Philippine Journal of Science
146 (2): 145-158, June 2017
ISSN 0031 - 7683
Date Received: ?? Feb 20??
Philippine Species of Parmotrema
(Ascomycota, Parmeliaceae)
Paulina A. Bawingan1*, Mechell P. Lardizaval1, Praxedes F. Rosuman2,
Weenalei T. Fajardo3, Andrea Azuelo4, John A. Elix5, and Jae-Seoun Hur6
1
Department of Biology, Saint Louis University, Baguio City, Philippines
2
St. Paul’s College, Vigan, Ilocos Sur, Philippines
3
Pangasinan State University, Lingayen, Pangasinan, Philippines
4
Central Mindanao University, Bukidnon, Philippines
5
Research School of Chemistry, Australian National University, Canberra, ACT 2601, Australia
6
Korean Lichen Research Institute, Suncheon National University, Suncheon, South Korea
This paper presents a taxonomic treatment of Parmotrema lichens (Ascomycota,
Parmeliaceae) collected in the Philippines at high altitudes where they abound,
particularly in the mountainous regions of northern Luzon and Mindanao. A
total of 30 Parmotrema species were identified: twelve being new records. Results
suggest that the Philippines has a potentially rich unknown lichen diversity
that remains to be further investigated. Many lichen species are restricted
to unexplored forested areas, which in the Philippines are facing alarming
degradation; hence, it is extremely important for these habitats to be conserved.
Key words: chemotaxonomy, foliose lichens, morphotaxonomy, parmotremoid lichens
INTRODUCTION
Parmotrema A. Massal. (1860) is a large genus of lichenized
fungi in the family Parmeliaceae. The name Parmotrema
refers to the perforate apothecia (Greek parmos = cup,
and trema = perforation) (Feige 1998). Parmotrema was
first segregated from Parmelia s.l. by Massalongo (1860)
based on P. perforata as type. Hale (1974a) resurrected the
genus and included in it all taxa incorporated in Parmelia
subgenus Amphigymnia (Vain.) C. W. Dodge, including
the Parmelia reticulata group previously included in
Parmelia Ach. subgenus Parmelia. Parmotrema was
subsequently subdivided by the separation of Rimelia
(Hale & Fletcher 1990) and Rimeliella (Kurokawa 1991)
or Canomaculina (Elix & Hale 1987; Elix 1997). A generic
*Corresponding author: pbawingan@gmail.com
reclassification of parmelioid lichens based on phylogenetic
studies, however, has recombined Rimelia, Canomaculina
(Rimeliella) as well as Concamerella into Parmotrema
(Blanco et al. 2005). Another phylogenetic analysis based
on morphological, molecular and chemical data showed
that the genera Flavoparmelia, Punctelia, Canoparmelia,
Flavopunctelia, and Nesolechia are seemingly also nested
within the Parmotrema clade (Crespo et al. 2010). More
studies, however, need to be undertaken to resolve generic
classification and synonymy among the parmelioid lichens.
Parmotrema sensu lato species are characterized by their
broad, rotund lobe apices, the absence of pseudocyphellae,
presence or absence of maculae, isidia or soredia, the
frequent occurrence of marginal cilia, the simple or
dimorphous rhizines, the presence of a broad erhizinate
or papillate marginal zone on the lower cortex (except
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Vol. 146 No. 2, June 2017
in the Rimelia and Rimeliella species group), and the
thick-walled, ellipsoid ascospores (Brodo et al. 2001;
Nash & Elix 2002). Cortical chemistry includes atranorin,
chloroatranorin, usnic acid and lichexanthone (Louwhoff
and Elix 1999). The medullary chemistry is highly
variable and many chemosyndromes have been observed
(Hale 1965; Krog & Swinscow 1981; Benatti et al. 2013).
There are currently more than 300 species of Parmotrema
known worldwide (Nash & Elix 2002). The center of
diversity is in the Neotropics, with numerous tropical
and subtropical species (Blanco et al. 2005; Crespo et
al. 2010). For the Philippines, Vainio (1909) described
several species now accommodated in Parmotrema. Hale
(1965, 1974b) also described five species of Parmotrema
from the Philippines. Recent studies of Philippine lichens
by foreign scientists assisted by Filipino biologists in the
1980s resulted in the discovery of several new species
and a number of new records. Aptroot and Sipman (1989)
reported Parmotrema clavuliferum (Räsänen) Streimann
and P. mellissii (C. W. Dodge) Hale both collected from
Mt. Santo Tomas, Benguet. Elix and Schumm (2001)
described P. negrosorientalum Elix & Schumm from
Negros Oriental and six new records for the Philippines;
P. cooperi (J. Steiner & Zahlbr.) Sérus, P. dilatatum (Vain.)
Hale, P. lobulascens (J. Steiner) Hale, P. permutatum
(Stirt.) Hale, P. rampoddense (Nyl) Hale, P. sancti-angelii
(Lynge) Hale, P. austrocetratum Elix & J. Johnst (syn.
Rimelia austrocetrata (Elix & Johnst.) Hale & Fletcher).
We started our lichen studies in 2000 and most of our
collections were made in high altitude areas of the
Cordillera Region where these broad-lobed foliose
lichens are abundant. Our initial collections resulted in
the description of Rimelia pustulata Elix & Bawingan
(= Parmotrema neopustulatum Kurok.) with P.
ultralucens (Krog) Hale, and P. vartakii Hale reported
as new records (Elix et al. 2002).
From our own collections made over the last decade as well
as other specimens reported from previous expeditions,
we have prepared this comprehensive taxonomic listing
of Parmotrema species in the Philippines (including
the species previously included in Rimelia). Diagnostic
characteristics as well as a key for the identification of
these found species of Parmotrema are also presented.
Few biologists study this group of lichenized fungi in the
Philippines or even in the entire Southeast Asia; hence,
this paper can be the first taxonomic report of these lichens
in the Region. Younger Filipino researchers are showing
interest on the study of lichens in the country; we hope
this paper can serve as guide in their taxonomic studies.
146
Bawingan et al.: Philippine Parmotrema Lichens
MATERIALS AND METHOD
Lichen Collection
After securing the gratuity permit from the Department
of Environment and Natural Resources (DENR) in 2000,
we explored various forests in the provinces of Benguet,
Mountain, and Ifugao. Collaboration with other universities
enabled us to collect in the provinces of Nueva Vizcaya,
Ilocos Norte, Pangasinan, Quirino, and Isabela. Dr. Andrea
Azuelo of Central Mindanao University provided duplicates
of collections from Mt. Apo and Mt. Kitanglad. All
specimens examined are deposited in the Father Braeckman
Museum of Natural History, Saint Louis University,
Baguio City. Dr. John Elix, one of the authors, and Dr.
Harrie Sipman of Freie University, Berlin, a lichenologistconsultant, confirmed most of the lichen identification.
Identification of the Lichens
Morphological and reproductive characters were observed
using Motic and Meiji stereomicroscopes. Sections of the
thallus and ascocarps were examined under the Motic
trinocular or Olympus compound microscopes. The
chemistry of the lichens was determined using spot tests.
Spot test involves the use of reagents such as potassium
hydroxide (K test), para-phenylenediamine (P test) and
calcium hypochlorite (C test). A drop of these solutions
was placed on the cortex and the medulla. The presence
or absence of color changes may indicate the presence of
certain lichen acids in the specimen. These determinations
were verified by thin layer chromatography using
solvents A and C following procedures described by
Elix and Ernst-Russell (1993) and Orange et al. (2001).
High-performance liquid chromatography (HPLC) was
also undertaken for some species. Two solvent systems
were used: (A) containing 1% aqueous orthophosphoric
acid and methanol in the ratio 3:7; and, (B) methanol.
The run started with 100% A and was raised to 58% B
within 15 min, then to 100% B within a further 15 min,
followed by isocratic elution in 100% B for a further
10 min. A photodiode array detector was used with
spectrophotometric reading at 254 nm with a flow rate of
1ml/min. The lichen acids were identified based on their
retention time (RT).
Based on the observed morpho-anatomic features and
chemistry, the specimens were identified guided by
available taxonomic keys and verified through lichen
descriptions from various references (Hale 1965; Krog &
Swinscow 1981, 1983; Elix & Johnston 1988; Elix 1994;
Louwhoff & Elix 1999; Spielmann & Marcelli 2009;
Sipman et al. 2013; Cyberliber not dated; Botanischer
Garten und Botanisches Museum Berlin not dated).
Lichen IDs and synonyms were obtained from Mycobank
(not dated).
Bawingan et al.: Philippine Parmotrema Lichens
Philippine Journal of Science
Vol. 146 No. 2, June 2017
RESULTS AND DISCUSSION
A total of 30 species of Parmotrema were identified
for the Philippines; 12 of these are new records for the
country (marked with *). Among the different collection
sites, Benguet Province in Luzon has the most number of
Parmotrema species followed by Mt. Province (Figure 1).
Few species were collected in the provinces of Cotabato
and Bukidnon in Mindanao; however, some species
are exclusively reported from the two provinces: P.
dilatatum in Bukidnon; P. cooperi, P. corniculans, and P.
permutatum in Cotabato; P. lobulascens in Bukidnon and
Cotabato. Other species collected only from one province
in the North include P. elacinulatum and P. saccatilobum
in Benguet, and P. vartakii and P. ultralucens in Mountain
Province. P. negrosorientalum was previously reported
only in Negros Oriental in the Visayas. This suggests the
need for further exploration of these and other provinces
to look for similar and other rare species.
Most of the lichens were collected in forested areas in the
different provinces; however, forest degradation in the
Philippines is described to be alarming (Asian Development
Bank 2009). The major cause is deforestation due to slashand-burn farming, illegal logging, mining, forest fires,
pest infestations, and typhoons. In March 2016, one of
the collection sites, Mt. Apo in Cotabato, Mindanao was
affected with massive forest fire due to hikers’ negligence
(Magbanua 2016). Another collection site, Mt. Santo Tomas
in Benguet, a favorite biodiversity laboratory in the past by
local and foreign scientists has now become a site of illegal
developmental projects which caused massive tree cutting
and man-made erosion (Aning 2014). Many similar activities
are happening throughout the country and these endanger the
natural habitats of lichens and other organisms.
Our data also shows that altitudinal ranges of the reported
Parmotrema species vary (Table 1). Four species only
were collected from areas with elevation below 1000
Figure 1. Percentage distribution of Parmotrema species according to province including Baguio City.
Table 1. Distribution of the different species according to elevation.
Elevation
Below 1000 m
1000 m to 2000 m
Above 2000 m
Parmotrema species collected in different altitudes
P. overeemi
P. hypotropum
P. saccatilobum
P. negrosorientalum
P. corniculans
P. elacinulatum
P. dilatatum
P. permutatum
P. ultralucens
P. vartakii
P. parahypotropum
P. praesorediosum
P. tinctorum
P. austrocetratum
P. cetratum
P. lobulascens
P. poolii
P. neopustulatum
P. rampoddense
P. subarnoldii
P. subrugatum
P. crinitum
P. cristiferum
P. gardneri
P. grayanum
P. maclayanum
P. mellissii
P. sancti-angelii
P. clavuliferum
P. reticulatum
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Bawingan et al.: Philippine Parmotrema Lichens
m while six species only from 1000 to 2000 meters in
elevation. Species with wider altitudinal range include
those collected from areas below 1000 to 2000 meters in
elevation (three species); eight species from areas with
elevation of 1000 to more than 2000 meters. Nine species
showing widest altitudinal range were collected from
all altitudinal zones. Altitudinal distribution shows that
Parmotrema can be found from different altitudinal zones
but most of the species are collected in higher altitude
areas, i.e. from 1000 m and above.
16 ° 52’ 48”N, 120 ° 50’ 52”E, 2186 m, P. Bawingan
SLU-L00122, SLU-L00217; Camp John Hay, Baguio
City, 16° 23’ 49.304”N, 120° 36’ 40.895”E, 1469 m,
R. Ansagay SLU-L00506; Sagada, Mt. Province 17°
5’ 52.87”N, 120° 54’ 26.294’’E, 1592 m, P. Bawingan
SLU-L00519, SLU-L00548; Camp John Hay, Baguio
City, 16° 23’ 49.304”N, 120° 36’ 40.895”E, 1469 m, S.
Tamayo SLU-L00611; Ampasit, Puguis, La Trinidad,
Benguet. 16° 26’ 54.137”N, 120° 34’ 32.836”E, 1324 m,
C. Feliciano SLU-L00794, SLU-L00802.
THE SPECIES
Parmotrema austrocetratum Elix & J. Johnst.,
Mycotaxon 31 (2): 495 (1988) [MB#133588] = Rimelia
austrocetrata (Elix & J. Johnst.) Hale & A. Fletcher (1990)
[MB#102560]; Parmelia austrocetrata Elix & J. Johnst.
[MB#373811]
Diagnosis: Thallus loosely adnate, upper surface
conspicuously reticulately cracked but not flaking off;
there is no or there is very narrow erhizinate marginal in
the lower cortex; black lower cortex with brown marginal
area; rhizines black, simple to squarrose; no soredia or
isidia; K+ yellow (atranorin), medulla K+ yellow to red,
C-, P+ red-orange (major acid, salazinic).
Specimens examined: Loakan Rd., Baguio City 16°
23’ 8.217”N, 120° 37’ 0.459”E, 1398 m, P. Bawingan
SLU-L00028; Mt. Santo Tomas, Benguet 16° 20' 4.9"N,
120° 33' 40.8"E, 2200 m, P. Bawingan SLU-L00203;
Sagada, Mt. Province 17° 5’ 52.87”N, 120° 54’ 26.294’’E,
1592 m, P. Bawingan SLU-L00312; Mt. Ugo, Benguet
16° 19’ 13.47”N, 120° 48’ 9.04”E, 2115m, J. Bugtong
SLU-L00583; Camp John Hay, Baguio City 16° 23’
49.304”N, 120 ° 36’ 40.895”E, 1469 m, G. Racca
SLU-L00884; Sinto, Bauko, Mt. Province 16° 51’ 42.0”N,
120° 52’ 25.9”E, 2118 m, P. Bawingan SLU-L01189;
Caliking, Atok, Benguet, 16° 35'N, 120° 42'E, 1450 m,
M. Lardizaval SLU-L01512, SLU-L01634; Monamon
Sur, Bauko, Mt. Province 16° 49’ 36.1”N, 120° 52’ 43.3”E,
2360 m, P. Bawingan & M. Lardizaval SLU-L02508; Mt.
Apo, Cotabato, 1866 m, A. Azuelo 71 (CMU, Bukidnon).
Other specimens: Cotabato Province: between Lake
Venado and Mt. Apo, Cotabato 6° 69’- 7° 00’N, 125°
16’- 125° 20’E, 2200-2800 m F. Schumm & U. Schwartz
CANB 6126; Mt. Apo, 7° 00’N, 125° 16’E U. Schwartz
5441 (Elix & Schumm 2001).
Diagnosis: Thallus loosely adnate, upper surface
conspicuously reticulately cracked, eventually becoming
areolate and flaking off exposing the medulla (Fig. 2A);
there is no or there is very narrow erhizinate marginal in
the lower cortex; black lower cortex with brown marginal
area; rhizines black, simple to squarrose; no soredia or
isidia; K+ yellow (atranorin), medulla K+ yellow to red,
C-, P+ red-orange (major acid, salazinic).
*Parmotrema cetratum (Ach.) Hale, Phytologia 28 (4):
335 (1974) [MB#343018]
= Parmelia cetrata Ach. (1814) [MB#373825]; Rimelia
cetrata (Ach.) Hale & Fletcher (1990) [MB#102561]
Specimens examined: Mt. Data, Bauko, Mt. Province,
148
Remarks: morphologically similar to P. austrocetratum
but maculae not forming areolae or flaking off.
Parmotrema clavuliferum (Räsänen) Streimann,
Bibliotheca Lichenologica 22: 93 (1986) [MB#129346]
= Parmelia clavulifera Räsänen (1944) [MB#368556];
Rimelia clavulifera (Räsänen) Kurok. (1991) [MB#128246]
Specimen examined: Mt. Cabuyao, Tuba, Benguet, 16°
21’ 33.363”N, 120° 33”E 52.841”E 2025 m, P. Bawingan,
P. Rosuman & Y. Flores SLU-L00042, SLU-L00200;
Virac, Itogon, Benguet, 16° 21’ 55.387”N, 120 ° 39’
11.031”E, 992 m, P. Bawingan & Y. Flores SLU-L00125,
A. Cuevas & V. Tabor SLU-L00322; Bana-ao, Tadian,
Mt. Province, 16° 55.769'N, 120° 49.639'E, 1340 m, P.
Bawingan SLU-L000271; Ampasit, Puguis, La Trinidad,
Benguet, 16° 26’ 54.137”N, 120° 34’ 32.836”E, 1324 m,
C. Feleciano, M. Lardizaval, & A. Rebogio SLU-L00798,
SLU-L00810; Sitio Sayet, Atok, Benguet, 16° 35'N, 120°
42'E, 1450 m, M. Lardizaval SLU-L01234, SLU-L01235,
SLU-L01237, SLU-L01240, SLU-L01375, SLU-L01377,
SLU-L011379, SLU-L01534, SLU-L01653, SLU-L01659;
Yamashita Shrine, Kiangan, Ifugao, 16.779299 oN,
121.081443oE, 700-800 m, A. Kiaki SLU-L02801; Sitio
Pula, Asipulo, Ifugao, 16° 41.4’ 28”N, 120° 4.9’ 23”E, 798
m, C. Oy-yeng & M. Balabag SLU-L02286.
Other specimen: Mt. Santo Tomas, Benguet, 16° 18’N,
120° 35’E, 2100 m, A. Aptroot & H. Sipman S 21812c
(Aptroot & Sipman 1989).
Diagnosis: Thallus loosely adnate, membranaceous
to coriaceous, laciniate, capitate soralia commonly
developing on tips of laciniae, upper surface with
reticulate maculae, sometimes becoming cracks but no
flaking off from the cortex (Fig. 2C); lower cortex black
with white to light brown or mottled marginal zone;
rhizines simple to squarrose; medulla K+ yellow to red,
Philippine Journal of Science
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Bawingan et al.: Philippine Parmotrema Lichens
Figure 2. Representatives of Parmotrema species and their features: (A) P. austrocetratum showing flaking areolae; (B) P. maclayanum
thallus and erhizinate marginal zone of the lower cortex; (C) P. clavuliferum showing capitate marginal soralia on laciniae;
(D) conspicuous cilia in P. subarnoldii; (E) lobulated laciniae and pycnidia in P. subrugatum; (F) thallus of P. crinitum; (G)
globose isidia of P. ultralucens; (H) isidiate lamina of P. tinctorum thallus; thallus of (I) P. overemii, (J) P. gardneri,
(K) P. vartakii, and (L) P. elacinulatum; (M) pigmented lower medulla of P. rampoddense; (N) K+ yellow to red medulla
of P. cristiferum; (O) P+ orange medulla of P. cristiferum; (P) KC+ brown medulla of P. rampoddense.
C-, P+ orange-red; major acid, salazinic.
Remarks: Morphologically similar to P. reticulatum
but distinctly laciniate with capitate soralia, and white to
mottled marginal zone in the lower cortex.
Parmotrema cooperi (J. Steiner & Zahlbr.) Sérus, The
Bryologist 87 (1): 4 (1984) [MB#107091] = Parmelia
cooperi J. Steiner & Zahlbr. (1926) [MB#397385]
Specimen examined (Elix & Schumm 2001): Cotabato
Province: near Ilomavis, 7° 02’N, 125° 11’E, 730 m, F.
Schumm & U. Schwartz CANB Schumm 5885.
Remarks: Elix (1994) describes this lichen to have
loosely adnate thallus, sinuate and ascending margins,
ciliate, soralia marginal to submarginal. Lower cortex
black with narrow to wide brown erhizinate zone; rhizines
simple. Medulla K-, C+ intense red, KC+ red, P-; major
acid, lecanoric.
*Parmotrema corniculans (Nyl.) Hale, Phytologia 28
(4): 335 (1974) [MB#343027]
= Parmelia corniculans Nyl. (1885) [MB#119691]
Specimen examined: Mt. Apo, Cotabato, 6° 56.652'N,
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125° 13.815'E, 1820 m, A. Azuelo193 (CMU, Bukidnon).
Diagnosis: Thallus coriaceous to membranaceous,
margins ciliate, lacking isidia and soredia; white medulla;
black lower cortex with narrow to wide erhizinate zone;
rhizines black, simple, rarely branched, some growing
upwards penetrating the thallus; medulla K-, C-, P-, lower
medulla K+ purple; major acid, alectoronic.
*Parmotrema crinitum (Ach.) M. Choisy, Bulletin
Mensuel de la Société Linnéenne de Lyon 21: 175
(1952) [MB#368891] = Parmelia crinita Ach. (1814)
[MB#119691]; Imbricaria crinita (Ach.) Arnold (1984)
[MB#387011]; Parmotrema crinita (Ach.) M. Choisy
(1952) [MB3273930]
Specimens examined: Burnham Park, Baguio City, 16°
24’ 34.33”N, 120° 35’ 41.345”E, 1430 m, B. Pangsiw, Y.
Flores, & P. Rosuman SLU-L00025; Mt. Cabuyao, Tuba,
Benguet 16° 21’ 23”N, 120° 33’ 32.8”E, 2025 m, Y. Flores &
P. Rosuman SLU-L00099; Mt. Data, Bauko, Mt. Province,
16° 52’ 48”N, 120° 50’ 52”E, 2186 m, P. Bawingan, Y. Flores,
& P. Rosuman SLU-L00220; Sagada, Mt. Province, 17° 5’
52.87”N, 120° 54’ 26.294’’E, 1592 m, P. Bawingan &M.
Lardizaval SLU-L00428; Camp John Hay, Baguio City,
16° 23’ 49.304”N, 120° 36’ 40.895”E, 1469 m, R. Ansagay
SLU-L00491, D. Fianga-an SLU-L00496, L. Cervantes
SLU-L00497; Mt. Data, Mt. Province, 16° 52’ 48”N, 120°
50’ 52”E, 2186 m, Y. Flores & P. Bawingan SLU-L00664;
Sitio Cayet, Caliking, Atok Benguet, 16° 35'N, 120° 42'E,
1450 m, M. Lardizaval SLU-L01636; Asipulo, Ifugao,
16° 41’ 9.08”N, 121° 4.1’ 23”E, 837 m, C. Oy-yeng, F.
Bengwayan & M. Balabag SLU-L02791.
Bawingan et al.: Philippine Parmotrema Lichens
57’ 57.886”N, 120° 48’ 55.568”E, 1387 m, P. Bawingan
& Y. Flores SLU-L00270; Ambangeg, Kabayan, Benguet.
16° 34’ 34.464”N, 120° 53’ 12.374”E, 2503 m, P. Bawingan
& Y. Flores SLU-L00366; Bontoc, Mountain Province,
17° 05' 14"N, 120° 58' 32"E, 1087 m, P. Bawingan & P.
Rosuman SLU-L00415; Bana-ao, Tadian, Mt. Province,
16 ° 55’ 41.997”N, 120 ° 49’ 20.169” E, 1340 m, P.
Bawingan SLU-L01591, SLU-L01593; Asipulo, Ifugao,
16o41’9.08”N, 121o4.1’23”E, 837 m, C. Oy-yeng, F.
Bengwayan & M. Balabag SLU-L02795, SLU-L02804,
SLU-L02808; Mankayan, Benguet 16° 52’N, 120° 47’E,
1320 m, C. Oy-yeng SLU-L02810; Adams, Ilocos Norte,
18° 27’ 21.6”N, 120° 54’ 22.4”E, 305 m, P. Bawingan &
M. Balabag SLU-L01027.
Diagnosis: Thallus loosely adnate to adnate, lobes
broadly rounded, main lobes eciliate, lateral lobes may be
sparingly ciliate, cilia rarely longer than 2.5 cm; margins
ascending, soralia linear, predominantly marginal; white
medulla; lower surface black with wide brown erhizinate
zone, rhizines sparse, black, simple; medulla K+ yellow
then dark red, C-, P+ orange to orange-red; (major acid,
salazinic; minor, consalazinic) (Fig. 2N, 2O).
Parmotrema dilatatum (Vain.) Hale, Phytologia 28 (4):
335 (1974) [MB#343038]
= Parmelia dilatata Vain. (1890) [MB#122344]
Specimen examined: Mt. Kitanglad, Bukidnon, 8° 10’N,
124° 56’E, 1870-2800 m, F. Schumm & U. Schwartz
CANB Schumm 6303 (Elix & Schumm 2001).
Diagnosis: Thallus adnate, margins crenate to isidiate;
isidia dense often ciliate at the apices, laminal and
marginal, becoming coralloid, occasionally dissolving
into soredia (Fig. 2F); margins sparsely ciliate; white
medulla; lower cortex black with wide brown erhizinate
zone, rhizines black, simple; medulla K+ yellow, C-, P+
orange (major acid, stictic acid; minor, constictic acid).
Diagnosis: Louwhoff and Elix (1999) described the thallus
to be membranaceous to coriaceous, margins entire to
crenate, sometimes sublaciniate, eciliate; soralia marginal
to submarginal, linear to subcapitate, soredia granular;
medulla white or pale yellow; lower surface black, rhizines
sparse, simple, black; medulla K+ dull yellow becoming
yellow-brown; C-, P+ orange-red; protocetraric and
echinocarpic acids.
Parmotrema cristiferum (Taylor) Hale, Phytologia 28
(4): 335 (1974) [MB#343031]
*Parmotrema elacinulatum (Kurok.) Streimann,
Bibliotheca Lichenologica 22:94 (1986)
= Parmelia cristifera Taylor, London Journal of Botany
6: 165 (1847) [MB#120260]
[MB#426776] = Parmotrema submerrillii Elix, Mycotaxon
47: 120 (1993) [MB#360126]
Specimen examined: Mt. Cabuyao, Benguet, 16° 21’ 23”N,
120° 33’ 32.8”E, 2025 m, Bawingan SLU-L00027; Banaao, Tadian, Mt. Province 16° 55.769'N, 120° 49.639'E
1117 m, P. Bawingan & C. Bawingan SLU-L00092,
SLU-L000103; Sagada, Mt. Province, 17° 5’ 52.87”N,
120° 54’ 26.294’’E, 1592 m, P. Bawingan & P. Rosuman
SLU-L00130, SLU-L00143; Bontoc, Mt. Province 17°
05' 14"N, 120° 58' 32"E, 1087 m, P. Bawingan & E.
Singson SLU-L00185; Tadian Central, Mt. Province, 16°
Specimen examined: Sitio Sayet, Caliking, Atok,
Benguet, 16⁰ 35'N, 120⁰ 42'E, 1450 m, M. Lardizaval
SLU-L01251.
150
Diagnosis: Thallus loosely adnate, coriaceous, lobes
subirregular to sublinear; lacking isidia, soredia, and
laciniae; margins with dense, conspicuous cilia; medulla
white; lower cortex black with wide erhizinate zone,
rhizines black, simple; medulla K-, C-, KC-, P+ orangered; major acid, protocetraric.
Philippine Journal of Science
Vol. 146 No. 2, June 2017
Remarks: no apothecia observed in the specimen
examined.
*Parmotrema gardneri (C.W. Dodge) Sérus, The
Bryologist 87 (1): 5 (1984) [MB#107092]
Specimens examined: Mt. Cabuyao, Tuba, Benguet, 16°
21’ 33.363”N, 120° 33”E 52.841”E, 2025 m, P. Bawingan,
P. Rosuman & Y. Flores SLU-L00018; Virac, Itogon,
Benguet, 16° 21’ 55.387”N, 120° 39’ 11.031”E, 992 m, P.
Rosuman & Y. Flores SLU-L00414; Sagada, Mt. Province,
17° 5’ 52.87”N, 120° 54’ 26.294”E 1592 m, M. Coloma
SLU-L00631; Asipulo, Ifugao 16° 41’ 9.96”N, 121° 4.1’
79”E, 796 m, F. Bengwayan & M. Balabag SLU-L02816.
Diagnosis: Thallus loosely adnate to adnate, coriaceous
to sometimes membranaceous, lobes subirregular to
sublinear, often sublaciniate, margin eciliate to very
sparingly ciliate; soralia marginal to submarginal, linear
to subcapitate; medulla white (Fig. 2J); lower surface
rugose, black with brown erhizinate zone; rhizines sparse,
black, simple, short (0.1-0.2 mm in length); medulla K+
pale yellow to pale brown, C-, KC+ red or brown, P+
orange-red (major acid, protocetraric).
*Parmotrema grayanum (Hue) Hale, Phytologia 28 (4):
336 (1974) [MB#343059]
= Parmelia grayana Hue (1899) [MB#397645]
Specimens examined: Mt. Cabuyao, Tuba, Benguet, 16°
21’ 33.363”N, 120° 33’ 52.841”E, 2003 m, P. Bawingan,
P. Rosuman & Y. Flores SLU-L00019; Loakan Road,
Baguio City, 16° 23’ 8.217”N, 120° 37’ 0.459”E, 1386 m, R.
Julian & E. Puguon SLU-L00024; Tadian, Mt. Province,
Philippines. 16° 57’ 57.886”N, 120° 48’ 55.568”E, 1340 m,
P. Bawingan &Y. Flores SLU-L00090; Mt. Sto. Tomas,
Tuba, Benguet. 16° 20’ 6” N, 120° 33’ 38.99” E, 2206 m,
Y. Flores, P. Rosuman, & P. Bawingan SLU-00263; Mt.
Data, Bauko, Mt. Province, 16° 51' 22.0"N, 120° 51'
14.0"E, 2294 m, Y. Flores & P. Bawingan SLU-L00264,
P. Bawingan Y. Flores, M. Lardizaval SLU-L00660; PMA
Compound, Baguio City, 16° 22’ 0”N, 120° 37’ 0”E, 1362
m, P. Rosuman & Y. Flores SLU-L00371, K. Dumlao,
R. Amoncio, & W. Leyson SLU-L01071; Virac, Itogon,
Benguet, 16° 21’ 55.387”N, 120° 39’ 11.031”E, 992 m,
M. Ramos & F. Rabena SLU-L00374; Mt. Ugo, Benguet,
16° 19’ 13.47”N, 120° 48’ 9.04”E, 2112 m, J. Bugtong,
J. Maslang, & M. Racoma SLU-L00582; Sagada, Mt.
Province, 17° 5’ 52.87”N, 120° 54’ 26.294’E, 1592 m, G.
Benedito & S. Cabral SLU-L00638, G. Benedito & P.
Bawingan SLU-L00638, SLU-L00640; Acop, Tublay,
Benguet. 16° 28’ 51.449”N, 120° 37’ 53.457”E, 1336 m,
P. Bawingan & M. Lardizaval SLU-00906; Kumyas, La
Trinidad, Benguet 16° 30' 28.0"N, 120° 38' 10.0"E, 1207
m, F. Bengwayan SLU-L02809.
Bawingan et al.: Philippine Parmotrema Lichens
Diagnosis: Thallus adnate to loosely adnate, coriaceous;
lobes rotund and crowded, margins ascending, crenate
with conspicuous cilia; upper surface commonly whitepruinose, emaculate; soralia marginal to submarginal,
linear to subcapitate; medulla white; lower surface rugose,
black, with brown erhizinate zone; rhizines sparse, black,
simple, short (1-2 mm in length); medulla K-, C-, KC-,
P-; with fatty acids in the medulla.
Remark: Most of the specimens we collected were
corticolous.
Parmotrema lobulascens (J. Steiner) Hale, Phytologia
28 (4): 337 (1974) [MB#343073]
= Parmelia lobulascens J. Steiner (1903) [MB#397816]
Specimen examined (Elix & Schumm 2001): Mt.
Kitanglad, Bukidnon 8° 10’N, 124° 56’E, 870-2800 m, F.
Schumm & U. Schwartz, CANB Schumm 6272; Mt. Apo,
Cotabato, 7° 00’N, 125° 16’E, 2200 m, F. Schumm & U.
Schwartz CANB Schumm 6071, 6084.
Diagnosis: Thallus coriaceous, loosely attached, with
ciliate margins, secondary lobules common along lobe
margin and on the lamina; upper surface maculate,
soralia linear, marginal to submarginal; some parts of
medulla ochraceous, margin of the lower surface brown
to mottled; medulla UV+ white (alectoronic acid) (Krog
& Swinscow 1981).
Parmotrema maclayanum (Müll. Arg) Hale, Phytologia
28 (4): 337 (1974) [MB#343078]
= Parmelia maclayana Müll. Arg. (1891) [MB#397833];
Parmotrema maclayanum (Müll. Arg) Phytologia 28 (4):
337 (1974) [MB#493068]
Specimens examined: Mt. Data, Mt. Province, 16 °
52’ 48”N, 120° 50’ 52”E, 2186 m, P. Bawingan & M.
Lardizaval SLU-L00415; Virac, Itogon, Benguet, 16° 21’
55.387”N, 120° 39’ 11.031”E, 992 m, A. Cuevas & V. Tabor
SLU-L00327; Sagada, Mt. Province, 17° 5’ 52.87” N,
120o 54’ 26.294’’ E 1592 m, M. Lardizaval & A. Rebogio
SLU-L00566; Sitio Sayet, Atok, Benguet, 16⁰ 35'N, 120⁰
42'E, 1450 m, S. Laguardia & K. Velasco SLU-L02786,
J. Domingo & E. Meniado SLU-L02787, J. Tambalong
& S. Laguardia SLU-L02788.
Diagnosis: Thallus moderately to loosely adnate, lobes
round, weakly imbricate; margins suberect, sparsely
ciliate, isidia and soredia absent; medulla white; lower
surface black with dark brown erhizinate zone, rhizines
black, simple, sparse (Fig. 2B); medulla K-, KC+ purple,
P+ red; major acid, alectoronic.
Parmotrema mellissii (C.W. Dodge) Hale, Phytologia 28
(4): 337 (1974) [MB#343083]
151
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Vol. 146 No. 2, June 2017
= Parmelia mellissii C. W. Dodge (1959) [MB#368691]
Specimens examined: Sagada, Mt. Province, 17o 5’
52.87”N, 120o 54’ 26.294’’E, 1592 m, Y. Flores, M.
Lardizaval & P. Rosuman SLU-L00147, P. Bawingan
SLU-L00639; Virac, Itogon, Benguet, 16o 21’ 55.387”N,
120o 39’ 11.031”E 992 m, P. Bawingan & P. Rosuman
SLU-L00326, V. Tabor & P. Rosuman, SLU-L00416;
Mt. Data, Mt. Province, 16o ° 52’ 48”N, 120° 50’ 52”E,
2186 m, Y. Flores & P. Bawingan SLU-L00418; Tinoc,
Ifugao, 16° 42’ 12.595” N, 120° 58’ 29.456”E, 1492 m, H.
Pugong & M. Taclobao SLU-L00425; Mt. Ugo, Benguet,
16° 19’ 13.47”N, 120° 48’ 9.04”E, 2112 m, J. Bugtong, J.
Maslang, & E. Pagayonan SLU-L00584; Burnham Park,
Baguio City, 16° 24’ 34.33”N, 120° 35’ 41.345”E, 1430
m, L. Magtoto SLU-L00592, SLU-L00942; Ampasit, La
Trinidad, Benguet, 16° 27' 0"N, 120° 34' 0"E, 1300 m,
C. Feleciano, M. Lardizaval, & A. Rebogio SLU-L00797;
Camp John Hay, Baguio City, 16° 23’ 49.304” N, 120° 36’
40.895”E, 1469 m, S. Tamayo & L. Magtoto SLU-L00886;
Yamashita Shrine, Kiangan, Ifugao, 16.779299 oN,
121.081443oE, 700-800 m, A. Kiaki SLU-L01253; Sitio
Sayet, Atok, Benguet, 16° 35'N, 120° 42'E 1450 m, M.
Lardizaval SLU-L01517, SLU-L01518, SLU-L01516,
SLU-L01519, SLU-L01639; Aparngao, Ifugao, 16° 56'
52.3"N, 121° 03' 20.5"E, 1463 m, A. Kiaki SLU-L02422;
Mt. Apo, A. Azuelo, CMU 33.
Other specimens: Mt. Santo Tomas, Benguet, 16° 18’N,
120° 35’E, 2100 m, A. Aptroot & H. Sipman A 20456, S
21808, S 21822, S 21823 (Aptroot & Sipman 1989).
Diagnosis: Thallus membranaceous, loosely adnate,
lobes rounded; margins ascending, ciliated, cilia long and
abundant; isidia on upper surface laminal to marginal,
simple or branching to coralloid, often ciliated; medulla
white with yellow-brown to orange brown pigmented
areas; lower surface black with wide brown erhizinate
zone; rhizines long, simple, black clustered in groups,
scattered; medulla K-, C-, KC+ red, P-; major acids,
alectoronic and α-collatolic.
Parmotrema negrosorientalum Elix & Schumm
Mycotaxon 79: 253-260 (2001) [MB#474503] =
Parmotrema negrosorientalis Elix & Schumm (2001)
[MB#476100]; Parmotrema negrosorientale Elix &
Schumm, Mycotaxon 79: 253 (2001) [MB#546186]
Specimen examined: Mt. Talinis, Negros Oriental,
9° 15’N, 123° 10”E, F. Schumm & U. Schwartz, herb.
Schumm 7521, 7463, 7465, 7466, 7571 (Elix & Schumm
2001).
Remarks: Described as a new species by Elix and
Schumm (2001). Holotype: herb. Schumm 7521. P.
negrosorientalum was reported to be morphologically
similar with P. rampoddense but differing in having a
152
Bawingan et al.: Philippine Parmotrema Lichens
large, coriaceous thallus, maculate upper surface, perforate
apothecial discs, and much larger spores; it closely
resembles P. lobulascens but the latter does not contain
skyrin (Elix & Schumm 2001).
Parmotrema overeemii (Zahlbr.) Elix, Australasian
Lichenology 42: 22-27 (1998) [MB#458345] = Parmelia
overeemii Zahlbr. (1928) [MB#397997]
Specimens examined: Sablan, Baguio City, 16 ° 29’
18.674”N, 120° 30’ 59.021”E, 853 m, J. Bagsawan
SLU-L00234; Adams, Ilocos Norte, 18° 27’ 18.74”N,
120° 54’ 46.1”E, 414 m, P. Bawingan & M. Balabag
SLU-L01030, 18° 27’ 12.7”N, 120° 54’ 17.9”E, 321 m, P.
Bawingan & M. Balabag SLU-L01028.
Diagnosis: Thallus moderately to loosely adnate,
membranaceous, partly blackened, lobes subirregular
to sublinear, 4-10 mm wide; margins entire to crenateincised, ciliated, occasionally lobulated (Fig. 2I); isidia
and soredia absent; lower surface black with wide brown
erhizinate zone, rhizines dense, black, simple; medulla K-,
C-, KC+ red, P+ deep orange; major acid, protocetraric.
*Parmotrema parahypotropum (W.L. Culb.) Hale,
Phytologia 28(4): 338 (1974) [MB#343101] = Parmelia
parahypotropa W.L. Culb. (1973) [MB#342869]
Specimen examined: Sitio Sayet, Atok, Benguet, 16°
35'N, 120° 42'E, 1450 m, M. Lardizaval SLU-L01521;
Sitio Pula, Asipulo, Ifugao 16° 41.4’ 28”N, 121° 4.9’ 23”E,
798 m, M. Balabag SLU-L02326; Asipulo, Ifugao, 16° 41’
9.08”N, 121° 4.1’ 23”E, 837 m, C. Oy-yeng, F. Bengwayan
& M. Balabag SLU-L02790.
Diagnosis: Thallus moderately to loosely adnate,
coriaceous; lobes sublinear, distinctly laciniate, labriform
soralia developing on tips of laciniae, soredia farinose;
lower surface black with white to mottled erhizinate
marginal zone especially on laciniae; rhizines sparse,
black, simple; medulla K+ yellow to red, C-, P+ yellow
then orange; major acid, salazinic.
Parmotrema permutatum (Stirt.) Hale, Phytologia 28 (4):
338 (1974) [MB#343104]
= Parmelia permutata Stirt. (1878) [MB#398037]
Specimen examined (Elix & Schumm 2001): Mt. Apo,
Cotabato, 7° 01’N, 125° 13’E, 1240 m, F. Schumm & U.
Schwartz, herb. Schumm 5941.
Diagnosis: Thallus loosely adnate, membranaceous to
coriaceous, lobes rounded, margins entire to crenate,
ciliated; with marginal to submarginal linear soralia; upper
medulla white, lower medulla yellow, orange-yellow, or
salmon pink; lower surface medulla K-, C+ pink, KC+
pale red, P-; with gyrophoric acid.
Philippine Journal of Science
Vol. 146 No. 2, June 2017
*Parmotrema poolii (C.W. Dodge) Krog. & Swinscow,
The Lichenologist 15 (2): 130 (1983) [MB#109156] =
Parmelia poolii C.W. Dodge [MB#373942]
Specimen examined: Barlig, Mt. Province, 17 ° 02’
23.97”N, 121° 08’ 36.7”E, 1860 m, P. Bawingan, Y.
Flores, & M. Lardizaval SLU-L00209; Camp John
Hay, Baguio City, 16° 23’ 49.304” N, 120° 36’ 40.895”E,
1469 m, R. Ansagay SLU-L00498; Mt. Data, Bauko,
Mt. Province, 16° 52’ 48”N, 120° 50’ 52”E, 2186 m,
P. Bawingan & Y. Flores SLU-L00663; Sitio Sayet,
Atok, Benguet, 16° 35'N, 120° 42'E, 1450 m, M.
Lardizaval SLU-L01227, SLU-L01365, SLU-L01367,
SLU-L011523, SLU-L01640, SLU-L01645.
Diagnosis: Thallus loosely adnate, coriaceous, lobes
rotund, wide, margin crenate-dentate, ciliated; with
marginal linear to subcapitate soralia; pycnidia scattered,
conidia sublageniform; lower surface black with broad
brown to dark brown erhizinate marginal zone; rhizines
black, simple; medulla K-, C-, KC+ red, P-; with
alectoronic acid.
Remarks: P. poolii resembles P. rampoddense except in its
sublageniform conidia and absence of skyrin in the medulla.
*Parmotrema praesorediosum (Nyl.) Hale, Phytologia
28 (4): 338 (1974) [MB#343106]
= Parmelia praesorediosa Nyl. (1891) [MB#398093]
Specimens examined: Lubon, Tadian, Mt. Province,
17° 1’ 40”N, 120° 4.7’ 49”E, 1200-1300 m, M. Taclobao
SLU-L00041; Bana-ao, Tadian, Mt. Province, 16°
55.769'N, 120° 49.639'E, 1340 m, P.Bawingan
SLU-L00087; Virac, Itogon, Benguet, 16° 21’ 55.387”N,
120° 39’ 11.031”E, 992 m, P. Bawingan & P. Rosuman
SLU-L00105; Sagada, Mt. Province, 17° 05' 03.6"N,
120° 54' 06.9"E, 1462 m, P. Bawingan, P. Rosuman, &
E. Singson SLU-L00231; Asipulo, Ifugao, 16° 41’ 9.08”N,
121° 4.1’ 23”E, 837 m, C. Oy-yeng, F. Bengwayan, &
M. Balabag SLU-L02793; Adams, Ilocos Norte, 18 °
27’ 18.7”N, 120° 54’ 46.1”E, 414 m, P. Bawingan & M.
Balabag SLU-L01031.
Diagnosis: Thallus adnate, coriaceous, lobes rounded,
margins entire or crenate, lacking cilia; with marginal
or submarginal soralia, linear to crescent-shaped; lower
cortex black with white or mottled or brown erhizinate
marginal zone; rhizines simple, black; medulla K-,
C-, KC-, P-; major acids, protopraesorediosic and
praesorediosic; with fatty acids.
Remarks: P. praesorediosum morphologically resembles
P. grayanum except for its eciliate margins.
Parmotrema neopustulatum Kurok., Journal of Japanese
Botany 81: 252 (2006) [MB#542306] = Rimelia pustulata
Bawingan et al.: Philippine Parmotrema Lichens
Elix & Bawingan (2002) [MB#484787]; Parmotrema
pustulatum (Elix & Bawingan) O. Blanco, A. Crespo,
Divakar, Elix & Lumbsch (2005) [MB#335080]
illegitimate name
Specimen examined: Mt. Data National Park, Mt.
Province1, 16° 52’N, 120° 52’E, 2305 m, P. Bawingan &
Y. Flores SLU-L00125; Bana-ao, Tadian, Mt. Province,
16° 55’N, 120° 49’E, 1516 m P. Bawingan SLU-L0073;
Camp John Hay, Baguio City, 16° 23’ 49.304” N, 120°
36’ 40.895”E 1469 m, S. Tamayo SLU-L00882; Sitio
Sayet, Atok, Benguet, 16° 35'N, 120° 42'E, 1450 m, M.
Lardizaval SLU-L01233, SLU-L01650, SLU-L01652.
Diagnosis: Thallus adnate, coriaceous, maculae reticulate
becoming areolate and flaking off; soredia pustulate,
pustules submarginal to laminal, soredia becoming
corticate to form pseudoisidia; medulla K+ yellow to red,
C-, KC+ red, P+ orange; major acid, salazinic.
Parmotrema rampoddense (Nyl) Hale, Phytologia 28 (4):
338 (1974) [MB#343114]
= Parmelia rampoddensis Nyl. (1900) [MB#398146];
Diploschistes rampoddensis (Nyl.) Zahlbr. (1924)
[MB#384471]
Specimens examined: Mt. Data, Tadian, Mountain
Province, Philippines. 16° 52’ 48”N, 120° 50’ 52”E, 2186
m, P. Bawingan SLU-L00216, V. Tabor, Y. Flores, M.
Lardizaval SLU-L00417, M. Lardizaval & M. Lardizaval
& P. Bawingan SLU-L00662; Mines View, Baguio City,
16.4197°N, 120.6273°E, 1400-1450 m, V. De Villa
SLU-L00368; Sagada, Mt. Province, 17° 5’ 52.87”N,
120° 54’ 26.294’’E, 1592 m, P. Rosuman & E. Singson
SLU-L00421, SLU-L00422, 17° 05' 03.6"N, 120° 54'
06.9"E, 1462 m, P. Bawingan SLU-L00650; Camp John
Hay, Baguio City, 16° 23’ 49.304”N, 120° 36’ 40.895”E,
1469 m, R. Ansagay & D. Fianga-an SLU-L00494, L.
Cervantes SLU-L00495, L. Magtoto, S. Tamayo, & G.
Racca SLU-L00765; Sitio Sayet, Atok, Benguet, 16°
35'N, 120° 42'E, 1450 m, M. Lardizaval SLU-L01228,
L-01229, L-1230, L-01231, L-1368, L-1369, L-01371,
L-01524; Aparnga-o, Ifugao, 16° 56' 52.4"N, 121° 03'
19.1"E, 1463 m, M. Lardizaval SLU-L02382; Kumyas,
La Trinidad, Benguet, 16° 30' 29"N, 120° 38' 6.0"E, 1238
m, F. Bengwayan SLU-L02805; Mt. Apo, Cotabato, A.
Azuelo 273 (CMU).
Diagnosis: Thallus loosely adnate, coriaceous, lobes
irregular, becoming involute, margins crenate with long,
conspicuous cilia; sorediate, soredia farinose, soralia
marginal, linear; medulla white, the area adjacent to the
lower cortex red-orange (with skyrin) (Fig. 2M); black
lower surface with wide brown erhizinate marginal zone,
rhizines black, simple; medulla K-, C-, KC+ red, P-; with
alectoronic acid and α-collatollic acid in the medulla.
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Philippine Journal of Science
Vol. 146 No. 2, June 2017
Remarks: P. rampoddense is morphologically similar
to P. poolii except that it has baciliform conidia and
pigmented lower medulla indicative of skyrin.
Parmotrema reticulatum (Taylor) M. Choisy, Bulletin
Mensuel de la Société Linéenne de Lyon 21: 175 (1952)
[MB#357464] = Parmelia reticulata Taylor (1836)
[MB#398165]; Parmelia laevigata var. reticulata (Taylor)
Linds. (1866) [MB#435088]; Rimelia reticulata (Taylor)
Hale & A. Fletcher (1990) [MB#102567]; Parmotrema
clavuliferum (Räsanen) Streimann (1986) [MB#129346]
Specimens examined: Mt. Data Hotel Compound,
16° 51’ 9”N, 120° 51’ 47”E, 2200 m, P. Bawingan & Y.
Flores SLU-L00233; Camp John Hay, Baguio City, 16°
23’ 49.304”N, 120° 36’ 40.895”E, 1469 m, D. Fianga-an
& R. Ansagay SLU-L00493; Sagada, Mt. Province. 17°
5’ 52.87”N, 120° 54’ 26.294’’E, 1592 m, P. Bawingan
SLU-L00635, SLU-L00653, SLU-L-00654; Ampasit,
Puguis, La Trinidad, Benguet, 16° 26’ 54.137”N, 120°
34’ 32.836”E, 1324 m, C. Feleciano, M. Lardizaval, &
A. Rebogio SLU-L00796; Mt. Ugo, Benguet, 16° 19’
13.47”N, 120° 48’ 9.04”E, 2112 m, Y. Flores & J. Maslang
SLU-L00838, J. Maslang & H. Wagas SLU-L00845;
Sinto, Bauko, 16° 51’ 41.8”N, 120° 52’ 25.5”E, 2128
m, M. Lardizaval & P. Bawingan SLU-L01191; Sitio
Sayet, Atok, Benguet, 16° 35'N, 120° 42'E, 1450 m, M.
Lardizaval SLU-L01236, SLU-L01238, SLU-L1244,
SLU-L01245, SLU-L01378, SLU-L01655, SLU-L01656;
Sitio Pula, Asipulo, Ifugao, 16° 41.4’ 28”N, 120° 4.9’
23”E, 798 m, M. Balabag & F. Bengwayan SLU-L02817;
Aparnga-o, Ifugao, 16o 59’ 58.4”N, 121o ° 03’ 19.5”E,
1447 m, R. Doplah SLU-L02381, SLU-L02389; Mt. Apo,
Cotabato, 1855 m, A. Azuelo 153 (CMU).
Diagnosis: Thallus loosely adnate, membranaceous to
coriaceous, lobes imbricate with subascending margins,
partly incised and laciniate; upper surface with reticulate
maculae, sometimes becoming cracks but no flaking
off from the cortex; sorediate, soralia marginal to
submarginal, labriform to linear to subcapitate, soredia
granular; lower surface black, rhizinate or papillate to the
margin or with narrow brown erihizinate zone; rhizines
simple or squarrose, long, dense, black; medulla K+
yellow to red, C-, P+ orange-red; major acid, salazinic.
*Parmotrema saccatilobum (Taylor) Hale, Phytologia
28(4): 339 (1974) [MB#343122]
= Parmelia saccatiloba Taylor (1847) [MB#398206]
Specimen examined: Virac, Itogon, Benguet, 16° 21’
55.387”N, 120° 39’ 11.031”E, 992 m, V. Valdez &G.
Sepulchre SLU-L00372.
Diagnosis: Thallus loosely adnate, coriaceous, lobes
rotund, broad; margins entire, lacking cilia; isidiate,
154
Bawingan et al.: Philippine Parmotrema Lichens
isidia moderately dense, cylindrical, simple or becoming
branched; lower surface black with wide erhizinate
marginal zone; rhizines simple, brown, sparse; medulla
K-, C-, KC+ red-brown, P+ brick-red; major acid,
protocetraric.
Parmotrema sancti-angelii (Lynge) Hale, Phytologia, 28:
339 (1974) [MB#343123]
= Parmelia sancti-angelii Lynge (1914) [MB#398215]
Specimens examined: Bontoc, Mt. Province, 17 ° 5’
28.522”N, 121° 0’ 38.063”E, 1087 m, P. Bawingan & P.
Rosuman SLU-L00197, P. Bawingan & M. Lardizaval
SLU-L00135; Bana-ao, Tadian, Mt. Province, 16° 55’
41.997”N, 120° 49’ 20.169”E, 1340 m, P. Bawingan
SLU-L00108, SLU-L00110; Sagada, Mt. Province, 17°
5’ 52.87”N, 120° 54’ 26.294”E, 1592 m, P. Bawingan &
P. Rosuman SLU-L00140, P. Bawingan SLU-L01179;
Barlig, Mt. Province, 17° 02’ 23.97”N, 121° 08’ 36.7”E,
1860 m, P. Bawingan, M. Decaleng, Y. Flores, & M.
Lardizaval SLU-L00314; Ambangeg, Kabayan, Benguet.
16° 34’ 34.464”N, 120° 53’ 12.374”E, 2503 m, P. Bawingan
& Y. Flores SLU-L00367; Virac, Itogon, Benguet, 16°
21’ 55.387”N, 120° 39’ 11.031”E, 992 m, V. Tabor & P.
Rosuman SLU-L00429; Sitio Sayet, Atok, Benguet, 16°
35'N, 120° 42'E, 1450 m, M. Lardizaval SLU-L01660;
Yamashita Shrine, Kiangan, Ifugao, 16.779299 oN,
121.081443 oE, 700-800 m, A. Kiaki SLU-L02810,
SLU-L02811, SLU-L02815.
Other specimens: Mt. Apo, Cotabato, 7° 01’N, 125° 13’E,
1240 m, F. Schumm & U. Schwartz herb. Schumm 5941
(Elix & Schumm 2001).
Diagnosis: Thallus loosely adnate to adnate,
membranaceous to coriaceous, lobes irregular, margins
crenate, ascending, ciliated; sorediate, soredia farinose,
soralia marginal, linear; lower surface black with brown or
mottled erhizinate marginal zone; rhizines dense, simple,
black; medulla KC+ pale red, KC+ pale red, P-; major
acid, gyrophoric.
*Parmotrema subarnoldii (Abbayes) Hale, Phytologia
28 (4): 339 (1974) [MB#343129]
= Parmelia subarnoldii Abbayes (1961) [MB#345522]
Specimens examined: Sagada, Mt. Province, 17 ° 5’
52.87”N, 120° 54’ 26.294’’E, 1592 m, P. Bawingan &
P. Rosuman SLU-L00142; Tadian, Mt. Province, 16° 52’
48”N, 120° 50’ 52”E, 2186 m, P. Bawingan SLU-L00265;
Mt. Data National Park, Mt. Province1, 16° 52’N, 120°
52’E, 2305 m, P. Bawingan & Y. Flores SLU-L00419;
Ampasit, Puguis, La Trinidad, Benguet, 16° 26’ 54.137”N,
120° 34’ 32.836”E, 1324 m, C. Feleciano, M. Lardizaval, &
A. Rebogio SLU-L00778, SLU-L00813; Sitio Sayet, Atok,
Benguet, 16° 35'N, 120° 42'E, 1450 m, M. Lardizaval
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Vol. 146 No. 2, June 2017
SLU-L01645; Aparnga-o, Ifugao, 16° 56' 52.4"N, 121°
03' 19.1"E, 1463 m, R. Doplah SLU-L02378;
Diagnosis: Thallus loosely adnate, membranaceous to
coriaceous, lobes rounded or irregularly incised; margins
entire or crenate, ciliated (Fig. 2D); sorediate, soredia
farinose, soralia linear, marginal; medulla white, partly
with yellow-orange pigment adjacent to lower cortex;
lower surface black with brown or mottled erhizinate
marginal zone; rhizines simple, rarely branched, sparse;
medulla K-, C-, KC+ red-brown, P+ orange-red; major
acid, protocetraric acid, with skyrin.
*Parmotrema subrugatum (Kremp.) Hale, Phytologia 28
(4): 339 (1974) [MB#343135]
= Parmelia subrugata Kremp. (1868) [MB#398366]
Specimens examined: Barlig, Mt. Province, 17 ° 02’
23.97”N, 121° 08’ 36.7”E, 1860 m, M. Dicaleng & M.
Lardizaval SLU-L00266; Mt. Ugo, Benguet, 16 ° 19’
13.47”N, 120° 48’ 9.04”E, 2112 m, J. Maslang & M.
Racoma SLU-L00586; Mt. Apo, Cotabato 1823 m, A.
Azuelo 170 (CMU, Bukidnon), 1820 m, A. Azuelo 242
(CMU, Bukidnon).
Diagnosis: Thallus loosely adnate, coriaceous, lobes
crowded, convoluted, lobulate laciniate at the margins,
with cilia (Fig. 2E); upper surface weakly maculate
lacking isidia or soredia; medulla white, red-orange near
the lower cortex; medulla K-, C-, KC+ red, P; major acid
alectoronic, with skyrin.
Parmotrema tinctorum (Despr. ex Nyl.) Hale, Phytologia
28 (4): 339 (1974) [MB#343140] = Parmelia tinctoria
Despr. Ex Nyl. (1872) [MB#273861]; Parmelia tinctorum
Despr. Ex Nyl. (1872) [MB#542088]; Parmotrema
tinctorium (Despr. Ex Nyl.) Hale, Phytologia 28 (4): 339
(1974) [MB#531735]
Specimens examined: Mt. Sto. Tomas, Tuba, Benguet,
16° 20’ 6”N, 120° 33’ 38.99”E, 1996 m, C. Fomaneg
& G. Negritto SLU-L00026; Bana-ao, Tadian, Mt.
Province, 16° 55.684'N, 120° 49.477'E, 1144 m, P.
Bawingan SLU-L00104; Bontoc, Mt. Province, 17° 05’
14”N, 120° 58’ 32”E, 1087 m, P. Bawingan, Y. Flores &
M. Lardizaval SLU-L00131; PMA Compound, Baguio
City, 16° 21’ 42”N, 120° 37’ 14”E, 1540 m, C. Fomaneg
& D. Lucas SLU-L00261; Virac, Itogon, Benguet, 16°
21’ 55.387”N, 120° 39’ 11.031”E, 992 m, V. Valdez
& G. Sepulchre SLU-L00321, A. Cuevas & V. Tabor
SLU-L00370; Tadian Poblacion 16° 45’N, 120° 5’E, 1340
m, P. Bawingan SLU-L00324; Mt. Lusod, Benguet, 16°
31’N, 120° 46’E, 1625 m, P. Bawingan & T. Colallad
SLU-L00570; Acop, Tublay, Benguet 16° 21’N, 120°
37’E, 1400 m P. Bawingan & M. Lardizaval SLU-L00907;
Burnham Park, Baguio City, 16° 24’ 35”N, 120° 35’
Bawingan et al.: Philippine Parmotrema Lichens
41”E, 1504.88 m S. Tamayo & G. Racca SLU-L00947;
Botanical Garden and Centennial Park, Baguio City, 16°
56.197'N, 121° 08.190'E, 1460 m, W. Leyson, R. Amoncio
& K. Dumlao, SLU-L01073; Sitio Sayet, Caliking, Atok,
Benguet, 16° 35'N, 120° 42'E, 1450 m, M. Lardizaval
SLU-L01252, SLU-L01253; Asipulo, Ifugao, 16° 41.4’
28”N, 120° 4.9’ 23”E, 798 m, C. Oy-yeng & M. Balabag
SLU-L02796, SLU-L02800; Yamashita Shrine, Kiangan,
Ifugao 16.779299oN, 121.081443oE, 700-800 m, A. Kiaki
SLU-L02797; Kumyas, La Trinidad, 16° 30’ 28”N, 120°
38’10”E, 1207 m, F. Bengwayan SLU-L02806.
Diagnosis: Thallus large, loosely adnate, lobes irregular,
margins linear or crenate, no cilia (Fig. 2H); isidia sparse
to abundant, mostly laminal, confluent to scattered, some
marginal; medulla white; lower surface black with wide
brown erhizinate marginal zone; rhizines short, simple,
black; medulla K-, C+ red, KC+ red, P-; major acid,
lecanoric.
Parmotrema ultralucens (Krog) Hale, Mycotaxon 1 (2):
108 (1974) [MB#342939]
= Parmelia ultralucens Krog (1974) [MB#342939];
Canomaculina ultralucens (Krog) Elix & J.B. (2003)
[MB#372302]
Specimens examined: Bontoc, Mt. Province, 17° 5’N,
120° 59’E, 1196 m, P. Bawingan, P. Rosuman, & E.
Singson SLU-L00131.
Diagnosis: Thallus loosely attached, lobes rounded to
subirregular, margins crenate, ciliated; isidia laminal,
simple to coralloid, often ciliated (Fig. 2G); medulla
white; lower surface black with brown, erhizinate zone,
rhizines, simple, black, varied in length; medulla UV+
yellow, K+ yellow to red, C-; major acids, salazinic and
lichexanthone.
Parmotrema vartakii Hale, Mycotaxon 5 (2): 441 (1977)
[MB#343144]
Specimen examined: Madongo, Sagada, Mt. Province,
17° 15’N, 120° 54’E, 1996 m, P. Bawingan, P. Rosuman
& E. Singson SLU-L00139.
Diagnosis: Thallus adnate, membranaceous to coriaceous,
lobes subirregular to linear, rugose, margins ciliated, cilia
sparse to numerous, slender; sorediate, soredia farinose,
scattered in the lamina (Fig. 2K); medulla white; lower
surface black with brown erhizinate zone; rhizines simple,
sparse, black; medulla K+ pale orange, C+ pale orange,
KC+ red orange, P-; major acid, gyrophoric.
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KEY TO THE PARMOTREMA SPECIES
1a Thallus without wide erhizinate marginal zone in the lower surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b Thallus with wide erhizinate marginal zone in the lower surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2a Soredia present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2b Soredia absent . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3a Soredia pustulate, marginal to submarginal, maculae forming areoles then flaking of . . . . . . . P. neopustulatum
3b Soredia not pustulate, marginal, labriform to linear, maculae not
forming areoles or flaking off . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. reticulatum
4a Maculae forming areoles, then flaking off . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. austrocetratum
4b Maculae not forming areoles or flaking off . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cetratum
5a Thallus lacking isidia and soredia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5b Thallus with isidia or soredia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6a Medulla entirely white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6b Medulla partly orange-pigmented near lower cortex, pigmented part K+ purple . . . . . . . . . . . . . . . . . . . . . . . . 9
7a Medulla KC-, P+ deep orange; protocetraric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7b Medulla KC+ purple, P+ red; alectoronic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. maclayanum
8a Cilia present, prominent; conidia bacilliform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. elacinulatum
8b Cilia present or absent; conidia sublageniform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. overeemii
9a Thallus lobulate-laciniate; conidia bacilliform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subrugatum
9b Thallus not lobulate-laciniate, conidia filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. corniculans
10a Thallus with isidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10b Thallus with soredia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
11a Eciliate or sparingly ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11b Lobes conspicuously ciliate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12a Medulla C+ red, P-; lecanoric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tinctorum
12b Medulla C-, P+ brick-red; protocetraric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. saccatilobum
13a Medulla K-, KC+ red; alectoronic and α-collatolic acids present . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mellissii
13b Medulla K+ yellow or red, KC- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14a Medulla K+ yellow; stictic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. crinitum
14b Medulla K+ yellow to red; salazinic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ultralucens
15a Lobes eciliate or sparsely ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15b Lobes conspicuously ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
16a Medulla K- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. praesorediosum
16b Medulla K+ yellow or yellow then red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17a Medulla K+ yellow then red; salazinic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cristiferum
17b Medulla K+ dull yellow to yellow-brown; protocetraric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a Upper surface grey; usnic acid and echinocarpic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. gardneri
18b Upper surface pale yellowish-grey; usnic acid and echinocarpic acids present . . . . . . . . . . . . . . . . P. dilatatum
19a Medulla K+ yellow then red, P+ orange; norstictic and salazinic acids present . . . . . . . . . . P. parahypotropum
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19b Medulla K- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20a Medulla C+ red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20b Medulla C- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
21a Lower medulla pigmented, medulla C+ pink; gyrophoric acid present . . . . . . . . . . . . . . . . . . . . P. permutatum
21b Medulla entirely white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22a Medulla C+ intense red; lecanoric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cooperi
22b Medulla C+ pink or pale red; gyrophoric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sancti-angelli
23a Medulla entirely white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23b Medulla with pigmented areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
24a Medulla KC-; fatty acids present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grayanum
24b Medulla KC+ red; alectoronic acid ± α-collatolic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25a Upper surface maculate; secondary lobules often present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lobulascens
25b Upper surface emaculate; secondary lobules absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. poolii
26a Medulla P+ orange; protocetraric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subarnoldii
26b Medulla P-; alectoronic acid ± α-collatolic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27a Thallus coriaceous; upper surface distinctly maculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. negrosorientalum
27b Thallus membranaceous; emaculate upper surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. rampoddense
ACKNOWLEDGMENT
The authors gratefully acknowledge the funding support
given by Vlaamse Interuniversitaire Raad (VLIR)
through the SLU-BSU-VLIR inter-university cooperation
(2000-2005), the Saint Louis University Research Grant
(SLU-URG) from 2010-2013, and the Sunchon National
University, Sunchon, Korea from 2014-2016.
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