Submitted Ph.D. thesis to Dr. R.M.L. Avadh University, in Botany Discipline. The research was carried out under the supervision of Dr. D.K. Upreti, Scientist, NBRI, Lucknow. Topic of the thesis is “Revisionary studies of Lecanora s. lat. from India”.
Sumary:
During the last five decades of past century a large number of macrolichens (foliose and fruticose) from India were revised or monographed. At present information on the occurrence of 697 species of macrolichens (crustose, suamulose, placodioid) of India belonging to 71 families is available. On the other hand, in contrast to macrolichens, microlichens from India are insufficiently revised or even investigated. Often descriptions from the century back publications were only information for several taxa. Most of such taxa belong to the lichen families Lecanoraceae and Lecideaceae. Awasthi in keys of microlichens of India has not keyed out Lecanora species, as the data according to the present concept of the genus were not available. The information on Indian Lecanora was scattered and a large number of specimens of the genus were lying unidentified in the lichen herbarium of National Botanical Research Institute, Lucknow (LWG). Thus there was an urgent need for the revision of the genus and consolidation of the information available so far on the species of Lecanora from India.
The lichen genus Lecanora Ach., is characterized by the crustose to effigurate or squamulose thallus containing Trebouxioid phycobiont, lecanorine apothecia and colourless simple ascospores which are small to medium sized. Lecanora is one of the largest genera with 600 species in the world, out of which 76 species are represented in India. It is a heterogeneous genus and widely distributed from Arctic, Antarctic to subtropical regions.
Lecanora
sensu stricto includes
species with crystals in the amphithecium, containing atranorin, filiform
conidia and with or without algal cell in amphithecium. Lecanora sensu lato
includes Lecanora sensu stricto and
all the other species under the genus Lecanora.
Lichen
genus Lecanora has two subgenus, Lecanora
subgenus Lecanora with purely crustose forms and Lecanora subgenus Placodium with
crustose to squamulose or placodioid forms.
According to the modern concept of lichen classification, the lichen genus Lecanora
belongs to family Lecanoraceae, suborder Lecanorineae, order Lecanorales, group
Bitunicate Ascohymenials, subclass Euascomycitidae and class Ascomycetes.
The
revisionary studies on lichen genus Lecanora
sensu lato in India is based on the observation of more than 1500 specimens
housed at LWG. The lichen herbarium LWG is richest in the South East Asia,
holding more than one lakh specimens belonging to about 1500 species and
representing all the phytogeographic regions of India. It includes personal
herbarium of Dr. D.D. Awasthi (AWAS), lichen herbarium of Lucknow University (LWU)
which have been transferred to LWG on permanent loan basis, thousands of
exsiccate materials and also several voucher specimens deposited by various
workers. The fresh collection for the present study of Lecanora
were also made from Himachal Pradesh, Sikkim, Uttaranchal and few localities of
Karnataka, Tamil Nadu and Uttar Pradesh.
The
taxa were investigated following the modern concept of the genus. All the
available information on the genus Lecanora
from India is consolidated. The Lecanora
specimens were identified by studying their morphology, anatomy and
chemistry. The chemistry of the specimens was performed by colour spot tests
(with usual reagents K, C, PD), thin layer chromatography (TLC) and high
performance liquid chromatography (HPLC) for few specimens. All the taxonomical
characters needed for the identification of species are discussed.
The
identity of the specimens were confirmed by matching them with type specimens,
exsiccate materials housed at LWG or consulting experts (Lumbsch, T.H.,
University of Essen, Germany).
The
description and discussion of different species are given. The list of different
chemical compounds has also been provided. The key to segregate different
species is given. The taxonomic treatment includes full citation of the
literature, basionym, synonyms, type collection and taxonomic description
(including Latin diagnosis and etymology in case of new species, morphology of
thallus and apothecia, anatomy of apothecia, chemistry, habitat, altitude,
distribution and list of specimens examined). Each species is compared with
similar taxa and the difference is discussed.
The
revisionary studies on lichen genus Lecanora
sensu lato in India revealed the occurrence of 76 species from India. A
total of five species are described as new to science, 20 are new records while
37 taxa are excluded for the country and the reason for the exclusion is
discussed,
New species:
1.
Lecanora caesiousa S. Nayaka and D. Upreti
2.
L. leuteomarginata S. Nayaka and D. Upreti
3.
L.
palniensis S. Nayaka and D. Upreti
4.
L.
sorediferous S. Nayaka and D. Upreti and
5.
L.
subpraesistens S. Nayaka and D. Upreti
New Records:
1. |
Lecanora
achroa Nyl. in J.M. Crombie |
11. |
L. kirra Poelt and Grube |
2. |
L. alba Lumbsch |
12. |
L. leproplaca Zahlbr. |
3. |
L. andina Räsänen |
13. |
L. marginata (Schaer.) Hertel and Rambold |
4. |
L. austrointumescens Lumbsch |
14. |
L. melanommata C. Knight |
5. |
L. flavidofusca Müll.Arg. |
15. |
L. polytropa (Ehrh.) Rabenh. |
6. |
L. formosula Lumbsch |
16. |
L. pseudargentata Lumbsch |
7. |
L. fuscococcinea Nyl. |
17. |
L. queenslandica C. Knight |
8. |
L. helva Stizenb. |
18. |
L. somervellii Paulson |
9. |
L. imshaugii Brodo |
19. |
L. subalbellina Vainio |
10. |
L. interjecta
Müll.Arg. |
20. |
L. wilsonii Müll.Arg. |
Excluded species:
1.
|
Lecanora
achroella Nyl. |
- distributed in Sri Lanka |
2.
|
L. adolfii Wei |
- distributed in Khumbu Himal, Nepal |
3.
|
L. amorpha Poelt |
- distributed in Khumbu Himal, East Nepal |
4.
|
L. badia (Pers.)
Ach. |
- transferred
to Protoparmelia badia (Hoffm.)
Half., distributed in N.W. Himalayas |
5.
|
L.
baicalensis Zahlbr. |
- distributed in Xinjiang Prov., China |
6.
|
L. carnulenta
Nyl. |
- reported from
E. Himalayas, but existence is doubtful |
7.
|
L. ceylanica Zahlbr. |
- distributed in Sri Lanka |
8.
|
L. collaevans
Nyl. |
- distributed in Sir Lanka |
9.
|
L. demissa (Flotow) Zahlbr. |
- distributed
in Jangri, Langtang and Khumbukama Himal, Nepal |
10.
|
L.
dispersoareolata (Schaer.) Lamy |
- distributed in Karakorum, Pakistan |
11.
|
L. dissipans Nyl. |
- distributed in Sri Lanka |
12.
|
L. emergens Müll.
Arg. |
- reported from
Manipur, but existence is doubtful |
13.
|
L. emodii H.
Vänska |
- distributed in Khumbu Himal, Nepal |
14.
|
L. freyi Poelt |
- distributed in Karakorum, Pakistan |
15.
|
L. furfurosa Tuck. |
- distributed in Sri Lanka |
16.
|
L. fusca Müll.
Arg. |
- reported from
Uttar Pradesh, but existence
is doubtful |
17.
|
L. glabrata (Ach.)
Malme |
- reported from
Madhya Pradesh, but miss identified for L. pulicaris |
18.
|
L.
laatakkaensis (Räsänen) Poelt |
- distributed in N.W. Himalayas of Pakistan |
19.
|
L.
lecanactina Zahlbr. |
- reported from
Madhya Pradesh and Uttar Pradesh, but miss identified for L.
tropica Zahlbr. |
20.
|
L. lesleyana Darb. |
- distributed in Mt. Everest, Nepal |
21.
|
L.
melacarpella Müll. Arg. |
- reported from
Andra Pradesh, but miss identified for L. fuscococcinea Nyl. |
22.
|
L. melanaspis
Ach. |
- reported from
N.W. Himalayas, but existence is doubtful |
23.
|
L.
pachyphylla H.
Magn. |
- distributed in Xinjiang and Kansu, Chaina |
24.
|
L.
pachyrihiza Poelt
and Grube |
- distributed in Karakorum, Pakistan |
25.
|
L. piniperda Körb. |
- now L.
albellula (Nyl.) Motyka, reported from Uttar Pradesh, occurrence in
India doubtful (personal communication with Dr. T.H. Lumbsch). |
26.
|
L.
placentiformis Steiner |
- distributed in Karakorum, Pakistan |
27.
|
L. populicola
(DC.
In Lam.) DC. |
- a temperate
to boreal species, occurrence in India is doubtful (personal communication
with Dr. T.H. Lumbsch). |
28.
|
L. scrupulosa
Ach. |
- reported from
N.W. Himalayas, but existence is doubtful |
29.
|
L. sherparum Poelt |
- distributed in Khumbu Himal, Nepal |
30.
|
L. sorediella
Nyl. |
- distributed in Sri Lanka |
31.
|
L. subcarnea (Lilj.)
Ach. |
- reported from
Madhya Pradesh, but miss identified for L. sulphurescens |
32.
|
L. subeffusa (L.)
Ach. |
- distributed
in Sri Lanka an E. Himalayas but occurrence is doubtful |
33.
|
L.
teretiuscula Zahlbr. |
- distributed in Langtang an Khumbakama Himal,
Nepal |
34.
|
L.
tschomolongamae Poelt |
- distributed in Langtang, Mahalangur Himal, Nepal |
35.
|
L.
verruculigera Hue |
- reported from
Uttar Pradesh, but existence is doubtful |
36.
|
L. warmingii Müll.
Arg. |
- transferred
to Vainiora warmingii (Müll.
Arg.) Kalb., reported from Jammu and Kashmir, and Maharashtra, occurrence
is doubtful |
37.
|
L. xylophylla
Hue |
- reported from
Goa, Jammu and Kashmir, Madhya Pradesh, Tamil Nadu and Uttaranchal, but
miss identified for some Lecideoid member. |
Out
of 76 species of Lecanora sensu lato 12
belongs to Lecanora subgenus Placodium while remaining 64 belongs to subgenus Lecanora.
Within the subgenus Lecanora the subfusca group is dominating with 57 species, followed by varia
group, represented by L. intricata, L.
polytropa, L. saligna, L. sambuci and L.
varia while dispersa group has
only two species such as L. albescens and
L. dispersa. The subgroups of subfusca
group, coronulans, pallida and marginata
are represented by eight, seven and one species respectively while remaining
41 species belongs to subgroup subfusca.
The
Western Himalayas exhibit the maximum diversity of Lecanora
with 66 species among the eight lichenogeogrphical regions of India. The
Western Ghats, Eastern Himalayas and Central India with 31, 20 and 18 species
respectively are the regions having moderate number of Lecanora
species. The Gangatic Plain, Eastern Ghats
and Deccan Plateau, Western Dry region and Andaman Nicobar Islands
exhibit scarce to poor diversity of Lecanora
with 10, nine, six and two species respectively.
Among
the different states Uttaranchal scores highest of 50 species followed by
Himachal Pradesh with 42 species and Tamil Nadu by 25 species.
The
taxa of Lecanora are substrate
specific. A total of 45 species found growing on bark (corticolous), 27 on rock,
boulders, stones (saxicolous) while L.
chondroderma, L. gypsacea and L.
himalayae were found either on moss (muscicolous) or soil (terricolous). L.
dispersa has a wide niche and grows on nutrient rich soil, bark and soil.
The type of the substratum is one of the important criteria for segregation of
different groups and taxa in Lecanora. The
name of the host plant is not mentioned on the herbarium packets thus it gives
very less scope to discuss the host specificity of different
taxa. However, by field experience and observation of the samples of bark in
the herbarium it is clear that Lecanora prefers
to grow on rough bark. The rough bark helps in retaining the moisture for longer
duration, easily trapping the lichen spores and provides better niche for
establishment. The major host plants in the Gangatic plain and Central India are
Mangifera indica and Shorea
robusta, in coastal areas the coconut trees, in Eastern Himalayas Cryptomeria
and Michelia trees, and in Western
Himalayas Quercus, Pinus and Cedrus
trees are mostly the rough barked trees. The Western Ghats has a great
diversity of host plants and their names are rarely mentioned on the herbarium
packets.
The
altitude plays an important role in the distribution of Lecanora.
The maximum number of Lecanora taxa
prefers to grow in temperate regions and the most favourable altitudinal range
is between 2001-2500 m. L. allophana, L.
cinereofusca, L. cinereofusca var.
himalayensis, L. indica, L. muralis var.
dubyi, L. muralis var. muralis, L.
perplexa, L. queenslandica, L. subimmersa and L.
sulphurescens are the taxa having a wide altitudinal range (Table 3). Among
the 12 placodioid Lecanora, 11 exhibit
their restricted distribution in temperate regions of Himalayas while only L.
indica distributed in other parts of the country. Most of placodioid Lecanora
prefers to grow in sub-alpine to alpine regions or in cold desert and mostly
on rock. The species L. andina, L.
fuscococcinea and L. intumescens are
mainly tropical in distribution. The availability of wide range of altitude
along with substratum is few of the reasons for high diversity of Lecanora
in Western Himalayas, Western Ghats and Eastern Himalayas.
To study the distribution of Indian Lecanora
vis-à-vis the world five distribution patterns can be recognized (Table 3);
1. cosmopolitan (C) - with world wide distribution, 2. restricted (R) –
restricted in distribution to one major region, such as Australia, Africa,
Europe, North or South America, northern or southern hemisphere, 3. semi-
restricted (SR) – distributed in mainly in one or two adjacent regions, 4.
endemic (E) – confined to India, and 5. new species (N) – presently known
from India. L. campestris ssp. gulmargia,
L. cinereofusca var. himalayensis, L.
fimbriatula and L. indica are
endemic to India while five new species described are presently known only form
their type locality except L. palniensis which
is also known from 2-3 localities in south India only. A total of 19 taxa were
cosmopolitan in distribution, 29 are restricted in distribution while 18 were
semi restricted in their distribution.
Lecanora is always found growing in association with other lichens and
bryophytes. However, strict association between any species and group could not
been established. The associated species varies from habitat to habitat. In
tropical regions the corticolous Lecanora
are found growing with the members of Physciaceae (Physcia, Heterodermia, Phaeophyscia), Graphidaceae and rarely with
Parmeliaceae. In moist, humid and shady areas the bryophytes (leafy liverwarts
and moss), Collema, Leptogium are the
common associates. In drier areas Arthonia,
Buellia, Chrysothrix accompany Lecanora.
In the evergreen forests of Western Ghats Graphidaceous, Pyrenocarpous and
Thelotremataceous lichens grow along with Lecanora.
The temperate climate such as Western Himalayas supports the good growth of
Parmelioid lichens (Bulbothrix,
Everniastrum, Hypotrachyna, Parmotrema, Punctelia, Rimelia, Relicina) that
usually grow along with Lecanora. The
species of Usnea, Ramalina are
occasionally found growing with Lecanora.
In temperate and sub alpine region the twigs of Barberis, Pyracantha crenulata bushes hosts the luxuriantly growth Buellia,
Caloplaca, Heterodermia, Ramalina, Usnea and Parmelioid lichens along with Lecanora.
On the cultivated trees species of Lecanora
usually grows along with Candelaria,
Heterodermia, Punctelia and Physcia.
The
saxicolous species of Lecanora on
open, exposed rocks grows along with species of Buellia,
Lecidea, Staurothele, Verrucaria and
Caloplaca while the shady rocks supports
Collema, Heterodermia, Leptogium, Physcia and Parmelioid lichens. The rocks
are the major substratum for the lichens in sub alpine and alpine regions. All
most all the placodioid Lecanora are
found growing in this climate. The different species of Lecanora grow together along with the species of Aspicilia,
Acarospora, Caloplaca, Rhizocarpon, Rhizoplaca, Staurothele, Xanthoria,
Xanthoparmelia and Verrucaria
It is too early to predict the affinities of Indian Lecanora
with other phytogeographic regions of the world, since not many revisionary
studies on the genus are available throughout the world. The different
lichenogeographical regions of the India such as Western Ghats, Western
Himalayas and Eastern Himalayas does not show any specific affinities with the
adjacent or neighbouring phytogeographic regions of the world. However, Lecanora
of India in general shows its affinity towards the Australian, European,
North American and partly Eastern Asian elements. The Lecanora
taxa of Western Himalayan regions have their affinity towards those of
Northern Hemisphere including Europe, Canada and Arctic.
L. argentata collected on 1st Dec. 1947
from Port Caming, West Bengal by D.D. Awasthi is the oldest herbarium that
have been observed during the study and it has been repeatedly collected
thereafter. Except for L. albescens, L. chondroderma, L. dispersa, L. intricata, L. sambuci and L. varia, all the other taxa are collected repeatedly or collected
in recent times. Since most these species are widespread and cosmopolitan in
distribution and can not be considered as extinct from India. They could not
been recollected since the exact locality and habitat of the species were
untraceable. None of the Lecanora taxa
falls under any IUCN category.