as
a new species is justified.
Mycovellosiella myrtacearum A. N. Rai, B. Rai & Kamal (Rai
et al., 1986), also has been reported on the same host.
(Accepted 19 December 1996)
866
Conidiophores Conidia
Fasciculate, 2–8 septate, not Indistinctly 3–8 septate,
branched, sparingly geniculate, 20±0–85±0¬3±0–5±6 µm
32±5–125¬3±0–5±5 µm
Fasciculate, sparsely (0–2) septate, Indistinctly (1–) 2–3 (–4) septate,
rarely branched, 1 or 2 rarely branched and catenate,
geniculate, 11–27±5¬2±0–4±5 µm 20–87±5¬2±5–3±5 µm
However, this species belongs in another genus having
entirely different morphological characteristics.
The authors are grateful to Dr Kamal, University of
Gorakhpur, for supervision and helpful suggestions in
taxonomic determinations.
REFERENCES
Braun, U. (1993). Taxonomic notes on some species of the Cercospora
complex. III. Mycotaxon 48, 275–298.
Deighton, F. C. (1976). Studies on Cercospora and allied genera. VI.
Pseudocercospora Speg. Pantospora Cif. and Cercoseptoria Peter. Mycological
Papers 140, 1–168.
Deighton, F. C. (1979). Studies on Cercospora and allied genera. VII. New
species and redispositions. Mycological Papers 144, 1–56.
Ellis, M. B. (1976). More Dematiaceous Hyphomycetes. Kew : Commonwealth
Mycological Institute.
Kamal, Moses, A. S. & Chaudhary, R. (1990). Two new species and a new
combination in Phaeoramularia from Uttar Pradesh, India. Mycological
Research 94, 714–717.
Liu, X. J. & Guo, Y. L. (1982). Studies on some species of the genus
Phaeoramularia in China. Acta Phytopathologica Sinica 12, 1–15.
Mehrotra, M. D. & Verma, R. K. (1991). Some new hyphomycetes associated
with leaf spots of trees in India. Mycological Research 95, 1163–1168.
Mutan4 ola, M. (1960). Algunos Hyphomycetes criticos, notas y descripciones.
Lilloa 30, 165–232.
Pavgi, M. S. & Singh, U. P. (1964). Parasitic fungi from North India. I.
Mycopathologia et Mycologia applicata 23, 188–195.
Pollack, G. (1987). An annotated compilation of Cercospora names. Mycologia
Memoir No. 12.
Rai, A. N. & Kamal (1986). A new species of Mycovellosiella Rangel. Current
Science 55, 798–799.
Rai, A. N., Rai, B. & Kamal (1986). A new species of Mycovellosiella Rangel.
Current Science 55, 412–413.
Singh, S. K., Chaudhary, R. K. & Morgan-Jones, G. (1995). Notes on
Hyphomycetes. LXVII. Three new species of Phaeoramularia from Nepal.
Mycotaxon 54, 57–66.
Singh, S. K., Chaudhary, R. K. & Morgan-Jones, G. (1996). Notes on
Hyphomycetes. LXXII. Further novel leaf spot-inducing species of
Phaeoramularia from the Indian sub-continent. Mycotaxon 58, 137–145.
Singh, S. K. & Singh, A. (1997). Additions to Phaeoramularia and Stenella from
the Indian sub-continent. Mycological Research. In press.
Yen, J. M. & Lim, G. (1980). Cercospora and allied genera of Singapore and the
Malay Peninsula. Garden’s Bulletin, Singapore 33, 151–263.