Two new species of Appendiculella (Meliolaceae) from ... - Mycologia

Mycologia, 99(4), 2007, pp. 544–552.
# 2007 by The Mycological Society of America, Lawrence, KS 66044-8897
Two new species of Appendiculella (Meliolaceae) from Panama
Délfida Rodríguez J.
Meike Piepenbring
Institute of Ecology, Evolution and Diversity,
J.W. Goethe-Universität Frankfurt am Main,
60054 Frankfurt am Main, Germany
Abstract: Two new species of Meliolaceae (black
mildews) are described based on specimens recently
collected in western Panama. Appendiculella lozanellae
on leaves of Lozanella enantiophylla is the first species
of Appendiculella known on Cannabaceae. Appendiculella
chiriquiensis on leaves of Cupania guatemalensis
is the first record of a species of Appendiculella on
Sapindaceae. These species differ from known species
on their respective host relationships by the presence
of larviform appendages attached to the perithecia, as
well as by characteristics of hyphae, appresoria and
ascospores. Sequence data of 18S and 28S rDNA are
published for A. lozanellae, which becomes the third
species of Meliolales and the first species of the genus
Appendiculella for which molecular data are available.
Key words: Cannabaceae, Cupania guatemalensis,
Lozanella enantiophylla, plant parasitic fungi, Sapindaceae,
systematics, taxonomy, Ulmaceae
INTRODUCTION
Species of Meliolaceae (about 1583 species in 20
genera, Kirk et al 2001; Meliolales, Ascomycota) are
tropical plant parasitic fungi infecting plant species
belonging to numerous families. Because of their
dark color and their growth on leaves and stems they
are called black mildews or dark mildews. They are
characterized by superficial, dark, thick-walled,
branching hyphae with appresoria and conidiogenous
cells. Appresoria adhere to the cuticle of leaves of
the host, penetrate the host cell wall and form
haustoria inside the host cell. Conidiogenous cells
(also called mucronate hyphopodia) are phialides
and produce rarely observed small spores of unknown
function. These spores might contribute to asexual
multiplication or sexual reproduction (Luttrell 1989,
Mueller et al 1991). Attempts to grow species of
Meliolaceae in culture have not succeeded. In
addition to appresoria and conidiogenous cells, the
dark hyphae develop superficial, dark perithecia
Accepted for publication 22 June 2007.
1 Corresponding author. E-mail: delfidaro@yahoo.es
544
containing clavate asci with thin walls. Asci typically
contain two brown ascospores with four septa each.
Setae can be attached to hyphae (Meliola) or can
arise from perithecia (Irenopsis). The position of
setae, wall structure and tips are morphologically
diverse among species of Meliolaceae. Therefore
details of position and structure of setae are important
tools to distinguish genera and species. Species of
the Meliolaceae traditionally have been assigned to
one of four genera on the basis of presence or
absence of setae and their form and location. Setae of
Meliola Fries, the largest genus of Meliolaceae with
about 1000 species, are attached to hyphae (mycelial
setae). Setae of Irenopsis F. Stevens (ca. 50 species)
arise from perithecia. Species of Asteridiella McAlpine
(ca. 250 species) lack setae, and species of Appendiculella
Höhn. (ca. 250 species) have perithecia with
larviform appendages (Hansford 1961, Kirk et al
2001). Larviform appendages differ from setae, being
aseptate and having thin, light brown, transversely
striate walls. The genus Appendiculella was proposed
by Höhnel (1919, type species A. calostroma [Desm.]
Höhnel on Rubus fruticosus L. [Rosaceace]) to
separate species of Irene Theiss. & P. Syd. (today
a synonym of Asteridiella) with larviform appendages
from species without appendages.
Species of the Meliolaceae traditionally have been
defined as being specific to host family (Hansford
1961). This hypothesis has never been analyzed
critically but is workable and likely to be adequate
because species of Meliolaceae interact with living
cells of the plants. This biotrophic process requires
specific physiological and molecular interactions.
Observations in the field show that species of black
mildews infect species of a single genus or of closely
related genera belonging to the same family (Rodríguez
2001; pers obs). For example Meliola mangiferae
Earle is known growing on leaves of two species of
the genus Mangifera (Anacardiaceae) (viz. Mangifera
indica L. [Amboin, Brunei, China, Colombia, Costa
Rica, Cuba, Dominican Republic, Guyana, Honduras,
India, Indonesia, Jamaica, Java, Malaysia, Panama,
Papua New Guinea, Peru, Philippines, Puerto Rico,
Singapore, Surinam, Taiwan, Trinidad and Tobago,
Venezuela, Virgin Islands fide Hansford 1961,
Schmiedeknecht 1989, Rodríguez and Minter 1998,
Farr et al 2006] and M. andamica Kinq. [Andaman
Islands fide Farr et al 2006]). But this species grows
also on leaves of Anacardium occidentale L. (Anacardiaceae)
in Costa Rica and Panama (Farr et al 2006).
On the other hand species of Meliolaceae are known

RODRÍGUEZ AND PIEPENBRING: NEW APPENDICULELLA SPECIES 545
to infect only one host species (e.g. Meliola anacardii
Zimm. on leaves of Anacardium occidentale L. in
Brunei, Costa Rica, Cuba, Dominican Republic,
Guyana, India, Malasia, Philippines fide Hansford
1961, Farr et al 2006).
Some species of Meliolaceae infect plants belonging
to a wide range of genera within a given, large
family. Meliola bicornis Wint. for example has been
reported from species of 29 genera of the Fabaceae
(Rodríguez 2006). Other species are known only from
one or a few host species. Few species are reported
from members of different families (Hansford 1961),
but these reports have not been subjected to a critical
revision.
DNA sequence data will help to resolve the
significance of host specificity to species delimitation.
Although mycologists in several laboratories tried to
isolate DNA from Meliolaceae (e.g. D. Triebel pers
comm) only a few attempts have been successful.
Because no cultures exist cells must be taken from
leaves that frequently are contaminated by numerous
other fungi. The dark pigment of the thick cell walls
also might interfere during the isolation of DNA.
Only Saenz and Taylor (1999) have published
sequence data for Meliolaceae, analyzing the phylogenetic
relationship between Meliola and other
ascomycetous fungi. They published the two existing
sequences of 18S rDNA of species of Meliolaceae
available in GenBank belonging to Meliola juddiana
F. Stevens and M. niessleana Wint. After numerous
attempts we obtained sequence data for a third
species of Meliolaceae, the new species Appendiculella
lozanellae.
In a monumental work Hansford (1961, 1963)
presented descriptions and illustrations of all thenknown
species of Meliolales. Many species were added
later, especially for India (e.g. Katumoto and Hosagoudar
1989, Hosagoudar et al 1994, 1997, 2001,
Hosagoudar 1996, 2002, Biju et al 2005) and China
(e.g. Katumoto and Hosagoudar 1989, Yang 1989,
Song et al 1996, 1997, 2003, Song 1998, Song and Li
2003, 2004).Today India has the highest number of
known species of Meliolaceae (ca. 461) followed by
China (with ca. 345) (Rodríguez 2006). Meliolaceae
from tropical America have been studied by Schmiedeknecht
(1989), Chacón and Cruz (1999) and
Rodríguez and Minter (e.g. 1998, 1999). From Brazil
we know a maximum of 240 species (Rodríguez
2006). For Panama, in the southern part of the
Central American isthmus, ca. 100 species have been
recorded, mainly by Stevens (1927, 1928). Because of
Panama’s high diversity of plant species (9520, Correa
et al 2004) many more species of Meliolaceae are
presumed to exist in this country. Here we describe
two new species of Appendiculella from Panama.
MATERIALS AND METHODS
Two new species were collected in 2004 during a survey of
the Meliolaceae in Chiriquí Province in western Panama.
Hyphae and perithecia were mounted in water or 5%
KOH for light microscopy (LM). Sizes of structures are
based on 20 measurements for ascospores and 10 measurements
for other cellular structures. For the observation
of larviform appendages perithecia were mounted in
water because the appendages become detached in KOH.
Longitudinal sections of ascomata, ca. 20 mm thick, were
made with a freezing microtome. Semipermanent preparations
of the sections were made by placing them directly
in a droplet on a microslide of this solution: distilled water
(60 mL), lactic acid (35 mL), glycerine (10 mL), polyvinyl
alcohol (10 g), chloral hydrate (50 g) and stain-blue water
(0.015 g). Air-dried specimens were observed by scanning
electron microscopy (SEM) with a Hitachi S-4500 at 5–20 kV
after being sputtered with fine gold for 60 s.
For the isolation of DNA numerous perithecia were taken
from leaves. In a vibratory mill the perithecia were triturated
1.5 min at 30 Hz. Afterward DNA was isolated, subjected to
PCR with the primers NL1/NL4 or FF1/FR1 and cleaned
following standard protocols (Cáceres et al 2006). PCRproducts
were sequenced by Scientific Research & Development
GmbH in Oberursel, Germany. Sequences were
edited and compared to sequences deposited in GenBank
with the BLAST search function.
TAXONOMY
Appendiculella lozanellae D. Rodríguez Justavino &
M. Piepenbr., sp. nov. FIGS. 1–11, 16
Plagulae hypophyllae, minus densae usque ad 2–6 mm
diam. Hyphae atrofuscae, septatae, leviter rectae usque ad
leviter undulatae, ramificationibus oppositis, cellulae 36–69
3 6–9 mm. Appresoria alternata, recta usque ad antrorsa,
29–52 3 13–17 mm; cellula basalis cylindrica, leviter curvata,
9–25 3 7–10 mm; cellula apicalis cylindrica usque ad leviter
lobata, 18–23 3 13–17 mm. Cellulae conidiogenae commixtae
appressoriis, oppositae vel unilaterales, ampullatae, 21–
31 3 7–10 mm. Perithecia numerosa, distributa in tota
plagula, nigra, usque ad 86–257 3 86–243 mm, quaeque
appendicibus larviformis plus 15 induta, pallide fuscis,
transversaliter striatis, rectis usque ad leviter curvatis, 44–
81 3 11–20 mm. Asci ovoidei cum 2 ascosporis. Ascosporae
atrofuscae, cylindricae, apicibus rotundatis, 4-septatae,
laeves, leviter constrictae in septis, 44–52 3 16–18 mm.
Colonies on the abaxial surface of leaves, subdense,
2–6 mm diam. Hyphae dark brown, septate, substraight
or slightly undulate, branching opposite,
mycelial cells 36–69 3 6–9 mm. Appresoria alternate,
straight to antrorse, 29–52 3 13–17 mm; stalk cell
cylindrical, slightly bent, 9–25 3 7–10 mm; head cell
cylindrical to slightly lobate, 18–23 3 13–17 mm.
Conidiogenous cells ampulliform, 21–31 3 7–10 mm,
opposite or unilateral, mixed with appresoria, conidia
not observed. Perithecia numerous, uniformly distrib-

546 MYCOLOGIA
FIGS. 1–11. Appendiculella lozanellae (holotype). 1–4. Hyphae with appresoria and conidiogenous cells. 5–7. Ascogeneous
hyphae and young asci. 8. Ascus with young ascospores. 9. Ascus with two almost mature ascospores. 10. Two mature
ascospores. One cell of the ascospore on the right side germinated. 11. Longitudinal section of an ascoma with appendages
and young asci. Scale bars: 1, 3, 4, 11 5 20 mm; 2, 5–10 5 10 mm.
uted over the entire colony, black, 86–257 3 86–
243 mm, surface cells conoid, with larviform appendages.
Larviform appendages more than 15 on one
perithecium, light brown, transversely striate, straight
to slightly bent, 44–81 3 11–20 mm. Asci ovoid with
two ascospores each. Ascospores cylindrical, rounded
at the tips, 44–52 3 16–18 mm, smooth, 4-septate,
slightly constricted at the septa, dark brown.
Etymology. In reference to Lozanella, a genus of the
Cannabaceae.
Holotype. PANAMA. PROV. CHIRIQUÍ: NationalPark
Volcán Barú, Los Quetzales trail, on Lozanella
enantiophylla (Donn. Sm.) Killip & Morton (Cannabaceae),
7.XI.2004, M. Piepenbring, R. Cáceres, R.
Mangelsdorff, I. Piepenbring & T. Trampe 3432
(holotype in PMA (18S rDNA, 632bp, GenBank

RODRÍGUEZ AND PIEPENBRING: NEW APPENDICULELLA SPECIES 547
DQ508301; 28S rDNA, 613 bp, GenBank DQ508302),
isotype in M).
Additional collections: PANAMA. PROV. CHIRIQUÍ: National
Park Volcán Barú, Los Quetzales trail, 2210 m,
21.VIII.2003, D. Rodríguez J. 38 (BPI, PMA; 28S rDNA
FIGS. 1–11. Continued.
sequence data of the host plant, 633 bp, GenBank
DQ508303); International Park La Amistad (PILA), El
Retoño Trail, 2270–2310 m, 3.III.2003, M. Piepenbring, R.
Kirschner et al 3194 (BPI, PMA); International Park La
Amistad (PILA), 2300 m, 2.III.2004, B. Koch 63 (M, PMA);

548 MYCOLOGIA
FIG. 12. Hyphae with appresoria and conidiogenous
cells of Appendiculella chiriquiensis. From the holotype.
Scale bars 5 10 mm.
International Park La Amistad (PILA), near Sendero La
Cascada, 2450 m, 22.IX.2005, R. Mangelsdorff 377 (BPI,
PMA).
Commentary. We are unaware of any species of
Meliolaceae described on members of the Cannabaceae
in the traditional sense (Hansford 1961, Farr et al
2006, references cited in the introduction). Because
many species of Cannabaceae (including Celtidaceae)
in the modern concept (Sytsma et al 2002) have been
classified in Ulmaceae, species of Meliolaceae known
on members of Ulmaceae have to be considered.
Seven species of Meliolaceae are known on species of
Trema, Celtis and Chaetachme (Hansford 1961). Six of
these are species of Meliola, characterized by the
presence of setae on their hyphae and thereby
markedly different from the species described above.
The seventh species is Asteridiella tremae (Speg.)
Hansf. on Trema guineense (Schum. & Thonn.)
Ficalho (Uganda, Sierra Leone, Ghana, Congo), T.
micrantha (L.) Blume (Argentina), T. orientalis (L.)
Blume (China, Java, Taiwan) and Trema sp. (Brazil,
Venezuela, Panama) (Hansford 1961, Farr et al 2006).
Asteridiella tremae differs fundamentally from the new
species by perithecia without larviform appendages.
These appendages might break off, but their presence
is still evident by large scars (cf. FIG. 17). In addition A.
tremae differs from the new species by smaller hyphal
cells (20–40 3 5–7 mm), smaller appresoria (20–
29 mm), smaller conidiogenous cells (13–20 3 6–
8 mm) and smaller ascospores (38–46 3 16–18 mm)
(Hansford 1961).
Efforts to isolate DNA from species of Meliolaceae
have been successful only for Appendiculella lozanellae
(specimen Piepenbring et al 3432). The partial 18S
rRNA gene sequence (632 bp, GenBank DQ508301)
is the third sequence of this gene available for species
of Meliolaceae in GenBank. It shows maximum
consensus of 97% with Meliola juddiana (AF021793)
and 93% with Meliola niessleana (AF021794, both by
Saenz and Taylor 1999). The partial 28S rRNA gene
sequence (613 bp, GenBank DQ508302) is the first
sequence of this gene for species of Meliolaceae in
GenBank.
Appendiculella chiriquiensis D. Rodríguez Justavino
& M. Piepenbr., sp. nov. FIGS. 12–15, 17
Plagulae amphigenae, laxae usque ad leviter densa, 1–
5 mm diam, confluentes. Hyphae atrofuscae, septatae,
leviter rectae usque ad leviter undulatae, ramificationibus
oppositis, cellulae 26–31 3 6–8 mm. Appresoria alternata,
recta usque ad antrorsa, 22–28 3 11–16 mm; cellula basalis
cylindrica, 6–10 3 6–8 mm; cellula apicalis, cylindrica usque
ad leviter lobata, 17–20 3 11–16 mm. Cellulae conidiogenae
commixtae appressoriis, alternatae, unilaterales, ampullatae,
18–27 3 7–9 mm. Perithecia distributa in tota plagula,

RODRÍGUEZ AND PIEPENBRING: NEW APPENDICULELLA SPECIES 549
FIGS. 13–15. Appendiculella chiriquiensis. 13. Appendages of ascomata. 14. Longitudinal, off center, section of an ascoma
with one appendage and young asci. 15. Mature ascospores. All from the holotype specimen. Scale bars: 13, 15 5 10 mm; 14 5
50 mm.

550 MYCOLOGIA
FIGS. 16–17. Ascomata of Appendiculella species. 16. A. lozanellae on Lozanella enantiophylla. 17. Appendiculella
chiriquiensis on Cupania guatemalensis Both taken from the holotype, SEM.
nigra, 119–210 3 116–206 mm, 6–8 appendicibus larviformis
pallide fuscis, transversaliter striatis, rectis usque ad leviter
curvatis, 73–128 3 18–22 mm. Ascosporae atrofuscae,
cylindricae, apicibus rotundatis, laeves, 4-septatae, leviter
constrictae in septis, 43–51 3 16–22 mm.
Colonies on both sides of leaves, scattered to
subdense, 1–5 mm diam. Adjacent colonies fusing to
black, thin layers. Hyphae dark brown, septate,
substraight to slightly undulate, branching opposite,
cells 26–31 3 6–8 mm. Appresoria alternate, straight

RODRÍGUEZ AND PIEPENBRING: NEW APPENDICULELLA SPECIES 551
to antrorse, 22–28 3 11–16 mm; stalk cell cylindrical,
6–10 3 6–8 mm; head cell cylindrical to slightly
lobulate, 17–20 3 11–16 mm. Conidiogenous cells
ampulliform, 18–27 3 7–9 mm, alternate, unilateral
mixed with appresoria. Perithecia uniformly distributed
over the entire colony, black, 119–210 3 116–
206 mm, surface cells conoid with larviform appendages.
Larviform appendages 6–8 mm on one perithecium,
light brown, transversely striate, straight to
slightly bent at the apex, 73–128 3 18–22 mm.
Ascospores cylindrical, 43–51 3 16–22 mm, rounded
at the tips, smooth, 4-septate, slightly constricted at
the septa, dark brown.
Etymology. Referring to Chiriquí Province, where
the fungus was collected.
Holotype. PANAMA. PROV. CHIRIQUÍ: Los Algarrobos,
close to the Río Majagua, on Cupania guatemalensis
(Turcz.) Radlk. (Sapindaceae), 23.XI.2004,
M. Piepenbring, I. Piepenbring & T. Trampe 3466
(holotype in PMA, isotype in M).
Commentary. Seventy-eight species and subspecies
of Meliolaceae are known on species of the Sapindaceae.
Sixty-seven species and subspecies belong to
Meliola, conspicuously differing from the species
described above by the presence of setae on the
hyphae. Four species of Irenopsis are characterized by
perithecia with setae (Hansford 1961). As drawn by
Hansford (1963) the setae of all four species differ
from larviform appendages by thick walls and the
presence of septa.
Seven species of Asteridiella are known from
Sapindaceae and could be confused with species of
Appendiculella if the larviform appendages of the
latter break off (see above). Three of these species
of Asteridiella are known on species of Cupania.
Asteridiella cupaniae (Toro) Hansf. on Cupania
americana L. (Puerto Rico) differs from the new
species by the cylindrical to clavate (not lobed) head
cells of appresoria and smaller ascospores (35–39 mm
long). Asteridiella guatemalensis Hansf. on Cupania
glabra Sw. ( Jamaica), C. guatemalensis (Turcz.) Radlk.
(Costa Rica), C. emarginata Cambess. (Brazil) and C.
americana L. (Venezuela, Trinidad) differ from the
new species by smaller hyphal cells (12–20 mm long),
mostly opposite appresoria with head cells cylindric
with broadly rounded apex (not lobed), as well as
smaller ascospores (34–40 mm long).
Asteridiella tersa (Cif.) Hansf. on Cupania americana
L. (Dominican Republic) differs from the new
species by opposite appresoria with head cells
cylindrical with rounded to subacuminate apex and
smaller ascospores (37–46 mm long) with the middle
cell often the largest. Asteridiella bonplandii (Speg.)
Hansf. on Sapindus saponaria L. is known from
Panama (Hansford 1961) and differs from the new
species by partly opposite appresoria and conspicuously
smaller ascospores (28 3 18 mm long). Asteridiella
chardoniana Hansf. on Serjania sp. (Venezuela)
differs by smaller ascospores (36–42 mm long).
Asteridiella dodonaeae Hansf. on Dodonaea triquetra
Wendl. (New South Wales) differs by smaller hyphal
cells (15–20 mm long) and often opposite appresoria
with subglobose to piriform head cells. Asteridiella
hypelates Hansf. on Hypelate trifoliata Sw. (Cuba)
differs by smaller colonies (1 mm diam) and smaller
hyphal cells (10–18 mm).
Although it is possible to justify new species as
presented above, only an investigation including
numerous specimens of Meliolaceae on closely related
hosts as well as molecular data will show whether
host relationships and small morphological differences
are sufficiently stable and indicative to distinguish
species. We are in great need of more
collections from tropical areas, detailed morphological
analyses as well as molecular data for Meliolaceae.
ACKNOWLEDGMENTS
We thank the German Academic Exchange Service (DAAD)
for a doctoral fellowship (DRJ) and continuous support for
our activities in Panama. The help of A. Monro (Department
of Botany, the Natural History Museum, London)
for the identification of Lozanella enantiophylla is gratefully
acknowledged. We thank R. Kirschner for correction of this
manuscript, M. Ruppel for his help at the scanning electron
microscope as well as R. Mangelsdorff for continuous help
during the investigation. D. Triebel’s collaboration for the
molecular study is acknowledged. R. Rincón (Autonomous
University of Chiriquí, Panama) helped in the field and with
literature. We also thank M. Vega, I. González and L. Vega
for help in the field as well as our team at the University of
Frankfurt, S. Späthe, J. Gossmann, T. Trampe, T. Hofmann,
D. Hahner and B. Koch.
LITERATURE CITED
Biju CK, Hosagoudar VB, Abraham TK. 2005. Meliolaceae
of Kerala, India XV. Nov Hedwig 80:465–502.
Cáceres O, Kirschner R, Piepenbring M, Schöfer H, Gené J.
2006. Hormographiella verticillata and an Ozonium stage
as anamorphs of Coprinellus domesticus. Antonie van
Leeuwenhoek 89:79–90.
Chacón S, Cruz F. 1999. Descripción de 13 nuevos registros
de mildiús negros (Meliolales) del estado de Veracruz,
México. Rev Mex Micol 15:23–26.
Correa AMD, Galdames C, de Stapf MS. 2004. Catálogo de
las plantas vasculares de Panamá. Primera Edición.
Bogotá, Colombia: Quebecor World. 599 p.
Farr DF, Rossman AY, Palm ME, McCray EB (n.d.). Fungal
databases, systematic botany & mycology laboratory,
ARS, USDA. Retrieved 17 Jan 2006, from http://nt.
ars-grin.gov/fungaldatabases/

552 MYCOLOGIA
Hansford CG. 1961. The Meliolineae. A monograph. Beih.
Sydowia 2:1–806.
———. 1963. Iconographia Meliolinearum. Beih. Sydowia
2:1–285.
Höhnel F. 1919. Fragmente zur Mykologie. Acad Wiss Wien,
Math Naturw Kl 138:556.
Hosagoudar VB. 1996. Meliolales of India. Calcutta:
Botanical survey of India. 363 p.
———. 2002. Meliolaceae of Kerala, India XIII. Sydowia 54:
53–58.
———, Abraham TK, Pushpangadan P. 1997. The Meliolineae—a
supplement. Kerala, India: Tropical Botanic
Garden and Research Institute. 201 p.
———, Biju CK, Abraham TK. 2001. Meliolaceae of Kerala,
India IX. J Econ Tax Bot 25:553–559.
———, Kaveriappa KM, Raghu PA, Goos RD. 1994.
Meliolaceae of southern India XVI. Mycotaxon 51:
107–118.
Katumoto K, Hosagoudar VB. 1989. Supplement to Hansford’s
‘‘The Meliolineae Monograph’’. J Econ Tax Bot
13:615–635.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Dictionary of
fungi. 9th ed. Wallingford, UK: CAB International. 655 p.
Luttrell ES. 1989. Morphology of Meliola floridensis.
Mycologia 81:192–204.
Mueller WC, Goos RD, Quainoo J, Morgham AT. 1991. The
structure of the phialides (mucronate hyphopodia) of
the Meliolaceae. Can J Bot 69:803–807.
Rodríguez HM. 2001. Acerca de la relación taxonomíaespecificidad
en Meliolales (Ascomycota). Rev Jard Bot
Nac Cuba 22:101–108.
———, Minter DW. 1998. Meliola mangiferae. IMI descriptions
of fungi and bacteria 1355.
———, ———. 1999. Meliola crucifera. IMI descriptions of
fungi and bacteria 1384.
Rodríguez JD. 2006. Meliolaceae aus Panama [Doctoral
dissertation]. Frankfurt am Main, Germany: J.W.
Goethe-Universität Frankfurt am Main. 268 p.
Saenz GS, Taylor JW. 1999. Phylogenetic relationships of
Meliola and Meliolina inferred from nuclear small
subunit rRNA sequences. Mycol Res 103:1049–1056.
Schmiedeknecht M. 1989. Meliolales aus Cuba. Wiss Zeitschr
Friedrich-Schiller-Univ Jena, Naturwiss R 38:185–209.
Song B. 1998. Studies on Appendiculella in China III.
Mycosystema 17:214–217.
———, Li TH. 2003. Six taxa of Meliolaceae from Yunnan
province in HKAS, China. Mycotaxon 87:421–424.
———, ———. 2004. Interesting taxa of Meliolaceae in
HMAS, China. Mycotaxon 90:129–132.
———, ———, Shen YH. 2003. Two new Meliola species
from China. Fungal Divers 12:173–177.
———, Ouyang Y, Hu Y. 1996. Studies on Appendiculella in
China. Acta Mycol Sinica 15:93–97.
———, ———, ———. 1997. Studies on Appendiculella in
China II. Mycosystema 16:244–246.
Stevens FL. 1927. The Meliolineae I. Ann Mycol 25:405–
477.
———. 1928. The Meliolineae II. Ann Mycol 26:165–388.
Sytsma KJ, Morawetz J, Pires JC, Nepokroeff M, Conti E,
Zjhra M, Hall JC, Chase MW. 2002. Urticalean rosids:
circumscription, rosid ancestry, and phylogenetics
based on rbcL, trnL-F, and ndhF sequences. Am J Bot
89:531–1546.
Yang JC. 1989. Preliminary studies on Meliolineae from
Guandgdong province of China. Acta Mycol Sinica 8:1–8.