American Journal of Plant Sciences, 2011, 2, 554-560
doi:10.4236/ajps.2011.24066 Published Online October 2011 (
Copyright © 2011 SciRes. AJPS
Conocephalum conicum (L.) Dumort. (Snake
Liverwort) Threatened in Bhaderwah (J & K) due
to Environmental Shock
Mudassar Iqbal1, Anima Langer1, Afroz Alam2
1Department of Botany University of Jammu, Jammu, India; 2Department of Bioscience and Biotechnology, Banasthali University,
Rajasthan, India.
Received December 28th, 2010; revised May 13rd, 2011; accepted June 6th, 2011.
Paper includes information on population status of Conocephalum conicum collected from diverse habitats in Bhad er-
wah (J & K state). Taxon exhibits tremendous diversity in morphoanatomical details of various gametophytic charac-
ters. Isozyme analysis was performed to find out whether variability in the taxon is only environmental or has genetic
basis as well.
Keywords: Bryophyta, Liverwort, Marchantiales, Conocephalum, Jammu & Ka shmir
1. Introduction
Bryophytes constitute one of the largest groups of land
plants among the plant kingdom. Himalayas rank first at
national [1] and third at global level in bryo-diversity [2].
During the last few decades, however, Himalayas have
suffered tremendously due to various tourism and ur-
banization related anthropogenic activities leading to the
disappearance of many of their inhabitants including
bryophytes. On basis of an observations [3] on occur-
rence of several hepatics in Western Himalaya (Nainital)
over the years, 9 threatened taxa [Athalamia pinguis,
Dumortiera hirsuta, Cryptomitrium himalayensis, Re-
boulia hemispherica, Stephensoniella brevipedunculata,
Weisnerella denudata, Fossombronia himalayensis,
Sewardiella tuberifera and Conocephalum conicum]
which were had observed to grow luxuriantly in the yes-
teryears, but had disappeared or undergone extreme re-
duction in population size, from the area. The suggested
specific reasons include anthropogenic activities like
rapid urbanization, land clearance, road construction,
deforestation, indiscriminate and wreck less collection
etc., as factors which could have contributed to the habi-
tat destruction and consequently, the disappearance of
these taxa. She, therefore, voice was raised for their con-
Against this backdrop, work on intraspecific variabil-
ity was initiated on three, Reboulia hemispherica [4],
Dumortiera hirsuta [5] and Conocephalum conicum [6]
of the nine hepatic taxa enlisted by Pant [1] in Depart-
ment of Botany, at University of Jammu, Jammu. Present
communication deals with our observations on popula-
tion status of Conocephalum conicum collected from
Bhaderwah in Jammu region, one of the areas of the
North-west Himalaya which used to host rich bryo-di-
versity in the yesteryears, because of the undisturbed,
stable habitats due to the inaccessibility of the area.
Bhaderwah lies between 32˚08 N - 35˚52 N latitude
and 75˚48 E - 75˚32 E longitude having an altitudinal
range of 821 - 4341 m. The climate of Bhaderwah is tem-
perate; temperature varies from minimum mean 17.3˚C
to maximum mean 29.9˚C in summers and minimum
mean –0.9˚C to maximum mean 02.8˚C in winters. It
experiences severe cold during winters with snowy sea-
son between the months of December to March with
temperature remaining subzero. The snow melts off by
April and weather becomes pleasant and remains so up to
May. In June and July, temperature increases upto the
maximum 29.9˚C. During August and September, the
temperature subsides and it again becomes cold after-
wards. The annual rainfall is up to 249.6 mm and maxi-
mum humidity is 84% [7].
In recent years, Bhaderwah has been subjected to tre-
mendous developmental and tourism related pressures,
like construction of bridges, highways, parks, roads,
Conocephalum conicum (L.) Dumort. (Snake Liverwort) Threatened in Bhaderwah (J & K) 555
due to Environmental Shock
tourism huts, deforestation, destruction of rocks by quar-
rying etc. Furthermore, a University Campus has recently
being established at Bhaderwah over a vast area. These
activities are likely to lead to habitat destruction and de-
pletion/disappearance of many of the inhabitants of the
area. Under such circumstances, only those organisms
which have accumulated genetic variability are likely to
survive, whereas genetically homogenous taxa are des-
tined to be wiped out by the natural selection. Cono-
cephalum conicum, a species reported to have disappeared
elsewhere in Western Himalaya in response to similar
reasons, was therefore selected for the present study.
2. Materials and Methods
Conocephalum conicum (L.) Dumort. is commonly
called snake liverwort is the study material collected
from Bhaderwah, Jammu & Kashmir, India.
3. Results and Discussion
Conocephalum conicum is commonly called snake liv-
erwort due to hexagonal areolae on its dorsal surface
(Figure 1(a)). It is also called cone headed liverwort
because of its conical archegoniophores (Figure 1(b)). It
is worldwide in distribution, as it is known to be distrib-
uted in America, Algeria, Alaska, Azores, Bhutan, Cau-
casus, Canary, China, Europe, Florida, India, Ireland,
Italy, Japan, Korea, Kuriles, Liukium Maderia, Nepal,
Pakistan, Rubki-Neylong, Seghalim and Siberia [8].
From India, the taxon is reported from Eastern [9-11] and
Western Himalaya [12-16].
In Jammu and Kashmir (North West Himalaya), it has
so far been collected from Poonch [17], Udhampur and
Kathua as stated by [18] districts of Jammu region and
Ladakh [19].
During present investigations, 46 accessions of C.
conicum have been collected growing at different sites
ranging in altitude from 1230 m - 2600 m. Populations
were collected growing on substrates with pH ranging
between 6.5 - 6.9. Plants have been collected from di-
verse habitats, such as epilithic (rock crevices, caves;
Figures 1(c) and (d)), under dripping chilled water (Fig-
ure 1(e)), under snow cover (Figure 1(f)), stream banks
(Figure 1(g)), submerged under water (Figure 1(h)),
non-epilithic (cool and moist shady soil, loamy soil)
(Figure 1(i)), and epiphytic on decaying log of Cedrus
deodara (Figure 1(j)).
Thalli of various populations displayed variability in
morpho-anatomy of almost every gametophytic character
(thallus size, rhizoids, scales, air pores, vegetative anat-
omy etc.) studied, that has been communicated separately.
In this paper focus is mainly on reproductive biology of
the taxon.
Figure 1. (a) A part of Thallus showing hexagonal areolae
on its dorral surface. (b) Thallus of C. conicum showing
conical archegoniophore. (c) & (d) Various habitats from
where populations of Conocephalum conicum have been
collected. (e) Under dripping chilled water. (f) Under snow
cover. (g) Stream banks. Submerged under water.
Non-epilithic. (h) Epiphytic. (i) On rotten log of Cedrus
C. conicum reproduces both, asexually as well as sexu-
ally; asexual reproduction occurs usually by regeneration
from thallus fragments; thallus being light in weight, is
easily transported by water; thallus produces ventral in-
novations, both apical and lateral (Figure 2(a)). During
Copyright © 2011 SciRes. AJPS
Conocephalum conicum (L.) Dumort. (Snake Liverwort) Threatened in Bhaderwah (J & K)
due to Environmental Shock
Figure 2. (a) Thalli of Conocephalum conium showing apical
and lateral innovations. (b) & (c) Male plants with 2 - 3
receptacles. (d) Female thalli with young (Note upto 4 re-
ceptacles/thallus). (e) & (f) Mature receptacles (1 in Figure
2(e) and 2 in Figure 2(f)). (g) Female thallus with healthy
receptacle and stalk. (h) Female thalli of Acc. MI 37 and MI
38 showing unhealthy/shrivelled re ceplacles and stalk.
present investigation, only 15 accessions were recorded
in fertile form, of which 12 comprised both male and
female thalli growing together, while remaining three
had only female plants. Dioecious; male receptacles ses-
sile, green, disciform, disc slightly raised above main
thallus, surrounded by thin growth of thallus; terminal or
subterminal; receptacles oval or circular having con-
spicuous papillae on their upper surface; borne on main
thallus as well as on apical innovations; number of re-
ceptacles per thallus varied from 2 - 3 (Figures 2(b) and
(c)); initiation of male receptacles occurred in April, and
they disappeared in July.
Although female thalli were recorded presently in 15
accessions, mature receptacles could be observed only
from 3; in the remaining 12 accessions, receptacles were
observed at young stages only, as they degenerated be-
fore reaching maturity. Female receptacle terminal or
slightly sub-terminal, 1 to 4 per thallus, sunken when
young in a cup formed by dorsal growth of the thallus
(Figure 2(d)); receptacles appeared in first week of May;
one flask shaped archegonium present per lobe with
broad venter and long neck; post fertilization changes
recorded after mid June, when neck of archegonium de-
generated; receptacles started emerging out of sunken
cup, stalk elongated slowly till it reached maximum of 11
cm (Figures 2(e)-(g)) in March next year. Mature recep-
tacles conical; raised on long stalk.
Receptacles were symmetric, healthy and 5 - 8 lobed
in single accession (Acc. MI 46; Figures 2(e)-(g)) and
unhealthy and asymmetric up to 5 lobed in two (Acc. MI
37 and MI 38; Figures 2(h)-3(d)). Stalk stout and
healthy in Acc. MI 46 (Figures 2(e)-(g)) but unhealthy
and shriveled in Acc. MI 37 and MI 38 (Figures
2(h)-3(d)). Mature sporophytes were present only in 1
and/or 2 lobes per receptacle in Acc. MI 37 and MI 38;
remaining lobes being extremely reduced, whereas in
Acc. MI 46 mature sporophytes were recorded in a
maximum of 5 lobes out of 8 (Figures 3(e)-(h)). One
sporophyte per lobe, having foot, reduced seta and pyri-
form capsule (Figure 3(i)). Sporophytes excised from
developed lobes of unhealthy receptacles exhibited re-
duced capsules (Figure 3(j)).
Mature spores observed only in Acc. MI 46 (Figure
4(a)); capsules of Acc. MI 37 and MI 38 possessed very
few viable spores (Figure 4(b); Table 1). Capsules from
even the fully mature receptacles of these unhealthy
populations possessed developing sporocytes and elaters;
ratio between the two being around 0.4:1. Capsules
showed very low spore output as compared to that of
elaters. Like the other two populations spore-elater ratio
in the healthy population (Acc. no. MI 46) was around
0.4:1 (Table 3). Out of the spores formed in Acc. MI 46,
another 10% were non viable. These were small (16 µm -
20 µm) × (12 µm - 16 µm), triangular and shriveled as
compared to viable ones which were large (44.5 µm -
102.3 µm) × (25 µm - 55.4 µm), green and healthy (Ta-
ble 2).
Elaters brown, healthy, bi- and tri- spiral, unbranched
and bluntly fusiform in Acc. MI 46 (Figure 4(c); Table
2). Elaters of abnormal morphology (branched and
blackish) were also present in capsules of Acc. MI 37
Copyright © 2011 SciRes. AJPS
Conocephalum conicum (L.) Dumort. (Snake Liverwort) Threatened in Bhaderwah (J & K) 557
due to Environmental Shock
Figure 3. (a) A female thallus of Acc. no MI 37 showing
asymmetric receptacles and stalk. (b-d) Female thallus of
Acc. no MI 37 and MI 38 with unhealthy/shrivelled stalk.
(e-h) Female thalli of Acc. no MI 46 bearing receplacles
with 1 (e), 2 (f), 4 (g) and 5 (h) sporophytes. (i) & (j) Healthy
(i) and unhealthy (j) Sporophyte exised from receptacle
lobes of MI 46 (i) and M 37 (j).
and MI 38 (Figure 4(d); Table 3). Size of elaters var-
Figures 4. (a) A few viable and non-viable spores excised from
matures capsule of healthy population. (b) Developing sporo-
cytes and claters, excised from capsule of unhealthy population
(Note the abundance of elaters). (c) Whole mount of healthy
elaters. (d) Whole moun t of abnormal elaters.
ied between (50 µm - 150 µm) × (15 µm - 70 µm)
(Table 3). Occurrence of elaters is a characteristic
unique to the liverworts. In contrast, they are totally
lacking in mosses. Archesporium throughout the class
hepaticopsida is endothecial in origin and differentiates
into sporocytes and elaterocytes; former undergo reduc-
tional division, each giving rise to 4 spores and the later
differentiating into elaters. Since both, sporocytes and
elaterocytes belong to the same generation, basic ratio
between spores and elaters expected is 4:1. This ratio
exists in a number of Marchantialean taxa like Reboulia,
Mannia, Targionia, Conocephalum etc. During the
course of evolution there has been a trend towards the
increase in spore output and consequently, higher
spore-elater ratios. Marchantia polymorpha displays
highest ratio (128:1) among Marchantiales. Schistochila
(Jungermanniales) exhibits still higher (200:1)
spore-elater ratio (Table 2). Sporocytes of all such taxa
where spore-elater ratio is higher than 4:1, undergo 1 or
more mitotic divisions, prior to meiosis. Interestingly,
these divisions have been accompanied by the reduction
in spore size. Production of large number of small sized
spores is an adaptation for wind dispersal of spores and
subsequently, successful under terrestrial habitat. Al-
though Conocephalum, Reboulia, Mannia and Tar-
gionia are theoretically expected to show 4:1 ratio, actual
atios are sometimes lower. Bischler (1998) re- r
Copyright © 2011 SciRes. AJPS
Conocephalum conicum (L.) Dumort. (Snake Liverwort) Threatened in Bhaderwah (J & K)
due to Environmental Shock
Copyright © 2011 SciRes. AJPS
Table 1. Number of viable/non-viable spores in receptacles with different sporophytes.
Size (µm)
Oval Circular
Number of Viable/Non-Viable Spores in Receptacles with Different Sporophytes
Acc. No.
L B diameter 1 2 3 4 5 6 7 8
MI 37 42.5 - 115
22 - 52
22.5 - 52.5
2848 - 2861
5344 - 5675
MI 38 42 - 110
25.5 - 49
22 - 50
2789 - 2915
5125 - 5478
Table 2. Variation recorded in spore characters.
Size (µm)
Oval Circular
Number of Viable/Non-Viable Spores in Receptacles with Different Sporophytes
Acc. No.
L B diameter 1 2 3 4 5 6 7 8
MI 37 42.5 - 115
22 - 52
22.5 - 52.5
2848 - 2861
5344 - 5675
MI 38 42 - 110
25.5 - 49
22 - 50
2789 - 2915
5125 - 5478
M1 46
44.5 - 120.3
16 - 20
25 - 25.4
12 - 16
23.2 - 55.3
2485 - 2725
50 - 110
5000 - 5350
150 - 225
7787 - 7985
315 - 375
9704 - 10492
400 - 425
12500 - 13505
450 - 480
Table 3. Variation recorded in elaters’ output.
Size (µm) Number of elaters in receptacle with different sporophytes No. of
spirals Spore/Elater ratio
L B 1 2 3 4 5 67823 4 1 2 3 4 5
MI 37 45 - 105
15 - 75
6106 - 6504
12212 - 12895
(12710.3) –––+– + 0.42:1 0.43:1 ――
MI 38 43 - 350
12 - 65
5895 - 5990
12427 - 12895
(12610.1) –––+– + 0.47:1 0.42:1 ――
MI 46 50 - 350
15 - 70
6335 - 6660
12317 - 12910
18318 - 18602
24920 - 25689
30982 - 31670
(31525.5) –––++ – 0.39:1 0.40:1 0.42:1 0.40:10.41:1
corded a similar situation in C. conicum and Pellia sp.,
and attributed it to endosporic, precocious spore germi-
nation in the species. Although in these species ratio was
lower than 4:1 yet the number of spores was far more
than elaters. Data obtained presently are in total contrast
to the expected numbers. Here, elaters were more abun-
dant than spores, leading to the ratio between spores and
elaters as low as 0.4:1 (Table 2). This is attributable to
the fact that a large number of sporocytes do not undergo
reduction division and therefore, the number of spores
does not multiply. This clearly indicates that although
apparently, C. conicum seems to be well established un-
der various habitats in the study area, all the populations
are asexually propagated only. Although female recepta-
cles were formed in 15 accessions, they reached maturity
only in 3 and out of these too, a few viable spores are
formed only in one.
4. Population Status
It can be inferred from the data obtained for various re-
productive characters (female gametangia, sporocytes
and spore-elater ratios) that there is minimal (if any)
contribution of sexual reproduction to the variability re-
corded among various populations. In order to further
assess the nature of variability, five populations of C.
conicum collected from diverse habitats i.e., epilithic
(under dripping chilled water, under snow cover), non
epilithic (moist shady soil, sandy soil) and epiphytic
(Cedrus deodara) were subjected to Polyacrylamide gel
electrophoresis for three enzyme systems (peroxidases,
estrases and phosphatases). The zymograms, thus ob-
tained revealed homogeneity in the occurrence of bands.
A total of 6 bands were obtained for phosphatases (Fig-
ures 5(a)-(d)), 5 for esterases (Figures 5(e)-(f)), one
Conocephalum conicum (L.) Dumort. (Snake Liverwort) Threatened in Bhaderwah (J & K) 559
due to Environmental Shock
Figures 5. Pictures of gels subjected to PAGE and explana-
tory figures thereof; Peroxidases (a-d), esterases (e,f) and
phosphatase (g,h).
for peroxidases (Figures 5(g) and (h)) in all the popula-
tions analyzed. Results thus obtained confirm our pre-
sumption that the variability recorded in the species is
environmental and lacks any genetic basis.
Geissler et al. [20] laid down following five criteria to
be fulfilled by a taxon before being termed threatened. a)
It should be threatened worldwide, b) should be confined
to a threatened habitat, c) should have a narrow range, d)
should not be overlooked and/or over collected and e)
should have a unique morphology/biology or occupy a
special evolutionary position.
5. Conclusions
Although C. conicum does not seem to satisfy criteria 1 -
4, populations studied exhibit unique reproductive biol-
ogy. It can, therefore, be concluded that the species is
threatened in Bhaderwah, as it has earlier been reported
for Kumaon Himalaya.
6. Acknowledgements
The authors are thankful to the Head, Department of
Botany, University of Jammu, Jammu and Vice Chen-
cellor, Banasthali Vidyapith, Rajasthan for providing ne-
cessary library and laboratory facilities.
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due to Environmental Shock
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