RESEARCH ARTICLE
Variation in Adaptive Traits of an Endemic
Meconopsis napaulensis DC. along an
Elevation Gradient in Alpine Himalaya,
Central Nepal
Narmada Rana, Suresh Kumar Ghimire
Tribhuvan University, Kathmandu, Nepal
*Corresponding Author: Narmada Rana: narmadarana123@gmail.com
Abstract
Citation: Rana N., Ghimire S. K.
(2016)Variation in Adaptive Traits
of an Endemic Meconopsis
napaulensis DC. along an Elevation
Gradient in Alpine Himalaya,
Central Nepal.Open Science
Journal 2(1).
th
Received: 4 Juy 2016
th
Accepted: 17 November 2016
th
Published: 6 February 2017
Copyright:© 2016 This is an
open access article 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.
Funding: The author(s) received
no specific funding for this work.
Competing Interests: The
author have declared that no
competing interests exists.
Understanding the pattern of variation in adaptive traits of rare
and endemic species along environmental gradient can suggest
important implications for developing optimal strategies for
species conservation and sustainable management. In this study
we aimed to study variation in life-history traits of an endemic
species M. napaulensis DC. along an elevation gradient in
Langtang National Park, Central Nepal covering three
populations. Each population was investigated two times (2013
and 2014) covering different seasons. Population sampling was
made to read habitat during the peak growing period (during
monsoon) except seed output, which was studied during the
late growing period (post monsoon). Entire area of each
population was extensively surveyed to record all the
individuals, including plants in flowering or fruiting and their
detailed vegetative characteristics and traits related to
population fitness. Analysis of the habitat features showed that
M. napaulensis exhibited high habitat specificity. M. napaulensis
was restricted to open and rocky habitats of high altitudes. M.
napaulensis growing sites had low vegetation cover indicating
decreased interspecific competition. M. napaulensis showed high
variation in traits due to altitudinal variation, climatic
conditions, and disturbances.
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Specifically, reproductive traits discriminated the populations. M.
napaulensis was suffered from human disturbance mainly from
livestock grazing, plant harvesting, and garbage pollution as the
study area is an important tourist destination and pilgrimage site.
Conservation of rare and endemic species such as M. napaulensis
requires strong provisions restricting human activities and
minimizing the impact of harvesting and grazing; and
implementing habitat restoration and population augmentation
programs.
Keywords: Endemic, Elevation gradient, Population ecology, Density,
Vegetative traits, Reproductive traits.
Introduction
Elevation gradient is correlated with several environmental factors. With the
increasing elevation, plant populations are subjected to a gradually decreasing
mean temperature, and a shorter growing season (Landolt, 1967; Körner, 2003).
Elevation gradient is therefore ideally suited for examining variations in species’
traits, which strongly influence fitness (Minden, 2010), including growth and
competition (Wright et al., 2006). Plant adaptive traits are evolved in response to
the changing environmental conditions, and therefore exhibit considerable
variations along elevation gradients (Westoby and Wright, 2006), including
gradients associated with disturbance (Daiz et al., 1999; Wana and Beierkuhnlein,
2009). When comparing plant populations growing on lower elevation areas with
that of higher elevation, an obvious adaptation of high altitude plants to the
adverse environment is found to be in the reduced size (Jenny-Lips, 1948; Körner,
2003). Plant species growing along the elevation gradient show considerable
variations in life history strategies, including the structure of their populations
and demography (Kim and Donohue, 2011). Studies pertaining to variations in
adaptive traits along the elevation gradient may provide opportunities to examine
performances of plant populations under environmental changes (Kim and
Donohue, 2011).
In a high altitude ecosystem, plant populations are subjected to an increasing
level of anthropogenic pressure, created by habitat destruction, deforestation, and
overexploitation (Ghimire et al., 2008; Sharma et al., 2009). Such areas are
particularly vulnerable to natural variations in climate (Sano et al., 2005;
Cavaliere, 2009; Salick et al., 2009). Studies have shown that populations of rare
endemic species exhibit reduced size, altered population structure, reduced
fitness, and ultimately face greater extinction risk in the response to altered
environmental conditions at high altitudes (Brune and Kress, 2002; Lavergne,
2005; Colling and Matthies, 2006; Dar et al., 2006). Reduced fitness in many rare
and endemic species is associated with limited success in sexual reproduction,
which can be attributed to reduced pollination, failure in the formation of viable
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seeds, increased herbivory and predation pressure (Morely, 1982; Menges et al.,
1986; Karran, 1987; Lavergne, 2005; Albert et al., 2005).
Seed size is the prominent life history trait that affects seed dispersal,
seedling establishment and survival (Leishman et al., 2000). Under the
increasingly adverse ecological conditions along an altitudinal gradient, seedlings
from heavy seeds might be more successful in establishment and survival
(Leishman et al., 2000; Moles and Westoby, 2004). Studies have shown that
alpine plant species exhibit a reduction in seed size with the increasing elevation.
Low temperatures and short growing seasons at high elevations are the two most
important factors affecting seed maturation and seed weight (Totland and Birks,
1996; Totland 1997a; Wagner and Reichegger, 1997; Baskin and Baskin 1998;
Blionis and Vokou, 2005).
In alpine environments, plant demography is often characterized by low
seedling recruitment and high mortality at early developmental stages compared
with lower-elevation populations (Billings and Mooney, 1968). Demographic rates
in plants are usually stage-dependent (Harper, 1977), the structure of a
population may be indicative for its demographic future. Moreover, lowered seed
output is one of the major threats to plant life-history processes influencing the
population age-stage structure directly, and may increase the probability of
extinction of populations and species in the long-run (Lennartsson, 2002).
Long-term persistence of a species population depends on the continuous
regeneration (Thakuri, 2010). A regeneration of a species is greatly influenced by
habitat conditions. However, a natural regeneration of a species largely depends
on production and germination of seeds and the establishment and survival of
seedlings. Potentialities of a species’ regeneration can be depicted through the
analysis of population structure (i.e. the proportion of plant individuals classified
into different stage/age classes) (Bharali et al., 2012). Study on the variation in
population structure along elevation gradients would be helpful in understanding
the influences of environmental factors on regeneration (Wang et al., 2004).
The present study was conducted to assess the responses of Meconopsis
napaulensis populations along the elevation gradient in north-central Nepal.
Meconopsis napaulensis is an endemic species, distribution of which is restricted
to the alpine areas of Langtang National Park, north-central Nepal. The
conservation and management of rare and endemic species is a major challenge in
the Himalaya. The genus Meconopsis as whole is one of the critically threatened
taxa (Sulaiman and Babu, 1996). Among the species in Meconopsis, rare and
endemic ones, such as M. napaulensis, are predicted to be more vulnerable. This
study explores habitat properties, population structure, and fitness-related traits
of M. napaulensis along the elevation gradient, which can provide a link between
population responses to the environmental change.
Methods
Study area
The study was undertaken in upper Trishuli valley between Lauribina danda
to Gosainkunda in north-central part of Langtang National Park (LNP).
Langtang National Park (LNP) was established in 1976 to conserve the unique
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flora and fauna of the region. LNP covers an area 1,710 sq. km. It is located at
o
o
28 28’20”N latitude and 85 15’86”E longitude. Elevation in the park ranges from
792 to 7245 m. The area is rich in terms of flora and vegetation. The focused
study area covers alpine zone with an elevation range of 4000-4500 m a.s.l.
Meconopsis species are the important component of vegetation on the alpine zone
in upper Trishuli valley.
Distribution
The genus of Meconopsis Vig. (Family: Papaveraceae) consists of 55 species
distributed in the Himalaya (Egan and Shrestha, 2011). In total, 22 species are
reported from Nepal, out of which 11 are endemic to the high altitude areas
between 2400-4900 m (Grey-Wilson, 2006). Most of the endemic species of
Meconopsis are restricted to Central Nepal; there are only three endemic species
in Western Nepal (i.e., M. chankheliensis, M. simikotensis and M. regia) and one
(M. dhwojii) in Eastern Nepal (Egan and Shrestha, 2011) (Appendix I). The
endemic M. napaulensis is very local in distribution and is known only from
Gosainkunda and Ganesh Himal area of Langtang National Park, and its
adjoining region in Rasuwa district, central Nepal (Grey-Wilson, 2006). It is
found in rocky, grassy slopes, open shrub berries, open rocky shrubs, rocky
grassland, and stream- margins at 3200-4500 m a.s.l. (Egan and Shrestha, 2011).
Ecology
The members of the genus Meconopsis are poorly known with respect to their
reproduction traits, fitness, and population ecology. Sulaiman and Babu (1996)
found that Meconopsis species are habitat specialist. In Meconopsis, a large
number of seeds are produced but most of them are exposed to a high level of
insects, fungal and viral infections (Sulaiman and Babu, 1996). Germination and
seedling recruitment have also been reported to be very low both in natural
habitat and under laboratory conditions (Xie et al., 2002). Several studies
conducted for different species of genus Meconopsis revealed the existence of
physiological dormancy hindering seed germination (Sulaiman 1993; Dar et al.,
2009).
Sampling
The study was completed in two field visits (the first visit was made in 2013
and the second visit in 2014). The distribution of M. napaulensis populations in
the study area was recorded with the help of GPS device and consultations with
local people. Based on the information obtained from the field survey, as well as
from literature and herbarium study, the whole of the distribution range of M.
napaulensis was divided into three distinct elevation levels each representing
distinct population: (i) lower elevation (4117-4125 m) in Lauribina danda, (ii)
mid-elevation (4200-4255 m) in Lauribina pass, and (iii) higher elevation (43964417 m) in Gosainkunda.
In each elevation level, M. napaulensis was sampled in four plots of 10 × 10
m size. The plots were laid systematically in the area where M. napaulensis
density was high. Each plot was divided into 4 subplots (5 × 5 m); and thus
altogether 48 such subplots were sampled from all the three elevation levels. Most
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of the population parameters were collected during the peak growing period in
July (during monsoon) except seed output, which was studied during the late
growing period in September (post monsoon season).
Data Collection
In each subplot, presence/absence of plant species associated with M.
napaulensis was recorded. In boththe years (2013 and 2014), sampling consisted
of recording all the individuals of M. napaulensis, classified into four different life
cycle stages, on the basis of the plant size, reproductive stage and the number
and size (length) of the largest leaf; small rosette (leaf number 1-7; largest leaf
length 1-6 cm), juvenile rosette (leaf number 3-28; largest leaf length 6-14 cm),
large rosette (leaf number 4-45; largest leaf length >14 cm), and reproductive
adult (with flowering/fruiting peduncle). Each of these individuals was tagged by
aluminium tag in 2013 and was monitored for the change in the population
structure in 2014.
Longitude, latitude, and elevation were recorded for each sampling plot with
the help of global positioning system device (GPS, eTrex Garmin). Elevation
data was cross-checked with an altimeter. Slope and aspect were recorded by a
clinometer-compass. Habitat parameters for each subplot further included soil
pH, soil moisture, litter depth, vegetation composition, and ground surface cover
by vegetation or physical components of the environment. Soil moisture and pH
were recorded by using a gauge (soil pH and moisture tester; model DM 15) with
a default scale of 1 to 8 for moisture and 1 to 7 for pH recording. Similarly, soil
depth was measured by inserting an iron peg. In each subplot, pH, moisture, and
depth were measured diagonally at three different points.
The anthropogenic disturbance was recorded in each plot. The disturbance
variables included harvesting, garbage pollution, and grazing. As Gosainkunda is
known for its pilgrimage, more pilgrims visit every year in August especially in
‘Janaipurnima’. They pluck flowers for offering to God and also pluck roots,
fruits, and seeds for medicinal purposes (based on interviews with local people) so
there is a high rate of harvesting. In addition, garbage pollution was recorded
along the trekking route due to more flow of people for trekking which turned
more use of things along the trekking route (based on interviews with local
people). The harvesting impact was recorded by direct observation of plant
uprooting or by observing the scars left after plucking of flowers and/or fruits.
Each type of disturbance was scored by categorical scale as no disturbance,
moderate disturbance, high disturbance, very high disturbance.
Vegetative and reproductive attributes
All the individuals from each subplot were marked in 2013 by aluminium tag.
Each individual was thoroughly inspected for recording a number of vegetative
and reproductive traits having significant adaptive value to the plant. Parameters
recorded from the individuals in rosette stage (non-reproductive) were the total
number of leaves, and the length of the largest leaf. Parameters recorded from the
individuals in reproductive adult stage included plant height, and the number of
buds, flowers, and fruits (capsules).
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The number of flowers and buds were counted in the flowering period and
the number of capsules in the fruiting period. During the flowering period, 5
flowering individuals from each population were randomly selected and their
flowers (n = 2) were collected for the study of pollen viability. Similarly, during
the fruiting period, 5 matured individuals were marked per population for capsule
harvesting. From each such plant, at least five matured but unopened capsules
were collected. Ten such capsules were randomly selected per population for the
measurement of seed size (in terms of seed mass) and seed viability. Capsule size
was measured with the help of Vernier calliper. Both length and diameter of a
capsule were measured. The diameter was recorded as the mean of upper, middle
and lower portions of the capsule.
Pollen and seed viability
Glycero acetocermine (1:1) mixture was used for the treatment of pollen
viability test assessed by Belling’s Iron-Aceto-Carmine staining method (Singhal
and Kumar, 2008). Pollen grains from each flower were treated with Glycero
acetocermine (1:1) mixture and were studied under a compound microscope.
Well–filled pollen grains with stained nuclei were regarded as fertile/viable, while
shriveled and unstained pollen were counted as sterile. Similarly, for the seed
viability, Triphenyl tetrazolium chloride (TTC) test (Baskin and Baskin, 1998;
Lin et al., 2001) was used. In this process, seeds of M. napaulensis, collected from
different populations, were cut into two equal halves in such a way that each part
got a portion of the embryo. Then treated with 1% solution of TTC (Dar et al.,
2009) and observed under stereo-microscope. Seed viability was revealed by pink
TTC precipitation produced by dissected seeds. Embryos that turned pink were
considered as viable and other as non-viable.
Data Analysis
Habitat characteristics of M. napaulensis were evaluated by non-parametric
Kruskal- Wallis one-way analysis of variance (ANOVA) as the data was found
not normal. Variables related to the anthropogenic impact were combined by
using Principle Component Analysis (PCA) to obtain an overall measure of
disturbance. In this process, the impact of harvesting and garbage pollution was
combined as an overall measure of the human impact and was obtained
explaining 63.49% variance and grazing as a measure of the livestock impact.
The numbers of associated species present per subplot were combined to
calculate abundance in an ordinal scale from 0 to 4, where 0 represents absence
in all subplots and 4 presence in all subplots. The abundance data of 71 associate
species from all 48 subplots were used to calculate their frequency and dominance
mean value in lower, middle, and higher elevation sites.
Variations in population density and structure were studied at subplot level
(5 x 5 m). Population structure indicated the proportions of individuals of
different life cycle stage (small rosette, juvenile rosette, large rosette, and adults).
Variation in population density among elevation sites was compared using
Kruskal-Wallis test.
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Vegetative and reproductive traits were computed for each adult individual
in the respective elevation site. The total data set comprised of 75 adult
individuals. The sample size for estimating capsule production comprised of 69
individuals of which 25 individuals from each, the lower and the middle site, and
19 individuals from the higher site, each individual’s seed weight was recorded.
Similarly, the sample size for viability test for seed and pollen comprised of 30
individuals of which 10 individuals from each, the lower elevation to the higher
elevation.
Kruskal-Wallis test and independent sample Wilcoxon test were used to
assess the variation in vegetative and reproductive traits, including seed and
pollen viability, among and between study sites (Table 1). SPSS version 16.0 was
used for all statistical analysis. Arc GIS 9.3 version was used for mapping.
Results
The habitats of M. napaulensis differed significantly in 6 out of 15 variables
studied. Coverage of herbs was greater at the higher elevation site and that of
shrubs was higher at the lower elevation site. The litter coverage was found
significantly higher at the lower elevation which showed that habitat at the lower
site was more fertile than at the upper site. Among the edaphic variables, the
value of soil depth was high at the lower site. As the elevation increases, more
rock and scree are found due to which the amount of soil and its depth reduced
at the high altitude. The level of human disturbance was high in plots on the
higher elevation sites. In general, higher proportions of sub-plots in populations
at Gosainkunda and Lauribina pass received higher levels of garbage pollution
and high livestock grazing pressure (Figure 1).
Figure 1. The level of disturbances in the three study sites. Bars represent
proportion of sub-plots receiving different levels of disturbance.
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Population density
The densities of small rosette (SR), juvenile rosette (JR), large rosette (LR),
and adult of M. napaulensis in the entire study site were found to be
0.072±0.016, 0.124±0.029, 0.194±0.027, and 0.080±0.018 (mean ± SE)
individuals per 25 m2 (Figure 2). The overall density combining all four stage
classes was 0.470±0.061 individuals per 25 m2. The value of total plant density
tended to be high at the higher elevation site, but the result was statistically
insignificant.
Population structure
0.12
(a)
0.12
(b)
0.1
0.1
0.08
0.08
0.06
0.06
0.04
0.04
0.02
0.02
0
0
SR
0.12
JR
LR
SR
REP
JR
LR
REP
0.12
(c)
(d)
0.1
0.1
0.08
0.08
0.06
0.06
0.04
0.04
0.02
0.02
0
0
SR
JR
LR
REP
SR
JR
LR
REP
Figure 2. Population Structure. Proportion (mean =SE) of small rosette (SR), juvenile rosette(JR), large rosette(LR)
and adult(REP) and overall pattern of M. nepaulensis at three elevation sites in Upper Trishuli Valley of Langtang
(a-c):(a) Lauribina danda (4117-4125m); (b) Lauribina pass (4200-4255m), (c) Gosainkunda(4396-4417m) and (d)
Overall Population Structure.
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Table 1. Variation in vegetative and reproductive traits, including seed and pollen viability
Variables
N
Mean ± SE at three sites
(LP)
Overall
Mean ± SE
Kruskal-Wallis test
among sites*
p-value
χ2
Z-values indicating difference
between¥
LD &LP
LD & GK
LP & GK
(GK)
(LD)
Plant height (cm)
No. of buds per plant
No .of flowers per plant
No. of capsules per
plant
Capsule length (cm)
Capsule diameter (cm)
Capsule mass in g
(with seeds)
Capsule mass in g
(without seeds)
Seed mass/capsule in g
No. of seeds §
75
189
101
69
78.00±1.53
17.2±1.8
8.4±0.7
24.2±2.38
74.2±2.26
13.4±1.83
8.00±0.86
19.6±2.47
66.00±3.71
8.05±1.01
4.63±0.72
9.95±1.66
73.32±1.52
13.30±1.05
7.22±0.48
18.61±1.48
6.547
15.166
9.527
17.359
0.038*
0.001**
0.009**
<0.001***
-1.968*
-1.716
-0.489
-1.716
-2.498*
-3.877***
-2.993**
-3.871***
-0.06
-2.305*
-2.394*
-2.991***
69
69
69
3.91±0.12
0.46±0.01
0.16±0.01
3.8±0.00
0.47±0.01
0.15±0.01
3.00±0.00
0.48±0.01
0.14±0.02
3.62±0.06
0.47±0.01
0.15±0.01
36.337
1.946
3.59
<0.001***
0.378
0.166
-1.57
-0.679
-0.466
-4.732***
-1.28
-1.885*
-6.557***
-0.948
-1.339
69
0.08±0.01
0.09±0.01
0.1±0.01
0.09±0.01
2.176
0.337
-0.398
-1.328
-1.211
69
15
0.07±0.01
154.8±41.08
0.04±0.01
226±25.64
0.06±0.01
169.67±19.29
7.474
4.994
0.024
0.082*
-0.68
000
-2.677**
-2.402*
-2.002*
-1.358
Seed viability (%)
Pollen viability (%)
30
30
0.08±0.01
128.2±16.8
7
4.1±0.35
7.6±0.49
3.5±0.34
7.3±0.54
2.6±0.22
5.9±0.28
3.4±0.21
6.93±0.28
8.734
6.475
0.013*
0.039*
-1.258
-0.501
-1.944*
-1.952*
-2.784**
-2.392*
Significance based on independent sample Wilcoxon test, rest of the values are based on Kruskal-Wallis test (nonparametric of one-way ANOVA).
§ Number of seeds based at a constant weight of 0.01gm (number of seeds were counted present at a constant weight
of 0.01 gm).
Asterisk indicate that the medians for a particular parameter between sites are significantly different from one another
at *p < 0.05, **p<0.01 and ***p<0.001.
Discussion
Plant population fitness is influenced by several ecological factors.
Populations of M. napaulensis occupied alpine habitats (4117- 4417 m a.s.l.)
experiencing strong ecological heterogeneity. In the majority of cases, plant
populations are grown in open rocky habitats in south-west facing slopes. The
similarities in most of the edaphic characteristics (such as soil pH, moisture, and
litter depth) and substrate types (such as solid rock, non-vascular plant cover,
grass cover, and total vascular plant cover) among sites revealed that M.
napaulensis is habitat specific. Previous studies (Sulaiman and Babu, 1996; Lesica
et al., 2006 and Poudeyal, 2010) revealed that certain species of Meconopsis are
restricted to extreme edaphic conditions (nutrient-poor, dry, and open rocky
substrates) where competition from dominant vegetation is reduced. Debussche
and Thompson (2003) reported habitat specialization is the important relative
traits for the rare endemic species. The habitat specific species like M.
napaulensis should have a strong correlation with edaphic endemism and other
ecological specificities (Ghimire, 2005; Poudeyal and Ghimire, 2011).
The statistically insignificant difference in the density value of individuals in
different stage/state classes (except reproductive size class) among three sites
further supports that the population structure and stage class distribution of
Meconopsis napaulensis largely depend upon the specific range of ecological
amplitude. The regeneration capacity of rare endemic species generally depends
upon the availability of suitable micro-habitat (Tilman et al., 1994). The patchy
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nature of M. napaulensis could also be linked to the limitation of suitable microhabitat sites in surrounding areas. Thus, the distribution of such rare endemic
plants are strongly influenced by the availability of suitable micro-habitat
(Tilman et al., 1994).
Regarding the ground cover vegetation, shrubs coverage was found more at the
lower altitude site which creates a high competition for the availability of soil
nutrients and for sufficient amount of light (Sulaiman and Babu, 1996). It might
be the reason of reduction of the density of reproductive individuals at the lower
altitude sites. The litter coverage was found significantly higher at lower
elevations which showed that habitat at the lower site was more fertile than on
the upper site. In this study, the moss coverage was found to be high at the lower
elevation site. It retains a high amount of moisture which enhances the vegetative
growth of plants (Korner, 2003; Devkota, 2009). Thus, the higher level of
vegetative growth of M. napaulensis was found to be associated with ecological
integration including edaphic factors at the lower altitude. Conversely, the seed
production rate was negatively associated with surrounding vegetation. Although
the high altitude plant population tries to counteract allocating higher
investment towards reproductive growth but due to harsh environmental
conditions including poor nutrient substrates, the overall growth of the individual
was substantially reduced in comparison to the lower altitude plants.
Distribution:
Populations of M. napaulensis are also influenced by various levels of
anthropogenic disturbance like in other parts of Himalaya (Ghimire et al., 2006
and 2008). The over-harvesting of medicinal plants, excessive grazing, and
trampling are the major anthropogenic factors which lead to the habitat
fragmentation, destruction, and ultimately cause local extinction. In the case of
M. napaulensis at Gosainkunda area, grazing seems to be a driving factor for
habitat fragmentation along with harvesting for medicinal and religious purposes.
The availability of pastoral land towards high altitudes provides the distribution
of the grazing resources (Fox et al., 1996; Ghimire et al., 2006). Present findings
also indicate that the anthropogenic effect was higher at Gosainkunda site in
compare to other sites. As indicators to evaluate disturbances, higher levels of
harvesting, grazing, and garbage accumulation in Gosainkunda site proved the
higher level of anthropogenic influence on the high altitude population.
Further, most the populations of M. napaulensis are along the trekking
(Lauribina and Gosainkunda) route and it is one of the important factors that
plant populations receive a significant amount of anthropogenic disturbances.
Similarly, the Gosainkunda Lake is one of the religiously important lakes among
Hindus and Buddhists. Nearly 30,000 pilgrims visit Gosainkunda Lake each year,
during the festival ‘Janaipurnima’ and generate significant human disturbances
(personal interviews with locals). During the festival, people who visit usually
pluck the flowers and offer them to the Gosainkunda deity. At the mean time
collection of capsules for medicinal use was also a prime factor for depletion of
the population. Previous study (Poudeyal, 2010) denoted that M. napaulensis
roots, leafs, and seeds are used to treat digestive system disorder, chest pain, sore
throat, and headache. Premature harvesting of capsules for medicinal purposes
was prevalent in Gosainkunda and Lauribina. Thus, an unsustainable harvesting
of M. napaulensis could be one of the major challenges for the long-term
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population viability. Low to moderate levels of grazing showed a positive
relationship with distribution and abundance but higher levels of grazing showed
a negative relationship due to the destruction and elimination of the species
(Ghimire et al., 2006). The management plan for the pasture land (for grazing)
and ecologically iconic species (for conservation) on the same area cannot go
independently, so a systematic approach incorporating social management to
accommodate the needs of different users at the landscape level is needed
(Ghimire et al., 2004 and 2006). Based on the results, M. napaulensis showed
identical densities in all stage classes, except reproductive class, among all the
three sites. The reproductive (stage) density was significantly high at Lauribina
pass. In comparison to the other sites, Lauribina pass received lesser
anthropogenic pressure because the site is located away from the human
settlements. A previous study suggested that the extent of disturbance in alpine
plant population is related to the distance from the settlements (Poudeyal and
Ghimire 2011). Thus, Lauribina danda and Gosainkunda sites which are located
nearer to the human settlements received a higher amount of disturbances.
Meconopsis napaulensis showed variation in most of the vegetative and
reproductive characteristics. The plant height is one of the discriminating
characteristics among all the sites. Plants at the lower altitude site (Lauribina
danda) were taller in comparison to the high altitude sites. Korner et al. (1983)
and Brown et al. (2003) reported that reduction in plant height mostly related to
the corresponding decline in temperatures and short growing period in the alpine
area. The poor availability of nutrients in the soil, thin soil profile, and further
higher level of anthropogenic influence also could be related to the stunted
growth of plants in the high altitude site (Gosainkunda). Similarly, the higher
allocation on vegetative investment in the lower altitude site supports the
production of a higher number of buds, flowers, and capsules. But in contrary to
the vegetative growth the number of seeds was significantly high at the high
altitude site. Xie et al. (2002) has reported that a large number of seed
production is the tendency of rare endemics. Similar is the case for M.
napaulensis that produces an innumerable number of tiny seeds. Plants at the
high altitude site counteracted with that of the lower altitude site by producing a
higher number of seeds and a higher allocation towards reproduction in
comparison to the vegetative growth. But seed viability tests suggest that the
high altitude site produced far less viable seeds. Thus the overall plant
performance was found to be always higher in the lower altitude populations. M.
napaulensis at high altitudes produced light and ill-developed seeds. Seed weight
was significantly reduced at higher altitudes if the plants from Gosainkunda
(4417 m) and Lauribina danda (4117 m) were compared. The maturation and
production of intact seeds in alpine plants is directly linked to the favorable
environment (Totland, 1997b). The study revealed that alpine plants are more
sensitive to temperature for seed feeling and maturation (Wagner and Reichegger,
1997). The frequently changing weather conditions, low temperatures, and short
growing seasons hindered the production of healthy and heavier seeds at the
higher altitude (Pluess et al., 2005; Wagner and Reichegger, 1997; Totland,
1997b; Totland and Birks, 1996; Corbet, 1990; Galen, 1985). The production of
intact seeds per plant was also related to the extent of disturbance. Escarre et al.
(1999) reported that floral damage by herbivores limits the seed production.
Similarly, suitable pollinator limitation, pollen limitation in a fragmented habitat
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Research Article
like M. napaulensis are also major challenges to develop well-firmed seeds in
alpine habitats (Morely, 1982; Menges et al., 1986; Karran, 1987; Lavergne, 2005;
Albert et al., 2005).
Conclusion
Meconopsis napaulensis prefers an open rocky substrate, nutrient-poor,
grassy slopes in south-west facing slopes. Habitat characteristics are directly
linked to the vegetative and reproductive attributes. Lower altitudes provide
fertile and nutrient-rich habitat for vegetative growth and reproductive outputs.
Vegetative and reproductive traits of Meconopsis napaulensis showed variation
along the elevation gradient. Lower altitude plants showed proliferation towards
the vegetative outputs in comparison to higher altitude which could be linked to
the favorable environment such as higher level of soil nutrients, moisture, and
higher temperature.
In contrary to the vegetative growth, high altitude plants tended to show a
higher number of seeds per capsule. Thus in this regard, a tradeoff among the
plant traits was observed in which lower altitude plants invest higher allocation
on vegetative growth and lesser investment on reproductive growth and
development. Contrarily, high altitude plants showed a higher investment on
reproductive output in comparison to the vegetative investment.
Plants in four stage/size classes did not showed marked differences in density,
but numerically the number of large rosettes is slightly higher than that of all
other stages. The population structure was identical in all sites studied. In
regards to density of the plants, reproductive plant density was markedly differed
among all the sites and stages. The reproductive density was far higher in
Lauribina pass which could be linked to the safe habitat in comparison to the
other two sites. The latter two sites were nearer to the human settlements and
received a higher level of disturbances.
Acknowledgements
We are thankful to Department of National Parks and Wildlife Conservation
(DNPWC) and Langtang National Park (LNP) for granting permission to carry
out this work inside the National Park Area under the Biodiversity Associates for
Research, Development and Action, Nepal (BARDAN). Our sincere thanks goes
to the local people of Deurali, Chandanbari, Cholangpati, Lauribina, and
Gosainkunda for their help and support, and for sharing their knowledge about
the studied species.
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