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The plant family Myrtaceae includes approximately 150 genera 

with over 5 500 species (Heywood et al. 2007), amongst which

are  some  important  forestry  species  (e.g.  Eucalyptus  spp.)

and  several  fruit  crops  such  as  guava  (Psidium guajava).

Some, such as Acca sellowiana  (common  name  feijoa;  in

Brazil,  goiabeira  da  serra)  are  only  minor  fruit  crops.  Acca 

sellowiana is a shrub or small tree native to southern South 

America (southern Argentina, Brazil, Paraguay, and Uruguay)

and, although only occasionally cultivated in South America, 

it has become more significant as a fruit crop in New Zealand

(Al-Harthy 2010). There are few published records of fungal

pathogens  associated  with  feijoa  (Farr  &  Rossman  2011,

Mendes & Urben 2011). However, during a recent search for

pathogens of feijoa in the southern Brazilian state of Rio Grande

do Sul, individuals of A. sellowiana in rural areas in the vicinity 

of Pelotas had foliage with intense tar-spot symptoms. Such

symptoms were typical of those caused by fungi belonging to 

the genus Phyllachora.  Examination  of  specimens  collected

and a literature and herbarium search were performed in order 

to clarify the identity of the fungus on feijoa, and the results of

these investigations are presented here.

MAterIAl ANd Methods 

Samples of diseased foliage of Acca sellowiana were collected 

in two localities. These were dried in a plant press and taken

to  the  laboratory  for  further  examination.  Representative

specimens were deposited in the local herbarium (Herbarium

Universidade  Federal  de  Viçosa,  VIC).  Examination  of

selected leaves bearing tar-spot symptoms with the help of an 

Olympus SXZ7 stereoscopic microscope revealed that fungal

structures were immersed in the leaf tissue and sections 

were prepared and mounted in lactophenol and lactofucsin 

for  further  examination.  Additionally,  sections  were  also

prepared  with  a  freezing  microtome  (Cryostat  Microm



520).  Observations,  photographs,  and  line  drawings  were

prepared with a light microscope Olympus BX51, fitted with

a digital camera (Olympus E-volt 330) and a drawing tube.


Phyllachora feijoae Rehm, Hedwigia 36: 370 (1897).

SynonymCatacauma feijoae (Rehm) Theiss & Syd.,

Ann. Mycol. 

13: 397 (1915).

(Fig. 1)


Brazil:  Rio Grande do Sul:  Pelotas,  Chácara  da

Brigada,  Cerro  da  Buena,  on  leaves  of  Acca sellowiana 

(Myrtaceae),  18  Aug.  2010,  R. W. Barreto  (VIC  31476  –

neotype designated here; B 70 0015054 – isoneotype).

Other specimen examined

Brazil:  Rio Grande do Sul:  Pelotas,

Capão do Leão, on leaves of Acca sellowiana (Myrtaceae), 18 Aug.

2010, R. W. Barreto (VIC 31766).


IMA FuNgus · voluMe 3 · No 1: 9–14

Reappraisal and neotypification of Phyllachora feijoae

Lilian C. Costa, Davi M. Macedo, and Robert W. Barreto

Universidade Federal de Viçosa, Departamento de Fitopatologia, 36570-000, Viçosa, MG, Brazil; corresponding author e-mail: Davi M. Macedo,


 Acca sellowiana (Myrtaceae), feijoa (in Brazil, goiaba da serra), is a native southern South America tree

that produces edible fruits which, although only occasionally cultivated in South America, became a significant fruit

crop in New Zealand. Recently, during surveys for fungal pathogens of feijoa in southern Brazil, several plants were

found bearing tar-spot symptoms caused by a species of Phyllachora. A literature search enabled us to identify the 

fungus as Phyllachora feijoae, a little-known species originally described in the 19


century by H. Rehm and later

transferred to the genus Catacauma. The name Catacauma feijoae, although now regarded as a later synonym of P. 

feijoae is still mistakenly in use (as, for instance, in the Brazilian list of fungi on plants). The type specimen was most

probably deposited in the Botanisches Garten und Museum Berlin-Dahlem (B) and lost or destroyed during World War

II, and could not be located. The recent recollection of abundant material of this fungus in the vicinity of Pelotas (Rio

Grande do Sul, Brazil) allowed its re-examination and neotypification. Phyllachora feijoae is also illustrated here for 

the first time.

Article info:

Submitted: 6 November 2011; Accepted: 13 March 2012; Published: 5 April 2012.

Key words: 



fruit crop





costa, Macedo & Barreto




i m a   f u n G u S

Fig. 1. Phyllachora feijoae (VIC 31766). A. Tar-spots on leaves of Acca sellowiana. B. Perithecium and clypeus. c. Ascospores. d. Paraphyses

and asci with ascospores.

e. Ascomata (left) and conidioma (right) and close up part of the conidioma. F–g. Conidiogenous cell and conidia.

Bars: B = 80 µm, C = 20 µm, D = 20 µm, E = 275 µm, F–G = 10 µm.

Reappraisal and neotypification of Phyllachora feijoae




v o l u m e   3   ·   n o .   1


Lesions  on living leaves, adaxially on all leaves at various 

developmental stages, initially punctiform, becoming irregular 

tar-spots, raised, with age surrounded by yellowish to reddish 

peripheral necrotic haloes,  widely distributed and leading 

to  foliage  distortions,  0.2–0.3  ×  2.1–3.0  mm  diam,  indistinct

abaxially.  Internal mycelium intra- and intercellular, hyphae 

2.0–3.0  μm  diam,  branched,  septate,  hyaline  to  pale  brown.

External mycelium absent. Stromata adaxial, clypeate, shield-

like, merged with the upper wall of the ascoma. Conidia formed 

within stromata externally indistinguishable from teleomorph 

stomata; flattened, lenticular to irregular (in section), epigenous,

subepidermal, single or in combination with ascomata, 

sometimes very broad occupying nearly the whole breadth of 

the stroma, 615–1729 x 100–184 µm walls of dark brown textura 

angularis,  38.5–69  µm  thick,  smooth;  conidiogenous cells 

subcylindrical, straight, 15–25(–40) x 2–3 µm, 0–1-septate, pale

brown;  conidia mucilaginous, enteroblastic, acicular, curved, 

lunate or sigmoid, 13–19 x 1.5 µm, aseptate, thin-walled, hyaline,

smooth.  Ascomata perithecial, epigenous, immersed, solitary, 

spherical to subsphaerical, somewhat to strongly depressed, 

short  papillate,  41–218  μm  diam,  inconspicuously  ostiolate, 

composed of thin-walled brown textura angularis, walls 6.5–44

µm, 7–11 cells thick, outer layers dark brown, inner layers pale

brown  to  subhyaline.  Interascal tissue of  paraphyses,  2.5–3

μm diam, longer than the asci, filiform, septate, hyaline, thin-

walled,  constricted  at  the  septae;  periphyses  well-developed,

filiform,  hyaline,  thin-walled.  Asci unitunicate, cylindrical to 

clavate,  short-stalked,  70.5–104  x  13–27  μm,  apex  broadly

rounded to nearly flat, thin-walled, 8-spored. Ascospores at first

uniseriate but sometimes partially biseriate, 15.5–22 x 8–14 µm,

ellipsoidal to cylindric-ellipsoidal, rounded at the ends, walls 2–3

μm thick, aseptate, hyaline, smooth, without a mucous sheath

or appendages.

Notes:  Very  little  information  is  available  on  Phyllachora 

feijoae. Only a very brief description is given in the original

publication of Rehm (1897). Later, Theissen & Sydow (1915)

prepared a more complete description of the fungus when 

combining it into Catacauma. This is, nevertheless, somewhat

incomplete and no illustrations were provided. Furthermore,

the  description  was  apparently  based  on  Rehm’s  material

collected  in  “Serra  Geral,  Minas  Gerais  –  Brazil”. The last 

publication  dealing  with  this  fungus  was  that  of  Jimenez  &

Hanlin (1992), where names of fungi described in Catacauma 

were  listed.  Although  the  authors  acknowledged  that  after

Petrak’s  (1924)  work  it  became  widely  accepted  that  the

distinction of Catacauma from Phyllachora was artificial, they

prudently did not propose that names in Catacauma should be 

immediately rejected or recombined into Phyllachora without 

a  careful  re-examination  of  types.  Since  that  publication,

mycologists have shown little interest in the names of fungi 

referred to Catacauma,  but some earlier fungal names in 

Phyllachora  have  been  reinstated.  That is the case of the 

name  C. feijoae, presently listed in MycoBank and Index 

Fungorum as a later synonym of P. feijoae. Nevertheless, this

name is still being used in other instances (e.g. the Brazilian

list of fungi on plants; Mendes & Urben 2011).

An expanded description based on the material recently 

collected  in  Brazil  is  provided  above.  This  is  also  the  first

time illustrations of P. feijoae have  been  published.  The

original  material  of  the  species  studied  by  Rehm  would

almost certainly have been deposited in the collections of the 

Botanisches Garten und Museum Berlin-Dahlem (B), but if so

it appears to have been lost or destroyed during World War II

as it could not now be found (H.J.M. Sipman, pers. comm.).

We  therefore  designate  one  of  the  recent  collections  as  a

neotype to fix the application of the name.


The fungus on Acca sellowiana exhibits all the typical 

features, both in terms of symptoms produced on the host 

and in its morphology, to members of the genus Phyllachora 

(Phyllachoraceae,  Phyllachorales).  Phyllachora is a large 

genus  including  approx.  1000  named  species  (Kirk  et 

al.  2008).  All  species  of  Phyllachora  are biotrophic plant 

pathogens, causing tar-spots on members of numerous plant 

families, but are particularly common on Fabaceae (Cannon

1991) and Poaceae (Parbery 1971). Besides the presence of

a well-developed, dark brown to black clypeus, other features 

such as the formation of the perithecia within the plant tissues, 

and hyaline, thin-walled, smooth and aseptate ascospores, 

are typical for the genus (Cannon 1991). Around 70 species of

Phyllachora have been described on members of Myrtaceae 

worldwide (Farr & Rossman 2011), with 21 species recorded

on this host-family in Brazil (Mendes & Urben 2011). Species

of Phyllachora associated with Myrtaceae have never been 


Some of the older records of Phyllachora on Myrtaceae 

were  later  recognized  as  mistakenly  placed  in  that  genus.

Some were found to belong to other genera, - such as P. 

pululahuensis (now regarded as a synonym of Vestegrenia 

multipunctata; von Arx & Müller 1954), and P. eucalypti (now 

recognized as  a synonym of Clypeophysalospora latitans;

Crous  et al. 1990).  Other  species  were  recombined  into

genera such as P. peribebuyensis which is now treated as 

Coccodiella peribebuyensis (Katumoto 1968). Several names

in  Phyllachora  that are listed on members of Myrtaceae 

were found to be later synonyms of already known species 

names:  P. conspurcata  (syn.  P. tropicalis

Saccardo 1883),

P. phylloplaca (syn. P ipirangae; Theissen & Sydow 1915b),

P. pseudostromatica  (syn.  P. melaleuca;  Sydow  &  Sydow

1904),  and  P. semillunata  (syn.  P. selenospora;  Petrak  &

Ciferri  1930).  Additionally,  P. langdonii is now treated as 

a subspecies of P. callistemonis,  P. callistemonis subsp. 

langdonii (Pearce & Hyde 1994).

In the case of species of Phyllachora recorded from Brazil, 

an issue to be taken into consideration is that numerous 

species names are included in Mendes et al. (1998), and have

also been kept in the database of fungi on plants in Brazil 

(Mendes & Urben 2011) but quoted as being “in press”. These

names, for which Medeiros & Dianese are given as authors,

have never been validly published and include the following 

costa, Macedo & Barreto




i m a   f u n G u S

table 1. Data on Phyllachora spp. described on hosts belonging to Myrtaceae.


Asci (µm)

Ascospores (µm)  host plants 


P. ambígua

50–60 x 8–12

9–11 x 6

Syzygium cumini (syn. Eugenia 


Theissen & Sydow (1915b)

P. angustispora

80–90 x 12–14

30 x 8–9

Eugenia sp.

Saccardo (1916)

P. bella

60–70 x 5–7

7.5–9 x 3–4

Syzygium australe (syn. E. australis)  Sydow (1937)

P. biareolata

90–95 x 6–9

12 x 5

Eugenia rhombea

Saccardo (1891)

P. biguttulata

50–65 x 8–10

10–12 x 5–5.5

Campomanesia rhombea 

Saccardo (1913)

P. brenesii

70–80 x 10–15

12–17 x 8–10

Eugenia guayaquilensis

Sydow & Petrak (1929)

P. callistemonis

115–210 x 12.5–16 18–27.5 x 7.5–10

Callistemon pallidus 

Pearce & Hyde (1994)

P. callistemonis subsp. Langdonii 100–154 x 12–20

18–25 x 6–9

Callistemon sp.

Pearce & Hyde (1994)

P. callistemonis subsp. Similis

117–173 x 18–27.5 18–29 x 7.5–12.5

Callistemon viminalis 

Pearce & Hyde (1994)

P. capensis

100–120 x 13–14

60–70 x 5–6

Eugenia zuluensis 

Doidge (1942)

P. cayennensis 

68–75 x 12–14

20–24 x 7–8

Psidium sp.

Theissen & Sydow (1915b)

P. clavata

110–140 x 15–18

39–45 x 3–6

Myrcia sp.

Garces Orejuela (1944)

P. curvulispora

60–80 x 10–20

17–20 x 5–7

Myrtaceae sp.

Saccardo (1925–1928)

P. distinguenda

60–70 x 18

18–20 x 4.5

Myrtaceae sp.

Saccardo (1899)

P. egenula

70–85 x 7–8

10–13 x 5-6

Leptospermum lanigerum 

Sydow (1938)

P. emarginata

80–130 x 18–28

16–20 x 10–12

Eugenia sp.

Petrak (1948)

P. eugeniae

60–75 x 7–9

8–10 x 4–4.5

Eugenia rhombea 

Chardón (1927)

P. feijoae

60 x 25

18 x 10

Acca sellowiana

Rehm (1915)

P. gentilis

120 x 4–12

18–20 x 8–9

Eugenia sp.

Saccardo (1895)

P. goyazensis

70–90 x 17–18

12–14 x 8–12

Myrtaceae sp.

Hennings (1895)

P. guavira

100–110 x 6–8

12 x 5

Psidium sp.

Theissen & Sydow (1915b)

P. ipirangae

30–90 x 10–12

15–16 x 8

Eugenia sp.

Theissen & Sydow (1915b)

P. lindmanii

80–90 x 13–16

16–24 x 13–16

Myrtaceae sp.

Theissen & Sydow (1915a)

P. maculata




Eucalyptus sp.

Cooke (1891)

P. manuka

10.5–13 × 6.5–8

Leptospermum scoparium 

Johnston & Cannon (2004)

P. melaleucae

66–84 x 8–11

Melaleuca spinosa 

Theissen & Sydow (1915a)

P. myrciae


Eugenia bimarginata

Saccardo (1883)

P. myrciae-rostratae

100–120 x 6–8

14–17 x 5–6

Myrcia splendens (syn. M. rostrata)  Viégas (1944)

P. muelleri

95–120 x 13–15

28–32 x 6–7

Eugenia dodonaeifolia 

Chardón et al. (1940)

P. myrrhinii

50–72 x 12–16

14–16 x 5

Myrrhinium atropurpureum var.


Theissen & Sydow (1915a)

P. nigerrimum

100–130 x 9

10–16 x 5

Campomanesea adamantium (syn.

C. caerulea)

Viégas (1944)

P. opaca

80–85 x 6–8

10 x 4–4.5

Myrtaceae sp. 

Berlese & Voglino (1886)

P. peglerae

120–140 x 17–20

20–23 x 12–13

Eugenia capensis 

Doidge (1942)

P. pettimenginii

85–105 x 14–18

2–8-32 x 8.5–11

Myrtaceae sp.

Maire (1908)

P. rhytismoides

14–19.5 × 12–15.5 Melaleuca cajuputi

Cannon (1991)

P. rickiana

68–78 x 14–15

10–13 x 6

Myrtaceae sp.

Theissen (1918)

P. rimulosa

85–100 x 10

14 x 8

Eugenia sp.

Saccardo (1925–1928)

P. samanensis

70–83 x 13–16.5

32–40 x 6–7.5

Eugenia sp.

Petrak & Ciferri (1932)

P. shivasii

136–225 x 10–15

15–22 x 6–8.5

Melaleuca viridiflora 

Pearce & Hyde (1995)

P. subcircinans

80–90 x 10–16

14–16 x 8–10

Psidium grandifolium 

Viégas (1944)

P. subopaca

75 x 10–15

12–14 x 7

Myrtaceae sp.

Saccardo (1899)

P. tachirensis

109–166 x 9.5–12 13–17 x 7–8

Eugenia sp.

Chardón & Toro (1934)

P. tropicalis

70–75 x 10–14

15–18 x 7–8

Psidium grandifolium 

Saccardo (1883)

P. truncatispora

70–90 x 16–24

22–26 x 7–8

Myrtaceae sp.

Viégas (1944)

P. urbaniana

70–90 x 16–18

14–15 x 6–8

Myrtaceae sp. 

Saccardo (1899)

P. verrucosa 

78–105 x 15–19

14–20 x 9–13

Melaleuca leucadendra 

Arx & Müller (1954)

Reappraisal and neotypification of Phyllachora feijoae




v o l u m e   3   ·   n o .   1


species designations associated with members of Myrtaceae

P. eugenii-complicatae, P. eugenii-punctifolie, P. myrciae-

decrescentis,  P.  myrciae-guianensis,  P.  myrciae-multiflorae, 

P.  myrciae-multiflorae,  P.  myrciae-pallescentis,  P.  myrciae-

tematae, P. myrciae-tortae, and  P. pampulhae.  Although  all

these designations are not validly published, most may well 

represent good taxonomic species which are still awaiting 

formal description. Most were collected in the Brazilian cerrado,

an area rich in endemic organisms of all kinds.

A  study  of  the  48  published  descriptions  of  taxa

(including  three  varieties)  of  Phyllachora  described from 

hosts belonging to Myrtaceae,  is  summarized  in  Table  1.

This shows that there are three species of Phyllachora with 

close morphological similarity to P. feijoae on A. sellowiana

P. brenesii,  P. emarginata, and  P. subcircinans.   Each  of

those species was found to have morphological differences 

from  P. feijoae.  Phyllachora. brenesii has perithecia with 

narrower  walls  (5  µm  thick),  and  asci  which  are  also

narrower  (10–15  µm  wide).  Phyllachora emarginata has 

thinner  ascospore  walls  (2  µm).  And  P. subcircinans has 

much  wider  perithecia  (250–500  µm  diam).  Additionally,

P. feijoae can be recognized as distinct from the other 

species known on Myrtaceae (Table 1) by a combination of

morphometric features; differences in perithecial diameter,

ascus width, and the absence of a mucilaginous sheath on 

the ascospores. Although no comparison of the morphology

of P. feijoae with other species on Myrtaceae was attempted 

in previous publications, our results indicate that this 

species is distinct from other Phyllachora  species on this 

host-family, and so deserves recognition as a separate 

species. No significant discrepancies were found between

the morphology of the neotype and the description provided 

in Theissen & Sydow (1915).


We acknowledge financial support from the Fundação de Amparo a

Pesquisa de Minas Gerais (FAPEMIG) and Conselho Nacional de

Desenvolvimento  Científico  e  Tecnológico  (CNPq)  and  also  would

like to thank Harrie J. M. Sipman for providing information on material

now held in the herbarium Botanisches Garten und Museum Berlin-



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