PLANT SCIENCE TODAY, 2021
Vol 8(4): 889–899
https://doi.org/10.14719/pst.2021.8.4.1265
ISSN 2348-1900 (online)
HORIZON
e-Publishing Group
RESEARCH ARTICLE
Pyrenocarpous lichens in Goa with five new records to India
Pallavi Randive1, Gaurav K. Mishra2, Sanjeeva Nayaka2*, D. K. Upreti2 & M. K. Janarthanam1
1
Department of Botany, Goa University, Taleigao Plateau, Goa 403 206, India
Lichenology Laboratory, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow 226 001, India
*Email: nayaka.sanjeeva@gmail.com
2
ARTICLE HISTORY
ABSTRACT
Received: 21 May 2021
Accepted: 27 August 2021
Available online: 29 September 2021
The pyrenocarpous lichens are the one which produces perithecial ascocarps. They are one of the
prominent groups of lichens in tropical forests. Frequent incidence of pyrenocarpous taxa in lichen
biota of Goa prompted us to take up the exclusive study of this group in the State. The study revealed
the occurrence of 79 species belonging to 15 genera and seven families. The family Pyrenulaceae had
the maximum number of 23 species, while 20 belonged to the family Porinaceae. In comparison to
North Goa, South Goa is fairly well explored for lichens representing 71 species from 11 localities. The
following five species are reported as new to India — Porina exserta, P. siamensis, Pyrenula
dissimulans, P. pyrenastrospora and P. rinodinospora. With the addition of five new records, the Goa
State now represents 165 species of lichens, out of which 48% is represented by pyrenocarpous
lichens. The present study will be useful for monographic studies on pyrenocarpous lichens and for
environmental monitoring studies in the area, as this can be considered as a key indicator species.
KEYWORDS
Arthopyreniaceae
Biodiversity
Porinaceae
Pyrenulaceae
Taxonomy
Western ghats
Introduction
The Western Ghats traverses through the State of Goa,
India and is a well-known biodiversity hotspot.
Extensive lichen exploration in the State has been
initiated recently, and so far, 160 species are reported
(1-6). Frequent occurrence of pyrenocarpous lichens
during the exploration in the State indicated their
dominance which prompted us to carry out an
exclusive study on this group. Pyrenocarpous are a
group of lichens having perithecia as their ascocarps.
Pyrenocarpous taxa commonly grow on the bark of
trees, sometimes on rocks, soil or leaves, mostly in
moist tropical regions of the world (7).
In the past, pyrenocarpous lichens collected from
Goa were included in various monographic works (814) and few new species such as Anthracothecium
goaense A. Singh [current name Pyrenula gibberulosa
(Vain.) Aptroot] and P. subacutalis Upreti [current
name Pyrenula maravalensis Vain.] were also
described. The recent floristic studies in the State
reported 14 pyrenocarpous taxa from the Cotigao
Wildlife Sanctuary (15) and 13 taxa from Bondla and
Bhagwan Mahavir Wildlife Sanctuaries (16). A list of
lichens from the Goa State reveals 118 species under
42 genera and 23 families, of which 44 were
pyrenocarpous lichens (1). Further, in an exclusive
survey
of
Cotigao
Wildlife
Sanctuary
36
pyrenocarpous taxa were reported under nine genera
(3). In the studies on foliicolous lichens, there
enumerated a total of 15 pyrenocarpous lichens (2, 4).
These reports clearly indicate that Goa State has a rich
diversity of pyrenocapous lichens. Therefore, carrying
out an exclusive study on pyrenocarpous lichens of
the State was inevitable. The ultimate aim of the study
was to produce a checklist and identification key for
pyrenocarpous lichens of Goa State by compilation of
previous reports and by more exploration in the State.
Materials and Methods
The present study is based on published literature,
previously collected specimens deposited in the
herbarium LWG of CSIR-National Botanical Research
Institute, Lucknow and freshly collected specimens
from Goa. About 350 specimens from 19 localities of
Goa (Fig. 1) were available for the study.
Morphological and anatomical characters were
examined using stereo zoom Leica S8APO and light
DM2500 microscopes attached to a camera. Thin
sections of perithecia were cut using a razor blade
under a stereo zoom microscope. All anatomical
measurements were recorded in plain water, while
© Randive et al (2021). This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use,
distribution and reproduction in any medium, provided the original author and source are credited (https://creativecommons.org/licenses/by/4.0/).
890
RANDIVE ET AL
Fig. 1. Map of Goa showing North and South districts and localities surveyed for present study.
10% KOH was used for the detailed study of asci and
ascospores. For spot tests, the usual reagents of K, C
and P were used. To identify lichens substance, thin
layer chromatography (TLC) was performed in solvent
system C following the standard method (17). Fresh
specimens were preserved in the herbarium of Goa
University (GU), and a set of voucher specimens were
deposited in herbarium LWG. The specimens were
identified up to species level with the help of keys of
earlier published literature (18-25). The classification
of lichens summarized in reference 26 was followed
for arranging species under their respective families.
The identity of species was confirmed by matching
with type specimens or well-identified specimens
available at LWG.
Results
The study revealed the occurrence of 79 species of
pyrenocarpous lichens belonging to 15 genera and
seven families in Goa. Five species were recorded for
the first time from the country (Table 1). The brief
descriptions for the five newly recorded lichens and
key for all the pyrenocarpous lichens encountered in
Goa are provided. The Pyrenulaceae family was most
dominant in the State, with 23 species followed by
Porinaceae (20 spp.) and Monoblastiaceae (13 spp.).
Among the genera, Pyrenula was dominant with 24
species, followed by Porina (19 spp.), Anisomeridium
(12 spp.) and Strigula (8 spp.). All the species
recorded were crustose except for Endocarpon
subrosettum A. Singh & Upreti, which was
squamulose. A total of 58 species were found to grow
on tree trunks, branches and twigs (corticolous),
followed by 16 species that grow on leaves
(foliicolous), while five species grow on rocks
(saxicolous). Astrothelium meristosporum (Mont. &
Bosch) Aptroot & Lücking and A. scoria (Fée) Aptroot
& Lücking exhibited their substrate specificity with
restricted occurrence on cashew nut (Anacardium
occidentale Linn.) trees. The species of Porina and
Pyrenula exhibit maximum substrate diversity as
they were found growing on various trees, leaves and
rocks, while all Strigula species exhibit luxuriant
growth only on leaves. Artocarpus integrifolius L.f.,
Anacardium occidentale L., Cocos nucifera L. and
Terminalia elliptica Willd. were the common
phorophytes for the growth of pyrenocarpous
lichens. Porina interestes (Nyl.) Harm., P. tetracerae
(Afz.) Müll. Arg., Pyrenula aspistea (Ach.) Ach., P.
oculata A. Singh & Upreti and Trypethelium eluteriae
Spreng. were the most common species of the State.
PLANT SCIENCE TODAY
891
Table 1. List of pyrenocarpous lichens recorded from Goa and their distribution [Note: + = present, - = absent, CR = Crustose, SQ =
Squamulose, C = Corticolous, S = Saxicolous, F = Foliicolous, BMWS = Bhagwan Mahavir Wildlife Sanctuary, CWS = Cotigao Wildlife
Sanctuary, BWS = Bondla Wildlife Sanctuary, Goa Uni. = Goa University, *New record for India]
Goa Uni.
Melauli
Panjim
Ponda
Radigati
Valpoi
Agonda
BMWS
Caranzol
Colem
CWS
Mollem
Pirla
Quepem
Quinamol
Rivona
Tambdisurla
P. subinterstes (Nyl.) Müll. Arg.
BWS
36
37 P. tetracerae (Afz.) Müll. Arg.
38 Trichothelium alboatrum Vain.
Pyrenulaceae
39 Lithothelium decumbens (Müll. Arg.) Aptroot
40 Pyrenula adacta Fée
41 P. aggregata (Fée) Fée
42 P. approximans (Kremp.) Müll. Arg.
43 P. aspistea (Ach.) Ach.
44 P. breutelii (Müll. Arg.) Aptroot
45 P. brunnea Fée
46 P. castanea (Eschw.) Müll. Arg.
47 P. dissimulans (Müll. Arg.) R.C. Harris*
48 P. fetivica (Krempelh.) Müll. Arg.
49 P. gibberulosa (Vain.) Aptroot
50 P. immissa (Stirt.) Zahlbr.
51 P. leucostoma Ach.
52 P. leucotrypa (Nyl.) Upreti
53 P. mamillana (Ach.) Trevis.
54 P. maravalensis Vain.
55 P. nitidula (Bres.) R.C. Harris.
Amona
Arthopyreniaceae
1
Arthopyrenia alboatra (Kremp.) Müll. Arg.
2
A. finkii Zahlbr.
3
A. grisea (Schierch.) Körb.
4
A. indusiata Müll. Arg.
5
A. nidulans Müll. Arg.
Monoblastiaceae
6
Anisomeridium albidoatrum (Nyl.) R.C. Harris
7
A. angulosum (Müll. Arg.) R.C. Harris
8
A. biforme (Borrer) R.C. Harris
9
A. complanatum (Makhija & Patw.) R. C. Harris
10 A. consobrinum (Nyl.) Aptroot
11 A. glaucescens (Müll. Arg.) R.C. Harris
12 A. indicum (Makhija & Patw.) R. C. Harris
13 A. palavanum (Vain.) R.C. Harris
14 A. subnexum (Nyl.) R.C. Harris
15 A. tarmugliense (Makhija & Patw.) R.C. Harris
16 A. terminatum (Nyl.) R.C. Harris
17 A. ubianum (Vain.) R.C. Harris
18 Monoblastia pellucida Aptroot
Porinaceae
19 Clathroporina mastoidea (Ach.) R.C. Harris
20 Porina americana Fée
21 P. atlantica (Erich.) P. M. Jørg.
22 P. chrysophora (Stirt.) R. Sant.
23 P. conica R. Sant.
24 P. epiphylla Fée
25 P. exserta Müll. Arg.*
26 P. internigrans (Nyl.) Müll. Arg.
27 P. interestes (Nyl.) Harm.
28 P. karnatakensis Makhija, Adaw. & Patw.
29 P. nitidula Müll. Arg.
30 P. ochrostoma Makhija, Adaw. & Patw.
31 P. pallescens R. Sant.
32 P. rufula (Kremp.) Vain.
33 P. siamensis P.M. McCarthy*
34 P. subcutanea Ach.
35 P. subhibernica Upreti
Habitat
Sl.
Family and species
No.
South Goa
Growth form
North Goa
CR
CR
CR
CR
CR
C
C
C
C
C
-
-
-
-
-
-
-
-
-
+
-
-
-
+
+
+
+
-
-
-
-
-
+
+
-
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
C
C
C
C
C
C
C
C
C
C
C
C
C
-
-
+
+
+
+
-
-
-
+
+
-
+
+
+
-
+
-
+
-
-
-
+
+
+
+
+
+
-
+
-
-
-
+
-
+
-
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
C
C
C
F
F
F
C
C
C
F
F
C
F
F
S
C
C
-
+
+
-
-
+
-
-
-
+
+
-
-
-
+
+
+
+
+
-
-
+
+
+
+
+
+
+
+
+
+
+
+
+
+
-
-
-
-
+
+
+
+
-
CR S
-
-
+
-
-
-
-
-
-
-
-
-
+
-
-
-
-
-
-
CR S
CR F
-
+
-
-
-
-
-
-
-
-
+
-
+
-
-
+
+
-
-
-
-
+
-
-
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
-
+
+
-
+
-
-
+
-
-
+
-
+
+
-
+
-
+
+
+
+
+
-
+
+
-
+
-
+
+
+
+
+
+
+
+
-
-
+
-
-
-
+
+
-
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
-
892
RANDIVE ET AL
56 P. ochraceoflava (Nyl.) R. C. Harris
57 P. oculata A. Singh & Upreti
58 P. quassiaecola (Fée) Fée
59 P. pyrenastrospora Aptroot*
60 P. rinodinospora Aptroot *
61 P. subumbilicata (C. Knight) Aptroot
Strigulaceae
62 Strigula antillarum (Fée) Müll. Arg.
63 S. concreta (Fée) R. Sant.
64 S. janeirensis (Müll. Arg.) Lücking
65 S. nitidula Mont.
66 S. phyllogena (Müll. Arg.) R.C. Harris
67 S. smaragdula Fr.
68 S. subelegans Vain.
69* S. subtilissima (Fée) Müll. Arg.
Trypetheliaceae
70 Astrothelium luridum (Zahlbr.) Aptroot & Lücking
71 A. meristosporum (Mont. & Bosch) Aptroot & Lücking
72 A. scoria (Fée) Aptroot & Lücking
Marcelaria benguelensis (Müll. Arg.) Aptroot, Nelsen
73
& Parnmen
74 Nigrovothelium bullatum Lücking, Upreti & Lumbsch
75 N. tropicum (Ach.) Lücking, M.P. Nelsen & Aptroot
76 Trypethelium eluteriae Spreng.
77 T. plicatorimosum Mahija & Patw.
Verrucariaceae
78 Endocarpon subrosettum A. Singh & Upreti
79 Staurothele fissa (Taylor) Zwackh
CR
CR
CR
CR
CR
CR
C
C
C
C
C
C
+
-
+
-
+
-
-
+
-
-
+
+
-
+
-
+
+
+
-
-
+
-
+
-
-
-
+
-
-
-
CR
CR
CR
CR
CR
CR
CR
CR
F
F
F
F
F
F
F
F
-
-
-
-
-
-
-
-
-
+
-
-
-
+
+
+
+
+
+
+
+
-
-
-
-
-
-
CR C
CR C
CR C
-
-
-
-
-
-
-
-
-
-
-
-
+
+
-
-
+
-
-
-
-
CR C
-
-
-
-
+
-
-
-
-
-
-
-
+
-
-
-
-
-
-
CR
CR
CR
CR
C
C
C
C
+
-
-
-
-
-
-
-
+
-
-
-
-
+
-
+
+
+
+
-
-
+
+
-
-
-
-
SQ S
CR S
-
-
+
-
-
+
-
-
-
-
-
-
-
-
-
-
-
-
-
-
The studied specimens belonged to nineteen
localities; eleven were from North Goa, while eight
localities were from South Goa. The maximum
diversity of pyrenocarpous lichens is recorded in
South Goa with 71 species, while North Goa recorded
29 species and 21 species are common between the
two districts. It may be noted that South Goa has a
more dense forest area which is also fairly well
explored compared to North Goa. Among the different
localities, Cotigao Wildlife Sanctuary (CWS) is the most
extensively surveyed and has maximum diversity of
pyrenocarpous lichens with 48 species, while
Bhagwan Mahavir Wildlife Sanctuary (BMWS) has 17
and Valpoi has eight species. Ponda and Quinamol
localities are poorly represented by pyrenocarpous
lichens with one species each. Compared to other sites
Ponda and Quinamol are anthropogenically more
disturbed which may be the reason for low
pyrenocarpous lichen diversity here.
Distribution and ecology:—This species is found
growing on the bark of Terminalia paniculata Roth in
Cotigao Wildlife Sanctuary between altitude of 100–
200 m. Earlier, this species was reported from
Thailand, Christmas Island, the Northern Territory,
eastern Queensland, Taiwan and Tahiti (27).
New records
Thallus saxicolous, rimose to areolate, smooth, matt,
pale greyish to green, 20–80 μm thick, ecorticate;
prothallus present, grey to black. Perithecia
numerous, mostly solitary, semi-immersed to ±
superficial, convex to hemispherical or subconical,
0.2–0.6 mm diam., greenish-brown to black, not
overgrown by the thallus; ostiole inconspicuous or in a
shallow depression; centrum subglobose to ovate,
0.10–0.25 mm diam.; excipulum hyaline to pale brown
or yellowish-brown, 10–15 μm thick, subhymenium
20–40 μm thick; paraphyses unbranched, 0.8–1 μm
wide, periphyses absent. Asci cylindrical to elongate,
72–86 × 8–9 μm, ascospores 3-septate, cylindrical to
fusiform, 14–24 × 3–4.5 μm, lacking perispore.
Pycnidia present. (Fig. 2B).
1. Porina exserta Müll. Arg. in Flora 71: 548. 1888.
Thallus corticolous, continuous to rimose, pale
brownish to green, smooth to minutely rugulose, 35–
80 μm thick, ecorticate; prothallus lacking. Perithecial
verrucae, hemispherical to subglobose, 0.35–0.96 mm
diam., brownish to blackish, apex rounded; ostiole
usually inconspicuous; excipulum 20–30 μm thick,
pale orange-brown, centrum 0.2–0.4 mm wide;
subhymenium
15–35
μm
thick;
paraphyses
unbranched, 0.7–1.2 μm wide, periphyses absent. Asci
elongate to cylindrical 110–1160 × 14–18 μm,
ascospores elongate to cylindrical, (11–)15–17 (–21)
septate, 44–70 × 3.5–5.5 μm. Pycnidia absent. (Fig. 2A).
Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen
substance present in TLC.
Specimen examined:—INDIA, Goa: South Goa, Cotigao
Wildlife
Sanctuary,
Endrem,
N14°59ʹ51.2ʺ,
E
074°11ʹ52.1ʺ, 24 February 2018, P. Randive GU-L 813
(LWG Acc. no. 36250!), on the bark.
Note:—Porina
exserta
closely
resembles
P.
bellendenica Müll. Arg. in having similar perithecial
verrucae and ascospores, but P. bellendenica differs
in having a black basal layer and 15 septate
ascospores.
2. Porina siamensis P. M. McCarthy in Lichenologist
31(3): 242. 1999.
Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen
substance present in TLC.
PLANT SCIENCE TODAY
893
Fig. 2. Habit (a), perithecial section (b) and ascospres (c) of newly recorded pyrenocarpous lichens. A. Porina exserta, B. P. siamensis, C.
Pyrenula dissimulans. D. P. pyrenastrospora. E. P. rinodinospora (Scale bars: habit A, C–E 2 mm, B 5 mm; perithecial section A–E 200 µm;
ascospores A. 50µm, B – E 30 µm).
Distribution and ecology:—Porina siamensis was
found growing on an open, exposed rock in Cotigao
Wildlife Sanctuary at about ±200 m. Previously this
species was known from Thailand (28).
diam.) and ascospores are slightly larger (16–32 × 4–6
μm). In comparison, P. fortunata differs in having
seven septate ascospores of size 18–33 × 5–4 μm and
perithecia of 0.3–0.4 mm diam.
Specimen examined:—INDIA, Goa: South Goa, Cotigao
Wildlife Sanctuary, Cuncolim, Agonda, N15°13ʹ13.0ʺ,
E073°97ʹ99.5ʺ, 28 December 2016, P. Randive GU-L434
(LWG Acc. no. 36251!), on rock.
3. Pyrenula dissimulans (Müll. Arg.) R.C. Harris in
More Florida lichens, Incl. 10 Cent Tour Pyrenol.: 110.
1995.
Note:—This species is similar to Porina chlorotica
(Ach.) Müll. Arg. and P. fortunata P. M. McCarthy &
Etayo in having black or dark reddish-brown
perithecia. P. chlorotica also has three septate
ascospores, but its perithecia are smaller (0.2–0.3 mm
Thallus corticolous, corticated, smooth, continuous,
brownish to yellowish, lacking pseudocyphellae.
Perithecia simple, dispersed, conical, emergent, 0.25–
0.6 mm diam., black; ostioles apical black;
hamathecium hyaline. Asci cylindrical to clavate, 4–8
spored, ascospores muriform, 25–55 × 14–22 μm,
894
RANDIVE ET AL
lumina mostly rounded, at least in the central part of
ascospores. Pycnidia not seen. (Fig. 2C).
Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen
substance present in TLC.
Distribution and ecology:—This species was found
growing on the tree bark of T. paniculata in the Goa
University campus approximately at an altitude of 100
m. Earlier, this species was known from North and
South America (29-31).
Specimens examined:—INDIA, Goa: North Goa, Goa
University campus, 02 October 2016, P. Randive, GUL305 (LWG Acc. no. 36252!) on bark, GU- L318 (LWG
Acc. no. 36253!) on bark.
Note:— Pyrenula dissimulans is close to P. oleosa R.C.
Harris in having corticated thallus, apical ostioles and
lacking oil inspersion in hamathecium, but the old
ascospores in the latter species are filled with oil.
4. Pyrenula pyrenastrospora Aptroot in Biblthca
Lichenol. 64: 165, 1997.
Thallus corticolous, corticated, smooth, brownish,
lacking pseudocyphellae. Perithecia mostly aggregated
with fused walls, conical, 0.3–0.6mm diam.; ostioles
apical or when ecentric all opining the same direction,
black; hamathecium not inspersed. Asci cylindrical to
clavate, 4–8 spored, ascospores brown, 3-septate,
fusiform, 16–25 × 6–10 μm, lumina in a straight line,
terminal lumina directly against the exospores wall.
Pycnidia absent. (Fig. 2D).
Distribution and ecology:—This species was found
growing on the bark of Ficus benghalensis L. in
Coatigao Wildlife Sanctuary and en route to SattariValpoi between altitude 100–200 m. Previously this
species was known only from Papua New Guinea (32).
Specimens examined:—INDIA, Goa: South Goa, Cotigao
Wildlife
Sanctuary,
Quinomol,
N15°13ʹ130ʺ,
E074°11ʹ88.7ʺ, 07 December 2016, P. Randive GU-L414
(LWG Acc. no. 36255!) on bark; North Goa, SattariValpoi, N15°31ʹ66.7ʺ E074°12ʹ41.1ʺ, 07 March 2017, P.
Randive GU-L 548 (LWG Acc. no. 36256!), on bark.
Note:—Pyrenula rinodinospora closely resembles
Pyrenula maravalensis Vain. in having similar
morphology, but the latter species differs by the
shorter ascospores of 20–25 µm long (18).
Key to the pyrenocarpous lichens of Goa
1a.
Thallus foliicolous ………………………………….…. 2
1b.
Thallus otherwise …………………………..………… 17
2a.
Asci functionally unitunicate, entirely thinwalled,
perithecia
covered
by
thallus,
ascospores transversely 3–9 septate….…..…….. 3
2b.
Asci functionally bitunicate (fissitunicate),
apically thick-walled, perithecia not covered by
thallus, ascospores transversely 1-septate..… 10
3a.
Perithecia with 5–10 (–12) setae, soft, usually
decurved, narrowly acute or bristle-like, whitish
or black with white distal halves, ascospores
fusiform to narrowly oblong, 7 septate, 25–35 ×
4–6 µm ………………. Trichothelium alboatrum
3b.
Perithecia lacking setae ….…………………………. 4
4a.
Area around the ostiole or perithecial surface
rough, slightly papillose or tomentose ……...… 5
4b.
Area around the ostiole and perithecial surface
glabrous ……………...……...………………………..…. 6
5a.
Perithecial wall colourless, ascospores 5 septate,
fusiform, 22–30 × 4–5 µm ... Porina pallescens
5b.
Perithecial wall upper part blackish, lower part
brownish, ascospores 5 septate, fusiform, 20–32
× 4–5.5 µm ……………….……...…Porina nitidula
6a.
Ascospores 3 septate …..…………………………….. 7
6b.
Ascospores 7–9 septate ……...…..………………….. 8
7a.
Perithecial apex conical, translucent, ascospores
18–27 × 3–5 µm …………………..… Porina rufula
7b.
Perithecia apex rounded, not translucent,
ascospores
oblong,
14–20
×
2–4
µm
…………………………………….Porina chrysophora
8a.
Perithecia conical, apex with prominent, short
cylindrical extension, ascospores narrowly
fusiform to oblong, 34–48 7 × 4–6 µm
………………..………………………..… Porina conica
8b.
Perithecia apex without any extension ……….. 9
9a.
Perithecia conical to wart-shaped when mature,
with black dot around ostiole, ascospores
narrowly fusiform, 33–63 × 4–6 µm, white
prothallus sometimes present ……………….………..
………………………...………Porina karnatakensis
Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen
substance present in TLC.
Distribution and ecology:—This species was found
growing on the tree bark of Tectona grandis L.f. in
Cotigao Wildlife Sanctuary at altitude ±200 m.
Previously this species was known from Papua New
Guinea (19).
Specimen examined:—INDIA, Goa: South Goa, Cotigao
Wildlife
Sanctuary,
Bela
lake,
N14°57ʹ23.7ʺ,
E074°09ʹ09.3ʺ, 15 January 2018, P. Randive GU-L655
(LWG Acc. no.36254!), on bark.
Note:—Pyrenula pyrenastrospora is closely related P.
minarum Vain., which differs in having inspersed
hamathecium and partly eccentric ostioles.
5. Pyrenula rinodinospora Aptroot in Lichenologist
44(5): 611–618. 2012.
Thallus corticolous, corticate, smooth, continuous,
thin, brownish, without pseudocyphellae. Perithecia
simple, dispersed, conical, emergent, 0.3–0.5 mm
diam., black, edges without thallus covering; ostioles
black, apical; hamathecium hyaline, densely inspersed
with oil droplets. Asci cylindrical to clavate, 8-spored,
ascospores brown, 3-septate, fusiform, without
constrictions, 20–30 × 11–12.5 μm, ends mostly
pointed, lumina mostly quadrangular, angles blunt,
terminal lumina elongated and not separated from the
end wall by an endospore layer. Pycnidia not seen.
(Fig. 2E).
Chemistry:—Thallus K–, C–, KC–, PD–, UV–; no lichen
substance present in TLC.
PLANT SCIENCE TODAY
9b.
Perithecia lens shaped to hemispherical,
without dark spot, prothallus absent, ascospores
oblong, 26–33 × 3–4 µm ..….. Porina epiphylla
10a.
Ascospores large, 35–70 × 4–8 µm, often
breaking into halves (each cell often with up to
3 secondary septa), oblong, with distinct
constriction at septum ....Strigula janeirensis
20a.
21a.
Distal cell of the ascospore enlarged.…………. 12
11b. Distal cell not enlarged ...…………………..……… 13
12a.
Thallus subcuticular, bright green, ascospores
irregularly biseriate, 15–25 × 4–6 µm …....………...
…………………..…………….... Strigula antillarum
12b. Thallus epiphyllous, pale greenish to bluish
grey, ascospores biseriate, 15–25 × 4–6 µm ……….
……………………..……………... Strigula subelegans
13a.
Ascospores breaking into halves either inside or
outside the asci …………..………………………...… 14
Thallus thin (8–15 µm), bright metallic green,
margin effuse, with a thin blackish line,
ascospores 8–12 × 2–3 µm ….Strigula nitidula
14b. Thallus thick (15–30 µm), pale greyish green,
margin crenulate to lobulate, lacking blackish
line, ascospores 8–12 × 2–3 µm ………………………..
…………………………………….… Strigula concreta
15a.
Thallus elobate, photobiont cells rectangular to
angular-rounded, perithecia pure black, conical,
delimited from surrounding thallus, ascospores
9–12 × 2–3 µm ……………….Strigula phyllogena
15b. Thallus crenulate lobate to distinctly lobate . 16
16a.
Thallus with distinct lobes leaving small to large
interspaces,
greenish-brown,
perithecia
completely exposed but covered by thin thallus
layer, ascospores oblong acicullar, with a slight
constriction at septum, 10–18 × 2–3 µm .………….
……..….………………………..Strigula subtilissima
16b. Thallus entire to crenulate or lobulate,
sometimes whole thallus lobate-laciniate, bright
green, perithecia immersed to erumpent,
covered by algiferous thallus tissue up to
ostiolum, ascospores 14–24 × 4–6 µm …….………..
…………………………………..Strigula smaragdula
17a.
Thallus saxicolous ……………………………………. 18
22a.
Thallus squmulose, squmules imbricate, up to
2.5 mm wide, perithecia 1–6 per squamule,
ascospores 2 per asci, brown, muriform, 30–40 ×
12–14 µm …………... Endocarpon subrosettum
18b. Thallus crustose, continuous to areolate …… 19
19a.
Ascospores muriform, 2 per acus, brown, 70–90
× 15–25 µm, perithecia sunken, algal cell in the
hymenium globose, thallus rimose areolate,
greyish brown …………......… Staurothele fissa
19b. Ascospores transversely septate, hyaline ….. 20
Ascospores hyaline …………………………………..23
22b. Ascospores greyish brown to brown …….…… 57
23a.
Ascospores simple, subglobose to ellipsoid,
surface ornamented with cristae, 9–13 × 7–9 µm,
thallus slightly squamulose ………..……………….....
……….………….………..….Monoblastia pellucida
23b. Ascospores septate …………………………………...24
24a.
Ascospores transversely septate ….……………. 25
24b. Ascospores muriform ……………………..……….. 56
25a.
Ascospores 1-septate ………….…………………….. 26
25b. Ascospores 3 or more septate..………………….. 41
26a.
Perithecial wall hyphal, containing bark cells,
pseudoparaphyses
branched,
but
not
anastomosing …………………………………..…….. 27
26b. Perithecial wall cellular, lacking bark cells,
pseudoparaphyses slender, branched and
anastomosing especially above the asci ...….. 30
27a.
Ascospores 2 per asci, 1-septate, sometimes
constricted at both ends and appearing as 3
septate, large (40–60 × 15–25 µm) …………….….…..
……………..…………………….. Arthopyrenia finkii
27b. Ascospores 8 per asci ………………………...……. 28
28a.
Distal cell of the ascospore is larger, ascospore
30–45 × 14–18 µm, perithecia solitary……………….
……………………………….Arthopyrenia nidulans
28b. Both the cells are almost equal in size …...…. 29
29a.
17b. Thallus corticolous ………………………………….. 22
18a.
Ostiole conspicuous, periostiolar region brown
to black, perithecia 0.5 – 0.8 mm diam., 7–9
septate, 23–34 × 5–9 µm, fusiform with rounded
ends …………………….…….. Porina subinterstes
21b. Ostiole usually inconspicuous, periostiolar area
pale to dark brown or blackish, perithecia 0.30.8 mm diam., ascospores 7 septate, 24–44 × 4–7
µm, cylindrical or narrowly obclavate……………..
………………………………...…….Porina tetracerae
13b. Ascospores not breaking ………………………….. 15
14a.
Ascospores 3 septate, cylindrical to fusiform, 14–
24 × 3–5 µm, perithecia 0.2–0.6 mm diam.,
greenish-brown to black, not overgrown by the
thallus, ostiole inconspicuous or in a shallow
depression …………………..…. Porina siamensis
20b. Ascospores 7–9 septate …………………………….. 21
10b. Ascospores small, up to 25 µm long ….……….. 11
11a.
895
Ostiole mammilate, perithecia solitary to 2–3
aggregate, ascospores 25–43 × 6–11 µm…………….
……………………………...Arthopyrenia indusiata
29b. Ostiole not mammilate, perithecia solitary,
ascospores 10–18 × 5–7 µm ………..……..…………….
………..…………….....….. Arthopyrenia alboatra
30a.
Thallus UV+ ……………………………………...…….. 31
30b. Thallus UV– ...…………………….……………………. 35
31a.
Distal cell of the ascospores is larger, ascospores
23–30 × 7–10 µm, perithecia single, ostiole
eccentric .………………..…………………………………….
…………………………...Anisomeridium ubianum
31b. Both the cells are of almost equal size ...…….. 32
896
32a.
RANDIVE ET AL
Ostiole eccentric, perithecia single, ascospores
27–35 × 8–10 µm . Anisomeridium angulosum
44a.
32b. Ostiole apical ………………………………...………... 33
33a.
Ascospores smaller, 14–19 × 4–65 µm, narrowly
fusiform, ascus mostly biseriate……………………...
……………………...Anisomeridium albidoatrum
33b. Ascospores >20 µm long ……………….………….. 34
34a.
Ascospores 25–30 × 9–13 µm, elliptical to
broadly elliptical, ascus mostly uniseriate………..
……………………..Anisomeridium consobrinum
34b. Ascospores 30–45 × 13–16 µm, ovoid to broadly
ovoid, ends somewhat pointed ………………………..
……………………………..Anisomeridium indicum
35a.
Ascospores 2–4 per ascus, 40–66 × 10–17 µm,
perithecia solitary, ostiole black with white
wide pore……...Anisomeridium tarmugliense
44b. Thallus smooth to uneven, olive-green to
yellowish-brown, perithecia 0.2–0.3 mm diam.,
prominent to sessile, subglobose to barrelshaped with a flattened top, ostiolar area
greyish, ascospores fusiform-ellipsoid, 20–25 ×
7–10 μm …………..… Nigrovothelium tropicum
45a.
Distal cell of ascospore larger in size ….……… 37
36b. Both the cells of ascospores are of almost equal
in size ………………………..………..…………………. 38
37a.
Ascospore 3 septate, perithecia solitary or 2–3
aggregated, ostiole apical, umbonate, 19–23 × 5–
7 µm, thallus endophloeodal…………………………...
……………………………….… Arthopyrenia grisea
45b. Ascospores more than 3 septate …………………. 46
46a.
35b. Ascospores 8 per ascus ………………………..…... 36
36a.
Thallus strongly verrucose-bullate, olive-green
to brownish, perithecia 0.2–0.3 mm diam.,
sessile, stioles visible as tiny black dots,
ascospores fusiform, 20–25 × 8–10 µm ……………..
…………………………...Nigrovothelium bullatum
Perithecia highly crowded to pseudostromatic
…………..…………………………………...…..…………. 47
46b. Perithecia solitary or aggregated, never in
pseudostroma, but in verrucae ……...…....…… 48
47a.
Ostiole eccentric, ascospores 19–30 × 7–12 µm
……...…….………… Anisomeridium palavanum
37b. Ostiole apical, ascospores 16–21 × 4–7 µm
………………..…………... Anisomeridium biforme
Perithecia in diffuse pseudostroma, brownblack, 0.5–1.0 mm, basally covered by thallus
but upper part exposed, flattened and discshaped, ascospores 12–17-septate, fusiform, 45–
55 × 6–8 μm, thallus olive-green to yellowish
……..…………… Trypethelium plicatorimosum
38b. Ostiole eccentric ……………………………………….39
47b. Perithecia in distinct pseudostroma, brownish
to dark brown, covered by yellow pigment, 1–2
mm diam., ascospores 9–13-septate, fusiform,
37–52 × 8–11 μm……………………………………………..
……….…….…………….... Trypethelium eluteriae
39a.
48a.
38a.
Ostiole apical, ascospores 23–27 × 9–11 µm,
fusiform, biseriate or irregularly arranged in
ascus ….……………... Anisomeridum subnexum
Ascospores granular ornamented, fusiform to
narrowly ovoid, 19–30 × 6–8 µm……………………...
……………….……..Anisomeridium terminatum
Ostiole K+ reddish, perithecia 0.45-0.6 mm
diam., ascospores 7 septate, elliptic, 19–30 × 4–7
μm …………………….Clathroporina mastoidea
39b. Ascospores smooth ……...…………………………...40
48b. Ostiole K- …………...…………………………………... 49
40a.
Ascospores fusiform, with pointed ends, 30–37 ×
9–11 µm ………... Anisomeridium glaucescens
49a.
40b. Ascospores 30–40 × 6–8 µm ……………………………..
…………………….Anisomeridium complanatum
50a.
41a.
42a.
Perithecia in distinct or diffuse pseudostroma,
laterally covered by thallus ……………..…….…. 43
42b. Perithecia solitary to crowded, fully exposed,
black ………………………...….……………………...… 44
43a.
49b. Ascospores 6–13 septa ……………………………….51
Ascospores lumina diamond-shaped ……….... 42
41b. Ascospores lumina rectangular-shaped ………45
Ascospores >13 septa ……………………………….. 50
Perithecia
brownish
blackish,
ostiole
inconspicuous, ascospores 11–21 septa, elongate
to cylindrical, 47–70 × 4–6 μm………………………….
…………………………………………….Porina exserta
50b. Perithecia greenish grey, area around ostiole
black, ascospores 13-16 septate, straight to
curved with pointed ends, 50–70 × 5–7 μm
…………………….………….… Porina subhibernica
51a.
Hamathecium
inspersed
with
oil
…………………………..……….……………….…………. 52
Perithecia in diffuse stroma, upper portion
blackish brown with whitish rim, erumpent, 0.30.5 mm diam., hamathecium inspersed,
ascospores 3 septate, 15–27 × 7–10 µm, fusiformellipsoid ………………...…...Astrothelium scoria
51b. Hamathecium
not
inspersed
………………………………...………….….….………….. 53
43b. Perithecia in distinct stroma, upper portion
dark, erumpent to prominent, 0.8-1.5 mm
broad, ostiolar region internally with yellow
pigment, hamathecium clear, ascospores 7–11
septate, fusiform, 55–88 × 13–28 µm………………...
…………………..……………Astrothelium luridum
52b. Ascospores 11–12 septate, fusiform, 89–92 × 16–
20 μm, ostiolar region black…………………………….
…………..………………………....Porina americana
52a.
53a.
Ascospores 7–9 septate, oblong ellipsoid, 60–85 ×
12–20 μm, ostiolar region brown black…………….
……………………………………...Porina subcutanea
Ostiolar region yellow to light brown,
surrounded by black priostiolar rim, ascospores
PLANT SCIENCE TODAY
7–9 septate, fusiform with acute ends, 43–50 × 4–
6 μm ……..………………....... Porina ochrostoma
63a.
53b. Ostiolar region variously coloured, peri ostiolar
rim lacking ………………………..………….………… 54
54a.
Ostiolar region brown-black, ascospores 7–8
septate, fusiform, acute ends, 32–48 × 4–7 μm…..
…………..…………………………….Porina interestes
Ostiolar region orange, pinkish, reddish to
brownish, rarely dark brown, ascospores 7–9 (–
13) septate, fusiform with rounded ends, 44–80 ×
7–13 μm ………………………...… Porina atlantica
55b. Ostiolar region orange-brown to black to black,
ascospores fusform round ended, 45–60 ×11–18
μm …………………………….. Porina internigrans
56a.
Thallus olive-green, yellow-orange pruinose,
UV+, perithecia flattened top, solitary to
irregularly grouped and confluent, covered with
yellow-orange pigment, hamathecium densely
inspersed, ascospores fusiform-ellipsoid, 50–80
× 17–23 µm ……………Marcelaria benguelensis
Thallus pseudocyphellate, yellow-brown buff,
perithecia solitary or 1–4 coalescing, ostiole
indistinct, ascospores oblong ellipsoid, 32–45(–
60) × 15–27 µm ………….…… Pyrenula immissa
63b. Thallus lacking pseuodcyphellae ………………. 64
64a.
54b. Ostiolar region otherwise, ascospores much
larger ……………………….……………………..……… 55
55a.
897
Ascospore 27–50 × 15–18 µm, oblong ellipsoid,
perithecia solitary, completely embedded in
thalline verrucae, ostiole indistinct, thallus
ochre-yellowish brown …... Pyrenula oculata
64b. Ascospore <30 µm long ……………………………..65
65a.
End lumina of the ascospore is elongated …..66
65b. All lumina ± rounded to angular …...…………. 67
66a.
Thallus brownish to pale brown, ascomata
emergent 0.3–0.5 mm diam., hemithecium
densely impressed with oil droplets, ascospores
20–25 × 6–10 µm .…… Pyrenula maravalensis
66b. Thallus ochraceous yellow to brownish,
ascomata semiemergent 0.5- 1.0 mm diam.,
hemithecium without oil droplets, ascospores
20–30 × 11–13 μm .... Pyrenula rinodinospora
67a.
Ostiole mamillate papillate, dull black,
perithecia solitary, ascospores oblong ellipsoid,
17‒20×7‒9 µm, thallus buff to yellow-brown …..
………….………………………….Pyrenula mamillan
56b. Thallus light olive-green, pigmented pruina
absent, UV+, flattened top, solitary to grouped,
covered by thallus up to ostiole, hamathecium
inspersed, ascospores oblong-fusiform, 120–220
× 25–40 µm, with thickened median septum……..
………………….… Astrothelium meristosporum
67b. Ostiole otherwise, centrum I ± perithecia
solitary or semi-solitary, ascospores ellipsoid,
thallus brownish yellow to brown …...………. 68
57a.
68a.
Ascospores transversely 3 septate ……..……… 58
57b. Ascospores submuriform to muriform ……… 74
58a.
Perithecia in pseudostroma, 1–12 carpic, thallus
yellowish-brown to brown, K+ red, ascospores
ellipsoidal, 13–23 × 6–10 µm ……..………………...
………………………………..… Pyrenula leucotrypa
58b. Perithecia solitary or aggregated, not forming
pseudostroma ……………..…..…..……...…………. 59
59a.
Ascomata mostly aggregated, with fused walls
but with separate ostioles, conical, 0.3–0.6 mm
diam., ostioles apical, black, ascospores brown,
3 septate, fusiform 16–25 × 6–10 μm………………...
…………………………Pyrenula pyrenastrospora
59b. Ascomata solitary ……………………………………. 60
60a.
Ascospores red-brown, lumina becoming
rounded when older, 15–20 × 9–10 µm, ellipsoid,
hamathecium inspersed, thallus olive-brown
shining…………….…….Lithothelium decumbens
68b. Psudocyphellae absent, ascocarp 1.0–2.5 (–3.0)
mm in diameter, flat, convex to conicodepressed, centrum I -, ascospores 15–17 × 5–7
µm ……..………………....……. Pyrenula castanea
69a.
Ostiole eccentric, perithecia solitary, ascospores
35–45 × 15–18 µm, terminal lumina separated
from the exospore wall by endospore thickening
………...….………………………….. Pyrenula adacta
61b. Ostiole apical ……………………………………….…. 62
62a.
Hamathecium inspersed ………..………………... 63
62b. Hamathecium not inspersed……………………….69
Thallus with pseudocyphellae, brownish to
olive-green, ascospore fusiform, 22–38 × 8–15
μm, lumina rounded to somewhat angular
…………..…….…….....…. Pyrenula quassiaecola
69b. Thallus lacking pseudocyphellae ….……...…… 70
70a.
Terminal lumina mostly separated from
exospore wall …….………………...……….………… 71
70b. Terminal lumina not separated from exospore
wall ….………….....……………………………..………. 73
71a.
60b. Ascospores grey to brown, rarely red-brown
and then lumina angular …...……..………..…… 61
61a.
Psudocyphellae present, ascocarp 0.8–1.0 mm in
diameter, conical- hemispherical, centrum I+
blue, ascospores 16–24 × 8–13 µm …………………...
………………...……………………..Pyrenula fetivica
Perithecia slightly larger, 0.6–1.0 mm diam.,
ascospores oblong ellipsoid, 12–17 × 5–7 μm …….
………………………………….…...Pyrenula brunnea
71b. Perithecia smaller ………….……………………….. 72
72a.
Perithecia 0.2–0.6 mm diam., ascospores oblong
ellipsoid, 17–22 × 9–10 μm …Pyrenula aspistea
72b. Perithecia 0.4–0.7 mm diam., ascospores oblong
ellipsoid, 15–21 × 8–10 μm……………………………….
…………………………………....Pyrenula aggregata
73a.
End lumina elongated, ascospore 20–29 ×9–11
μm ……...………….….…. Pyrenula approximans
898
RANDIVE ET AL
73b. All lumina ± rounded to angular, ascospores 13–
16 × 7–8 μm ………………....… Pyrenula nitidula
74a.
Ostioles eccentric, fused, perithecia usually in
groups of 2–6, ascospores submuriform, with 5
transverse septa and few longitudinal septa,
fusiform, with pointed ends, 24–32 × 10–13 μm,
thallus yellowish to olive-green ……………............
……………………………. Pyrenula subumbilicata
74b. Ostiole apical, not fused, perithecia mostly
solitary ……………..…………..……………...……….. 75
75a.
Thallus and perithecia yellow to orange,
indistinctly to faintly papillate, ascospores
ellipsoidal, 12–18 × 7–10 μm …………….……………..
………………………….... Pyrenula ochraceoflava
75b. Thallus and perithecia otherwise ……………….76
76a. Old ascospores filled with orange oil, muriform,
25–30 × 10–15 μm, pantropical distribution
…………………………………...… Pyrenula breutelii
76b. Old
ascospores
lacking
orange
oil,
submuriform..…….………………….…………..…... 77
77a. Thallus yellow-brown, pseudocyphellate, UV+
yellowish, ostioles indistinct to mamillatepapillate,
hamethecium
not
inspresed,
ascospores 23–35 × 10–12 μm long ……………..…...
…………………………...…… Pyrenula gibberulosa
77b. Thallus otherwise, ostioles apical, ascospores
larger, up to 45–55 μm long ……………………... 78
78a. Lumina of the ascospore mostly round,
ascospore 25–55 × 14–22 μm, ostioles apical,
naked,
black,
thallus
yellowishbrown……………………….Pyrenula dissimulans
78b. Lumina relatively large and angular, ascospore
24–46 × 9–20 μm, ostiolar region naked,
brownish-black to black, thallus greenish straw
coloured, yellow to yellow-brown …………………...
…………………………………..Pyrenula leucostoma
Discussion
The semi-evergreen forest in the Goa region exhibits
maximum diversity of pyrenocarpous species. The
trees along the streams in moist, shady habitats bear
luxuriant growth of pyrenolichens which mostly
belong to Porina and Pyrenula species. The occurrence
of pyrenocarpous lichens indicates the abundant
presence of smooth-barked trees in the State. Further,
the occurrence of a large number of foliicolous lichens
indicates that the State has several healthy and
undisturbed forests. Bhagwan Mahavir Wildlife
Sanctuary as a whole and within Cotigao Wildlife
Sanctuary, Avem, Bela to Zombolim,, Endrem to
Tulshimol, Kuske waterfall area, Marlem, Nadkem and
Ravan Donger are few such pristine habitats. With the
addition of five new records, the total number of
lichens species in Goa State increases to 165 species,
out of which pyrenocarpous lichens represent almost
half (48%).
Ghats forests of Goa. It can be noted that among the
protected areas within Goa, Cotigao Wildlife
Sanctuary represents more number of pyrenocarpous
lichens. Further, along with the forest areas within
Goa, the coconut, arecanut orchards and several
cultivated plants also provide suitable habitats for
lichen growth. The present study will be highly useful
for monographic studies on pyrenocarpous lichens of
India or the world and for environmental monitoring
studies in the area.
Acknowledgements
We are thankful to Director, CSIR-NBRI, Lucknow, for
providing the laboratory facilities for the research, to
Publication Ethics Committee of CSIR-NBRI for
carefully scrutinizing the manuscript and approving it
for publication with ms number MS/2020/08/03.
Authors PR and MKJ are thankful to Goa University for
financial assistance under the Ph.D. programme and
to the Forest Department of Goa for permission to
survey Cotigao Wildlife Sanctuary. PR is also thankful
to DST for financial assistance under Women Scientist
Scheme (SR/WOS-A/LS-354/2016). GKM is thankful to
SERB for financial assistance under the NPDF scheme
(PDF/2017/000356).
Authors’ contributions
PR collected fresh samples of pyrenocarpous lichens
from Goa and carried out their preliminary
identification. GKM studied pyrenocarpous lichen
specimens preserved in herbarium LWG and drafted
the manuscript. SN authenticated identity of some
pyrenocaprous lichens reported in the manuscript,
improved the manuscript and supervised the whole
study. DKU authenticated identity of some
pyrenocarpous lichen specimens and suggested
improvement in the manuscript. MKJ conceptualized
the idea of studying lichens of Goa, supervised the
work of author PR and provided intellectual inputs to
the study.
Conflict of interests
The authors do not have any conflict of interests to
declare.
References
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Randive P, Nayaka S, Janarthanam MK. An updated checklist of
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Randive R, Joseph S, Nayaka S, Janarthanam MK. Notes on foliicolous lichens from Western Ghats part of Goa, India. Indian
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PLANT SCIENCE TODAY
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Pyrenula. Lichenologist. 2012;44(1):5-53. https://doi.org/10.1017/
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To cite this article: Randive P, Mishra G K, Nayaka S, Upreti D K, Janarthanam M K.
Pyrenocarpous lichens in Goa with five new records to India. Plant Science Today.
2021;8(4):889–899. https://doi.org/10.14719/pst.2021.8.4.1265
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