Cryptogam Biodiversity and Assessment
Cryptogam Biodiversity and Assessment
Special Volume (2018), e-ISSN :2456-0251, 37-46
New records and an updated key of Pyrenula from India
Komal K. Ingle1,2*, Vindhyeshwari Uppadhyay1,2, Sanjeeva Nayaka1, Suman Trivedi2 and Dinabandhu Sahoo3
1
Lichenology Laboratory, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow- 226 001, India
Department of Botany, Motilal Vigyan Mahavidyalaya, Bhopal- 462 026, India
3
Institute of Bioresources and Sustainable Development, Imphal, India
2
Publication Info
Article history:
Received : 10.12.2017
Accepted : 11.12.2017
DOI: https://doi.org/10.21756/
cab.esp7
ABSTRACT
Pyrenula andina, P. atropurpurea, P. bahiana , P. caracasana, P. macrospora, P. massariospora,
P. pseudobufonia , P. subducta, P. subgregantula and P. tristissima are described as new records
for Indian lichen flora. An updated key to all Pyrenula species so far known from India is also
presented.
Key words:
Pyrenulaceae, India, Taxonomy,
Lichenized Fungi
*Corresponding author:
Email: ingle.komal@gmail.com
INTRODUCTION
layer of endospore, and presence or absence of dark bands
between the septa (Aptroot 2012; Mendonça et al. 2016).
The lichen genus Pyrenula typically found on smooth
and shaded bark mostly in evergreen forest is represented
by ca. 226 species in the world (Aptroot 2012; Aptroot et al.
2013, 2015; Mendonça et al. 2016; Aptroot and Common
2017), of which 92 are known from India (Upreti 1998, Singh
and Sinha 2010, Jagadeesh Ram and Sinha 2010). Following
the recent world key of the species of Anthracothecium
and Pyrenula by Aptroot (2012), at present only 67 species
are known from India. The addition of present ten new
records of Pyrenula species further incresed the number of
species to 77.
Most of the species of Pyrenula described from India
were found growing on bark exhibit their distribution mostly
in the Himalayan regions and Western Ghats of India. The
species prefers to grow on smooth bark trees of Alnus,
Quercus and Ficus spp., on trunk, thick branches and twigs.
Among the Quercus species, Q. semecarpifolia having
smooth twigs and bark provides excellent habitat for many
species.
The present investigation resulted addition of ten
species of Pyrenula viz. Pyrenula andina Aptroot, P.
atropurpurea (Eschw.) Müll. Arg., P. bahiana Malme, P.
caracasana Müll. Arg., P. macrospora (Degel.) Coppins &
P. James, P. massariospora (Starbäck) R.C. Harris, P.
pseudobufonia (Rehm) R.C. Harris, P. subducta (Nyl.) Müll.
Arg., P. subgregantula Müll. Arg. and P. tristissima Vain. as
new record to the Indian lichen flora. An account on new
records together with an updated key of genus Pyrenula
from India is also provided.
Pyrenula is mainly a tropical and subtropical genus
(Mendonça et al. 2016), and neotropics are the centre of
diversity of the genus (Aptroot 2012). The genus is
characterized by whitish, browninsh to yellowish, ecorticate
or corticate, UV– or UV+ yellow thallus, with or without
pseudocyphellae, with or without lichexanthone or
anthraquinones; trentepohlioid type of alga; perithecioid
ascomata with uniformely carbonized wall, conical to
globose or flattened, solitary or aggregated with common
walls but separate ostiole or joined with partly common
ostioles; hamathecium with or without oil globules;
uniseriate or biseriate asci; ellipsoid or fusiform to elongate,
grey to brown, with or withour cilia, transversely septate to
sub-muriform or muriform ascospores with rounded or
pointed ends, rounded, diamond-shaped or elongated
lumina, directly against the exospore wall or separated by a
MATERIALS AND METHODS
The present study is based on collection in LWG as
well as fresh specimens collected on a field trip to Manipur
and Nagaland, and deposited in the herbarium of CSIRNational Botanical Research Institute, Lucknow (LWG).
Morphological characters were examined on dry material
under a stereo-zoom microscope, Leica S8APO. Thin handmyresearchjournals.com
37
Ingle et al.
cut sections of apothecia and thallus were mounted in plain
water, cotton blue, 10% KOH and iodine solution and
observed under a compound microscope Leica DM500.
ascospores, 12–14 × 8–10 µm, with rounded to pointed ends
having lumina in a straight line and termianl lumina mostly
separated from the exospore by an endospore layer.
The New Records
Remarks: Pyrenula atropurpurea (Eschw.) Müll. Arg. is
close to P. aspistea (Ach.) Ach., but the latter has smaller
ascomata (0.7 mm) and shorter ascospores (4–6 µm). This
pantropical species has been reported from humid and
tropical climate of Western Ghats region, where it was found
growing on Ficus tree trunk at an altitude of 1449 m.
Pyrenula andina Aptroot, in Aptroot, Lücking, Sipman,
Umaña & Chaves, Biblthca Lichenol. 97: 91 (2008) Fig. 1A
This species is characterized by yellowish-grey,
smooth, UV+ yellow thallus, immersed to semi-immersed,
globose to sub-globose perithecia, 0.2–0.6 mm diam. with
apical ostiole, completely carbonized laterally spreading
involucrellum, 22–34 µm thick at the base and 64–98 µm
thick laterally, colourless hamathecium without oil globules,
8-spored asci and 3-septate, fusiform ascospores, 24–40 ×
13–17 µm, with rounded ends having lumina in a straight
line and terminal lumina mostly separated from exospore
wall by an endospore layer.
Remarks: Pyrenula andina Aptroot, is closely resembles
P. dermatodes (Borrer) Schaer., which also has corticated,
UV+ yellow thallus, apical ostiole and hamathecium without
oil globules, but differs in the presence of shorter ascospores
(24 µm). This neotropical species was found growing on
tree bark between altitude of 773–1842 m in humid and
tropical climate of North-East Himalayan region.
Specimen examined: India, Tamil Nadu, Coimbtore, Siruvani
waterfalls, N 10º 52' E 76º 41', alt. 1449 m, on Ficus, 11 January
2008, H.T. Lumbsch, D.K. Upreti, P. Divakar and J. Tandon
19706/O/1 (LWG).
Pyrenula bahiana Malme, Ark. Bot. 22A(no. 11): 26 (1929)
Fig. 1C
This species is characterized by a greenish-grey,
smooth, UV– thallus, immersed to semi-immersed,
subglobose perithecia, 0.2–0.5 mm diam with apical ostiole,
completely carbonized involucrellum, 56–80 µm thick,
colourless hamathecium without oil globules, 8-spored asci
and 3-septate, fusiform ascospores, 29–38 × 13–18 µm, with
orange oil, ronded ends and terminal lumina separated from
the exospore wall by an endospore layer.
Remarks: Pyrenula bahiana Malme is similar to P.
thailandica Aptroot in having corticated thallus, apical
ostiole and 3-septate ascospores with orange oil, but the
latter differs in having longer ascospores (35–51 m long).
This pantropical species was found growing on tree bark
between altitude of 665–1641 m in humid and tropical climate
of North-East Himalayan region.
Specimen examined: India, Manipur, Ukhrul district, Samsai,
o
o
N 25 09' 14.7" E 94 23' 33.2", alt. 1338 m, on bark, 04 June
2017, Komal K. Ingle 17-031125/A, 17-031125/B (LWG),
Hungpung village, Nungshang river area, near cave, N 25o
00' 09.6" E 94o 19' 55.9", alt. 1641 m, on bark, 11 June 2017,
Komal K. Ingle 17-031126 (LWG); Bishnupur district,
o
Inspection bunglow, Keibul Lamjao National Park, N 24 28'
32.9" E 93o 48' 47.3", alt. 773 m, on bark, 18 June 2017, Komal
K. Ingle 17-031127 (LWG); Tamenglong district, Kabonram
village, N 25o 07' 31.0" E 93o 37' 41.5", alt. 1367 m, on bark, 24
September 2017, Komal K. Ingle 17-031128 (LWG). Nagaland,
Kohima district, Vaisema to Dzukou valley, N 25o 33' 24.9" E
o
94 08' 03.0", alt. 1842 m, on bark, 25 June 2017, Komal K.
o
o
Ingle 17-031129 (LWG), N 25 33' 25.5" E 94 08' 24.0", alt.
1723 m, on bark, 25 June 2017, Komal K. Ingle 17-031130
(LWG).
Specimen examined: India, Manipur, Ukhrul district,
Hungpung village, Nungshang river area, near cave, N 25o
00' 09.6" E 94o 19' 55.9", alt. 1641 m, on bark, 11 June 2017,
Komal K. Ingle 17-031131 (LWG); Senapati district, Govaganj
village, 25o 09' 43.8" E 93o 45' 50.5", alt. 665 m, on bark, 25
September 2017, Komal K. Ingle 17-031132 (LWG).
Pyrenula caracasana Müll. Arg., Flora, Regensburg 68: 337
(1885) Fig. 1D
This species is characterized by a light brown to fawn,
smooth, UV– thallus, convex, plane to rounded perithecia,
0.5–0.8 mm diam with apical ostiole, completely carbonized
laterally spreading involucrellum, 92–135 µm thick at the
base and 180–260 µm thick laterally, colourless hamathecium
inspersed with oil globules, 8-spored asci and 4-7 septate,
fusiform ascospores, 35–65 × 16–22 µm, with papillate ends
and termianl lumina mostly not separated from the exospore
by an endospore layer.
Pyrenula atropurpurea (Eschw.) Müll. Arg., Flora,
Regensburg 67(35): 665 (1884) Fig. 1B
This species is characterized by a yellowish-grey to
yellowish-green, smooth, UV– thallus, immersed to semiimmersed, globose to sub-globose perithecia, 0.5–0.8 mm
diam. with apical ostiole, completely carbonized laterally
spreading involucrellum, 35–40 µm thick at the base and
70–102 µm thick laterally, colourless hamathecium without
oil globules, 8-spored asci and 3-septate, fusiform
38
Cryptogam Biodiversity and Assessment
Fig. 1. A–F. Habitus. A: Pyrenula andina Aptroot; B: Pyrenula atropurpurea (Eschw.) Müll. Arg.; C: Pyrenula bahiana Malme;
D. Pyrenula caracasana Müll. Arg.; E: Pyrenula macrospora (Degel.) Coppins & P. James. F: Pyrenula massariospora
(Starbäck) R.C. Harris. Scale bars: A = 0.5 mm. C & F = 1 mm. B, D & E = 2 mm.
39
Ingle et al.
Remarks: Pyrenula caracasana Müll. Arg. resembles P.
wilmsiana Müll. Arg., which differs by shorter ascospores
(30 m long). This pantropical species was found growing
on the bark of tree at an altitude of 772 m in Eastern Himalayan
region.
carbonized involucrellum, 38–120 µm thick, hamathecium
inspersed with oil globules, 8-spored asci and 3-septate
fusiform ascospores, 20–35 × 10–18 µm with rounded to
pointed ends having lumina in a straight line and termianl
lumina mostly separated from the exospore by an endospore
layer.
Specimen examined: India, West Bengal, Darjeeling district,
Chunabhatti, N 26º 50' 45.3" E 88º 20' 16.5", 772 m, on bark,
14.04.2014, R. Bajpai 14-022008 (LWG).
Remarks: Pyrenula massariospora (Starbäck) R.C. Harris
appears close to P. acutalis R.C. Harris (Starb.) R.C. Harris
in corticated thallus, hamathecium inspersed with oil
globules, similar size and septation of ascospores with
elongated central lumina, but differs in pointed ends of
ascospores and biseriate arrangement in the ascus. This
pantropical species was found growing on tree bark from
North-East Himalayan region, between altitudes of 1598–
2570 m.
Pyrenula macrospora (Degel.) Coppins & P. James,
Lichenologist 12(1): 107 (1980) Fig. 1E
This specie is characterized by a light-brown to
greenish-grey, smooth, somewhat waxy, UV– thallus with
pseudocyphellae, immersed to erumpent, globose perithecia,
0.5–1.0 mm diam with apical ostiole, completely carbonized
involucrellum, 53–120 µm thick, colourless hamathecium
without oil globules, 8-spored asci and 3-septate, fusiform
ascospores, 30–38 × 11–14 µm, with rounded to pointed
ends having lumina in a straight line and termianl lumina
mostly separated from the exospore by an endospore layer.
Specimen examind: India, Manipur, Ukhrul district, Talui, N
o
o
25 11' 43.6" E 94 19' 53.6", alt. 1598 m, on bark, 15 June 2017,
Komal K. Ingle 17-031133 (LWG). Tuensang district, Mount
o
o
Saramati, N 25 46.175' E 94 58.272', alt. 2570 m, on bark, 15
June 2017, Komal K. Ingle 17-031134 (LWG).
Pyrenula pseudobufonia (Rehm) R.C. Harris, Evansia 2(3):
46 (1985) Fig. 2A
Remarks: Pyrenula macrospora (Degel.) Coppins & P.
James exhibits its close resemblance to P. complanata
(Mont.) Trevis. in having UV– thallus, hamathecium without
oil globules and similar size and septation of ascospores,
but the latter differs in having smaller ascomata size (0.7
mm diam) and thallus without pseudocyphallae. This
temperate species is distributed throughout the South and
West British Island and also known from Portugal and South
Scandinavia (Smith et al. 2009). It is reported from the
temperate Central Himalayan region, where it was found
growing on Alnus tree and also from temperate Eastern
Himalayan region, where it was found growing on bark of
tree between altitude of 1627-1800 m.
This species is characterized by pale yellowish-grey,
smooth, UV+ yellow thallus, immersed to semi-immersed,
globose perithecia, 0.5–1.0 mm diam with apical ostiole,
completely carbonized laterally spreading involucrellum, 75–
94 µm thick at the base and 113–172 µm thick laterally,
colourless hamathecium inspersed with oil globules, 8spored asci and 3-septate fusiform ascospores, 16–23 × 9–
13 µm with rounded ends having lumina in a straight line
and termianl lumina all against the exospore wall.
Remarks: Pyrenula pseudobufonia (Rehm) R.C. Harris is
closely related P. cayennensis Müll. Arg., which differs in
having UV– thallus and shorter ascospores (16 m long).
This species is distributed in northern temperate regions of
the world (Aptroot 2012). It is reported from humid and
tropical climate of North-East Himalayan region, where it
was found growing on bark of tree at an altitude of 1059 m.
Specimen examind: India, Uttarakhand, Rudraprayag, 4 Km
before Chopta, near HAPPRC medicinal nursery, on Alnus,
01.10.2013, D.K. Upreti & R. Bajpai 13-019928 (LWG). West
Bengal, Darjeeling district, Kalimpong, Dello hill, N 27º 05'
11.8" E 88º 30' 32.8", 1630 m, on bark, 18.04.2014, R. Bajpai
14-022057, 14-022058/A (LWG); Kurseong, N 26º 54' 21.2" E
88º 17' 30.5", 1627 m, on bark, 14.04.2014, R. Bajpai 14-022096
(LWG); Kurseong, 8 mile, N 26º 57' 08.4" E 88º 17' 30.7", 1800
m, on bark, 14.04.2014, R. Bajpai 14-022028, 14-022029;
Mungpoo, 4 Km away, N 26º 58' 24.9" E 88º 21' 01.2", 1642 m,
on bark, 16.04.2014, R. Bajpai 14-022069 (LWG).
Pyrenula massariospora (Starbäck) R.C. Harris, Mem. N. Y.
bot. Gdn 49: 95 (1989) Fig. 1F
Specimen examined: India, Manipur, Tamenglong district,
Dailong village, forest area, N 25o 05' 42.1" E 94o 21' 41.0",
alt. 1059 m, on bark, 21 September 2017, Komal K. Ingle 17031135 (LWG).
Pyrenula subducta (Nyl.) Müll. Arg., Flora, Regensburg
67:666 (1884) Fig. 2B
This species is characterized by a brown to reddishbrown, smooth, UV– thallus, immersed to erumpent, globose
perithecia, 0.3–0.8 mm diam with apical ostiole, completely
carbonized involucrellum, 38–120 µm thick, colourless
This species is characterized by a brownish to olivegreen, smooth, UV– thallus, immersed to erumpent, globose
perithecia, 0.3–0.5 mm diam with apical ostiole, completely
40
Cryptogam Biodiversity and Assessment
Fig. 2. A–D. Habitus. A: Pyrenula pseudobufonia (Rehm) R.C. Harris; B: Pyrenula subducta (Nyl.) Müll. Arg.; C: Pyrenula
subgregantula Müll. Arg.; D: Pyrenula tristissima Vain. Scale bars: A & B = 2 mm. C & D = 0.5 mm.
hamathecium without oil globules, 8-spored asci and 3septate, fusiform ascospores, 36–45 × 11–20 µm with
rounded to pointed ends having lumina in a straight line
and termianl lumina mostly separated from the exospore by
an endospore layer.
Specimen examined: India, Uttarakhand, Pithoragarh
district, near Maupani way to Askote, 5500 ft., D.D. Awasthi
3978 (LWG-AWAS); Manipur, Ukhrul district, Samsai, N 25o
09' 14.7" E 94o 23' 33.2", alt. 1338 m, on bark, 04 June 2017,
Komal K. Ingle 17-031136 (LWG), Lunghar village, N 25o 10'
o
22.0" E 94 26' 16.9", alt. 1789 m, on bark, 14 June 2017,
Komal K. Ingle 17-031137 (LWG), Bishnupur district,
Inspection bunglow, Keibul Lamjao National Park, N 24o 28'
32.9" E 93o 48' 47.3", alt. 773 m, on bark, 18 June 2017, Komal
K. Ingle 17-031138, 17-031139 (LWG)
Remarks: Pyrenula subducta (Nyl.) Müll. Arg. is very close
to P. complanata (Mont.) Trevis. in having corticatd thallus,
apical ostiole and hamathecium without oil globules, but
the latter differs in having longer ascomatal size (0.7 mm)
and smaller ascospores (29–40 µm long). This neotropical
species has been reported from the temperate Central
Himalayan regions of India, where it was found growing on
tree bark at an altitude of 5500 ft. It is also reported from
North-East Himalayan regions of India where it was found
growing on bark of tree trunk between altitude of 773–1789
m.
Pyrenula subgregantula Müll. Arg., Bull. Soc. R. Bot. Belg.
30: 94 (1891) Fig. 2C
This species is characterized by a yellowish to olivegreen, smooth, UV– thallus with pseudocyphellae, immersed
to erumpent, fused perithecia, 0.3–0.6 mm diam with lateral
ostiole, completely carbonized involucrellum, 30–52 µm thick,
41
Ingle et al.
colourless hamathecium without oil globules, 8-spored asci
and 3-septate, fusiform ascospores, 18–25 × 8–10 µm, with
rounded to pointed ends having lumina in a straight line
and termianl lumina all against the exospore wall.
This species is characterized by greenish-grey,
smooth, UV– thallus, immersed to semi-immersed, globose
perithecia, 0.1–0.3 mm diam with apical ostiole, completely
carbonized involucrellum, 39–66 µm thick, colourless
hamathecium without oil globules, 8-spored asci and 3septate, fusiform ascospores, 18–20 × 6–8 µm, with rounded
to pointed ends having lumina in a straight line and terminal
lumina mostly separated from the exospore by an endospore
layer.
Remarks: Pyrenula subgregantula Müll. Arg. shows its
resemblance to P. circumfiniens Vain., in having lateral
ostiole, similar size and septation of ascospores with
terminal lumina all against the exospore wall, but differs in
having single ascomata. The species is previouly known
from Cuba (Rosabal et al. 2012). It is reported from Western
Ghats of India where it was found growing on bark of tree
trunk at an altitude of 1449 m.
Remarks: Pyrenula tristissima Vain. is similar to P. minor
Fée, but the latter differs in having smaller ascospores (18
m long). This neotropical species is reported from humid
and tropical climate of North-East Himalayan region, where
it was found growing on bark of tree at an altitude of 1071 m.
Specimen examind: India, Tamil Nadu, Coimbtore district,
Siruvani waterfalls, N 10.52' E 76.41', 1449 m, on Ficus tree
trunk, H.T. Lumbsch, D.K. Upreti, P.K. Divakar & J. Tandon
19706/e/1, 19706/o/2 (LWG).
Specimen examined: India, Manipur, Tamenglong district,
Kabonram village, N 25o 07' 15.3" E 93o 37' 51.0", alt. 1071 m,
on bark, 24 September 2017, Komal K. Ingle 17-031140
(LWG)
Pyrenula tristissima Vain., Ann. Acad. Sci. fenn., Ser. A
15(no. 6): 345 (1921) Fig. 2D
Key to the Indian Pyrenula species
1.
1a.
2.
2a.
3.
3a.
4.
4a.
5.
5a.
6.
6a.
7.
7a.
8.
8a.
9.
9a.
10.
10a.
11.
11a.
12.
12a.
13.
Ascospores muriform/ or submuriform .................................................................................................... ............. 2
Aspores transversely septate .................................................................................................... ........................ 25
Ascospores submuriform .................................................................................................... ................................ 3
Ascospores muriform .................................................................................................... ...................................... 5
Ostiole apical .................................................................................................... .................................................. 4
Ostiole lateral, Ascospores 24–32 m long ......................................... Pyrenula subumbilicata (C. Knight) Aptroot
Ascospores 23–35 m long, hamathecium not inspersed .............................. Pyrenula gibberulosa (Vain.) Aptroot
Ascospores 23–33 m long, hamathecium inspersed ............ Pyrenula darjeelingensis Jagadeesh Ram & G.P. Sinha
Thallus with anthraquinone .................................................................................................... ............................. 6
Thallus without anthraquinone .................................................................................................... ........................ 7
Ascospores 10–23 m long ................................................................... Pyrenula ochraceoflava (Nyl.) R.C. Harris
Ascospores 23–35 m long ................................................................ Pyrenula ochraceoflavens (Nyl.) R.C. Harris
Ostiole lateral .................................................................................................... .................................................. 8
Ostiole apical .................................................................................................... ................................................ 11
Ascospores < 70 m long .................................................................................................... ................................ 9
Ascospores 70 m long .................................................................................................... ............................. 10
Ascosmata partly fused .................................................................................................... ................................ 11
Ascomata solitary, ascospores 15–33 m long .............................. Pyrenula microspora (Nagarkar & Patw.) Upreti
Ascospores 2 per ascus, 135–200 m long ........................................................... Pyrenula lyoni (Zahlbr.) Aptroot
Ascospores 4-8 per ascus, 50–90 m long ............................. Pyrenula ceylonensis (Ajay Singh & Upreti) Aptroot
Ascospores 25–45 m long ........................................................................... Pyrenula astroidea (Fée) R.C. Harris
Ascospores 45–70 m long ......................................................................... Pyrenula ravenelii (Tuck.) R.C. Harris
Ascospores <25 m long .................................................................................................... ............................... 13
Ascospores 25 m long .................................................................................................... .............................. 15
Thallus UV+ yellow ......................................................................................... Pyrenula confinis (Nyl.) R.C. Harris
42
Cryptogam Biodiversity and Assessment
13a.
14.
14a.
15.
15a.
16.
16a.
17.
17a.
18.
18a.
19.
19a.
20.
20a.
21.
21a.
22.
22a.
23.
23a.
24.
24a.
25.
25a.
26.
26a.
27.
27a.
28.
28a.
29.
29a.
30.
30a.
31.
31a.
Thallus UV– .................................................................................................... ................................................. 14
Ascospores 15–22 m long ........................................................... Pyrenula parvinuclea (Meyen & Flot.) Aptroot
Ascospores 7–10 m long ...................................................................... Pyrenula nanospora (Ajay Singh) Upreti
Ascospores with orange oil ....................................................................... Pyrenula breutelii (Müll. Arg.) Aptroot
Ascospores without orange oil .................................................................................................... ...................... 16
Ascospores 80 m long, mostly 2-spored ascus .............................................................................................. 17
Ascospores 80 m long, mostly 4-8 spored ascus ........................................................................................... 20
Hamathecium inspersed, Ascospores 90–200 m long .................................... Pyrenula globifera (Eschw.) Aptroot
Hamathecium not inspersed .................................................................................................... .......................... 18
Thallus not pseudocyphellate, Ascospores 80–140 m long ................... Pyrenula platystoma (Müll. Arg.) Aptroot
Thallus pseudocyphellate .................................................................................................... ............................. 19
Ascospores 80–110 m long ............................................................................ Pyrenula neosandwicensis Aptroot
Ascospores 115–180 m long ........................................................................... Pyrenula duplicans (Nyl.) Aptroot
Hamathecium inspersed .............................................................. Pyrenula sublaevigata (Patw. & Makhija) Upreti
Hamathecium not inspersed .................................................................................................... .......................... 21
Lumina angular .................................................................................................... ........ Pyrenula leucostoma Ach.
Lumina rounded .................................................................................................... ............................................ 22
Ascospores 50 m long .................................................................................................... .............................. 23
Ascospores 50 m long .................................................................................................... .............................. 24
Ascospores ends rounded ................................................................... Pyrenula pyrenuloides (Mont.) R.C. Harris
Ascospores ends pointed ............................................................................... Pyrenula papillifera (Nyl.) Aptroot
Ascospores 25–35 m long ................................................... Pyrenula welwitschii (Upreti & Ajay Singh) Aptroot
Ascospores 11–15 m long ...................................................................................... Pyrenula thelomorpha Tuck.
Ascospores 2-septate ......................................................................................... Pyrenula lineatostroma Aptroot
Ascospores 3 or more septate .................................................................................................... ....................... 26
Ascospores more than 3-septate .................................................................................................... ................... 27
Ascospores 3-septate .................................................................................................... ................................... 30
Ascospores 4-7 septate .................................................................................................... ................................. 28
Ascospores 7-11 septate ............................................................ Pyrenula subcylindrica Jagadeesh Ram & Upreti
Ascospores 30 m long .................................................................................................... .............................. 29
Ascospores 29 m long .................................................................................... Pyrenula caracasana Müll. Arg.
Ascospores with oragne oil ....................................................................... Pyrenula sexlocularis (Nyl.) Müll. Arg.
Ascospores without orange oil .............................................................................. Pyrenula wilmsiana Müll. Arg.
Ostiole lateral .................................................................................................... ................................................ 31
Ostiole apical .................................................................................................... ................................................ 34
Terminal lumina all directly against the exospore wall ......................................................................................... 32
Terminal lumina mostly separated from exospore wall .............................................................................................
Pyrenula adacta Fée
32. Ascomata at least partly fused with joint ostiole .............................................. Pyrenula subgregantula Müll. Arg.
32a. Ascomata single .................................................................................................... ........................................... 33
33. Ascospores 16–25 m long ...................................................................................... Pyrenula circumfiniens Vain.
33a. Ascospores 25–35 m long .................................................................... Pyrenula cuyabensis (Malme) R.C. Harris
34. Ascospores 50 m long ..................................................................................... Pyrenula immissa (Stirt.) Zahlbr.
34a. Ascospores 50 m long .................................................................................................... .............................. 35
43
Ingle et al.
35.
35a.
36.
36a.
37.
37a.
38.
38a.
39.
39a.
40.
40a.
41.
41a.
42.
42a.
43.
43a.
44.
44a.
45.
45a.
46.
46a.
47.
47a.
48.
48a.
49.
49a.
50.
50a.
51.
51a.
52.
52a.
53.
53a.
54.
54a.
55.
55a.
56.
56a.
Ascomata mostly aggregated .................................................................................................... ........................ 36
Ascomata solitary .................................................................................................... ......................................... 40
Hamathecium inspersed ................................................................................................... Pyrenula minarum Vain.
Hamathecium not inspersed .................................................................................................... .......................... 37
Ascospores 25 m long ........................................................................ Pyrenula zeylanica Upreti & Ajay Singh
Ascospores 25 m long .................................................................................................... .............................. 38
Ascospores 15–20 m long .................................................................................................... ........................... 39
Ascospores 20–25 m long ............................................................................... Pyrenula leucotrypa (Nyl.) Upreti
Ascospores mostly 15 m long ........................................................................... Pyrenula anomala (Ach.) Vain.
Ascospores 15 m long ...................................................................................... Pyrenula arthoniotheca Upreti
Thallus ecorticate ................................................................................................ Pyrenula microcarpa Müll. Arg.
Thallus corticate .................................................................................................... ........................................... 41
Old ascospores with orange oil .................................................................................................... ...................... 42
Old ascospores without orange oil .................................................................................................... ................. 43
Ascospores 25–35 m long ........................................................................................... Pyrenula bahiana Malme
Ascospores 35–51 m long .................................................................................... Pyrenula thailandica Aptroot
Terminal lumina all directly against the exospore wall .......................................................................................... 44
Terminal lumina mostly separated from exospore wall ......................................................................................... 54
Thallus UV+ yellow ............................................................................ Pyrenula pseudobufonia (Rehm) R.C. Harris
Thallus UV– .................................................................................................... ................................................. 45
Hamathecium inspersed .................................................................................................... ................................ 46
Hamathecium not inspersed .................................................................................................... ......................... 51
End lumina elongated .................................................................................................... .................................... 47
All lumina ± rounded to angular .................................................................................................... ..................... 48
Ascospores 20–25 m long ....................................................................................... Pyrenula maravalensis Vain.
Ascospores 30–50 m long ........................................................................... Pyrenula kurzii Ajay Singh & Upreti
Ascospores 16 m long ................................................................................... Pyrenula cayennensis Müll. Arg.
Ascospores 16 m long .................................................................................................... .............................. 49
Ascospores 40 m long ................................................................................... Pyrenula subcamptospora Upreti
Ascospores 40 m long .................................................................................................... .............................. 50
Hamathecium inspersed only in upper part. ............................................................... Pyrenula acutalis R.C. Harris
Hamathecium totally inspersed ................................................................ Pyrenula fetivica (Krempelh.) Müll. Arg.
End lumina elongated .................................................................................................... .................................... 52
All lumina ± rounded to angular .................................................................................................... ..................... 53
Ascomata 0.5–0.7 mm diam. ....................................................................................... Pyrenula finitima Müll. Arg.
Ascomata 0.5 mm diam. ................................................................ Pyrenula approximans (Krempelh.) Müll. Arg.
Ascospores 16 m long .............................................................................. Pyrenula nitidula (Bres.) R.C. Harris
Ascospores 16 m long ............................................................................. Pyrenula mastophoriza (Nyl.) Zahlbr.
Hamathecium inspersed .................................................................................................... ................................ 55
Hamathecium not inspersed .................................................................................................... .......................... 62
Central lumina strongly elongated .......................................................... Pyrenula subelliptica (Tuck.) R.C. Harris
Central lumina not strongly elongated .................................................................................................... ........... 56
Ascomata mostly 0.7 mm diam. ................................................................ Pyrenula oculata Ajay Signh & Upreti
Ascomata mostly 0.7 mm diam. .................................................................................................... ................... 57
44
Cryptogam Biodiversity and Assessment
57.
57a.
58.
58a.
59.
59a.
60.
60a.
61.
61a.
62.
62a.
63.
63a.
64.
64a.
65.
65a.
66.
66a.
67.
67a.
68.
68a.
69.
69a.
70.
70a.
71.
71a.
72.
72a.
73.
73a.
74.
74a.
75.
75a.
76.
76a.
77.
77a.
Ascospores mostly 30–45 m long .................................................................................................... ................ 58
Ascospores 30 m long .................................................................................................... .............................. 59
Ascomata conical, laterally spreading ................................................................ Pyrenula interducta (Nyl.) Zahlbr.
Ascomata globose, not spreading laterally................................................ Pyrenula mastophoroides (Nyl.) Zahlbr.
Ascospores 21 m long .................................................................................................... .............................. 60
Ascospores 20 m long .................................................................................................... .............................. 61
Ascospores 17 m long, biseriate in the ascus ......................................... Pyrenula castanea (Eschw.) Müll. Arg.
Ascospores 17–21 m long, uniseriate in the ascus .......................................... Pyrenula mamillana (Ach.) Trevis.
Ascospores rounded, uniseriate in the ascus ...................................... Pyrenula massariospora (Starb.) R.C. Harris
Ascospores at least at one end pointed, biseriate in the ascus ................................... Pyrenula acutalis R.C. Harris
Thallus UV+ yellow .................................................................................................... ....................................... 63
Thallus UV– .................................................................................................... ................................................. 64
Ascospores 24–40 m long. ........................................................................................... Pyrenula andina Aptroot
Ascospores 24 m long ............................................................................ Pyrenula dermatodes (Borrer) Schaer.
Ascospores 25 m long .................................................................................................... .............................. 65
Ascospores 25 m long .................................................................................................... .............................. 69
Ascospores 36–45 m long.. ........................................................................... Pyrenula subducta (Nyl.) Müll. Arg.
Ascospores 40 m long. .................................................................................................... ............................. 66
Ascomata 0.7 mm diam. .................................................................................................... .............................. 67
Ascomata 0.7 mm diam .................................................................................................... ............................... 68
Thallus pseudocyphellate ........................................................ Pyrenula macrospora (Degel.) Coppins & P. James
Thallus not pseudocyphellate ..................................................................... Pyrenula complanata (Mont.) Trevis.
Ascospores mostly 32–42 m long. .................................................................... Pyrenula punctella (Nyl.) Trevis.
Ascospores mostly 25–37 m long.. ................................................................... Pyrenula quassiaecola (Fée) Fée
Ascospores 21–25 m long .................................................................................................... ........................... 70
Ascospores 21 m long .................................................................................................... .............................. 71
Ascomata K+ red ................................................................................................... Pyrenula nitida (Weigel) Ach.
Ascomata K– ................................................................................. Pyrenula submastophora Ajay Singh & Upreti
Ascospores mostly 15 m long .................................................................................................... ................... 72
Ascospores 15 m long .................................................................................................... .............................. 74
Ascomata 0.7 mm diam ..................................................................... Pyrenula atropurpurea (Eschw.) Müll. Arg.
Ascomata 0.7 mm diam .................................................................................................... ............................... 73
Ascospores 6–8 m long .................................................................................................... Pyrenula brunnea Fée
Ascospores 4–6 m long ........................................................................................ Pyrenula aspistea (Ach.) Ach.
Ascomata 0.7 mm diam .................................................................................................... ............................... 75
Ascomata 0.7 mm diam. .................................................................................................... .............................. 76
Lumina rounded .................................................................................................... Pyrenula scutata (Stirt.) Zahlbr.
Lumina angular .......................................................................................... Pyrenula balia (Krempelh.) R.C. Harris
Ascomata 0.4 mm diam. .................................................................................................... .............................. 77
Ascomata 0.4 mm diam ........................................................................................ Pyrenula aggregata (Fée) Fée
Ascospores 18–20 m long ........................................................................................... Pyrenula tristissima Vain.
Ascospores 18 m long. .................................................................................................... .. Pyrenula minor Fée
45
Ingle et al.
ACKNOWLEDGEMENTS
new species of Pyrenula from South America, with a note on
overmature ascospores. Lichenologist 45: 169–198. https://
doi.org/10.1017/S0024282912000734
We are grateful to the Director, National Botanical
Research Institute, Lucknow for proving laboratory facilities
and authorities of Forest Departments of Manipur and
Nagaland for allowing collection in the area. Financial
support from IBSD, Imphal and DBT, Government of India
is gratefully acknowledged. Authors are grateful to Dr. D.
K. Upreti for valuable guidence in identification of the
specimens and preparation of the manuscript.
Jagadeesh Ram TAM and Sinha GP (2010). A new species and
new records of Pyrenula (Pyrenulaceae) from India.
Lichenologist 42(1): 51–53. https://doi.org/10.1017/
S0024282909990132
Mendonça CO, Aptroot A and Cáceres MES (2016). Six new
species of Pyrenula (Pyrenulaceae) from Northeast Brazil.
Phytotaxa 286 (3): 169–176. http://dx.doi.org/10.11646/
phytotaxa.286.3.4
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DR. SANJEEVA NAYAKA