International Society
for Tropical Ecology
Tropical Ecology
https://doi.org/10.1007/s42965-020-00109-2
RESEARCH ARTICLE
Environmental determinants of plant associations and evaluation
of the conservation status of Parrotiopsis jacquemontiana in Dir,
the Hindu Kush Range of Mountains
Fazal Manan1 · Shujaul Mulk Khan1
Majid Iqbal1 · Abdullah1
· Zeeshan Ahmad1 · Saqib Kamran1 · Zahoor Ul Haq1 · Fatima Abid1 ·
Received: 20 December 2019 / Revised: 27 August 2020 / Accepted: 7 September 2020
© International Society for Tropical Ecology 2020
Abstract
Hindu Kush is the largest mountain range of Central Asia that forms part of a vast alpine zone that stretches across the Eurasia from east towards the South Asia. We studied vegetation structure and the role of edaphic and topographic factors on
distribution and formation of plant associations with specific emphais on Parrotiopsis species of the Districts Dir regions in
the Hindu Kush Mountains. We also assessed the conservation status of Parrotiopsis jacquemontiana, an endemic species
of the western Himalayan floristic province. We hypothesized that edaphic and climatic factors were responsible for the
formation of different plant associations each with distinct indicators. A combination of transect and quadrat based methods were used for sampling. We used two way cluster analysis (TWCA), cluster analysis (CA), indicator species analysis,
detrended correspondence analysis and canonical correspondence analysis to analyze and elaborate the vegetation pattern and
formation. We used Google Earth Path software (V 1.4.6) for the calculation of extant of occurrence (EOO) and area of
occupancy (AOO) for evaluation of conservation status of P. jacquemontiana. A total of 142 plant species were reported
belonging to 62 families. CA and TWCA clustered four plant associations within altitudinal range of 1556–2313 m. Parrotiopsis jacquemontiana should be designated as endangered species under ‘EN A2acd; B1B2 bc (i, ii, iii) of IUCN red list
categories and criteria in the region. We found that high phosphorous and potassium concentration, elevation, aspect, slope,
lower pH, electrical conductivity and soil texture were significant environmental variables that play an important role in the
determination of vegetation structure, formation of plant associations and its indicators in the region. This information will
be useful for conservation and management practices for endemic and rare plant taxa, and evaluation of vegetation structure.
Keywords Conservation · Hindu Kush mountain range · Western Himalayan Province · IUCN · Multivariate statistics ·
Parrotiopsis jacquemontiana · Plant associations
Introduction
Plant species represent the hierarchical expression of vegetation as they are affected by environmental factors (Gaston
2000; Shaheen et al. 2017). Various types of analyses have
been used to find out important plant species and communities/associations formation (Fujiwara 1987; Dhyani et al.
2019). Multivariate statistical approaches can help ecologists
describe and display the vegetation structure and identify
* Shujaul Mulk Khan
shujaqau@gmail.com
1
Department of Plant Sciences, Quaid-i-Azam University
Islamabad, Islamabad, Pakistan
effects of environmental gradients on vegetation or whole
group of species in effective ways (Hair et al. 2006; Ahmad
et al. 2019). Statistical tool can provide efficient means to
reduce the complexity inherent in natural vegetation and to
detect important environmental factors that explain this complexity (Dufrêne and Legendre 1997; Khan et al. 2011; Haq
et al. 2020). Plant communities, which can be described both
physiognomically and floristically have defined structure in
relation to both abiotic and biotic factors (Kent 2011). The
development of vegetation structure is typically influenced
by environmental gradients such as topographic and edaphic
factors (Ismail et al. 2019; Khadanga and Jayakumar 2020).
Various environmental variables such as climate and soil
conditions can affect the distribution, recognition and classification of indicator species (Khan et al. 2013; Ahmad
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et al. 2016b). Plant associations are organized on the basis
of both phytosociological attributes and environmental factors (Cook and Kairiukstis 2013; Ullah et al. 2015; Zeb
et al. 2020). Different ecological associations are formed
when their formation is based on indicator values. Regarding
different environmental gradients, the predictable richness
of every species are designated by indicator values (Braak
and Barendregt 1986; Dufrêne and Legendre 1997). Each
habitat type usually has one or more indicator species. These
sensitive species can be used to monitor changes in plant
communities that result from environmental or management
changes. An effective and meaningful description of plant
associations and their component communities can inform
management decisions for diverse ecological assemblages
that may be stenotopic and eurytopic (Iqbal et al. 2018; Rahman et al. 2020).
Poorly planned urbanization, deforestation and overexploitation of natural resources are occurring at an alarming scale in the developing countries (Sheikh et al. 2002).
Human population is increasing at a rate of 2.1% annually,
which directly increases pressure on vegetation and wildlife. A total of 1572 genera and 5521 species of flowering plants have been documented from Pakistan (Ali and
Qaiser 2010b), but no comprehensive national red lists
have been complied. Threatened plants of Pakistan are not
fully explored (Jan and Ali 2009; Ali and Qaiser 2010a, b).
According to Nasir (1991) the number of nationally threatened flowering plants were 580–650 (i.e. 12% of the total
flora). However, this estimation was not based on any criteria. According to Walter and Gillett (1998) there are 14
threatened species of flowering plants in Pakistan. Only 26
species are on the Red list 2010 from Pakistan. Eight Himalayan medicinal plants were assessed for their conservation
status by Muhammad (2003), but this method was based on
consumption and availability in markets (i.e. data on population size, extent of occurrence and area of occupancy was
not obtained).
Parrotiopsis jacquemontiana is an important endemic
plant of the Western Himalayan floristic province (Takhtajan et al. 1986). This species is facing serious threats in the
region (Hazrat and Wahab 2011; Ali et al. 2018). Therefore,
our study included an assessment of its occurrence, area of
occupancy and estimate of population size. Endemic taxa
need special attention because any unfavorable alteration
in habitat may lead to quick extinction. We hypothesized
that edaphic and climatic factors affect the formation of
different plant associations each with a distinct indicator.
Our main objective was to document, classify and quantify
plant species into various associations to analyze influence
of different environmental variables on vegetation structure.
This study can be used as a baseline for further ecological
research for the formation of various plant association or
communities and in identification of its associated indicators
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via multivariate statistical approach. It could also be used to
determine the conservation status of any species in general
and endemic species in particular via extent of occurrence,
area of occupancy and data on population size techniques.
Materials and methods
Study area
Dir is the combination of two districts, Dir Lower and
Dir Upper located in Khyber Pakhtunkhwa Province of
Pakistan between 35° 50′ to 34° 22′ N latitude and 71°
2′ to 72° 3′ E longitude of Hidukush range. The study
area is surrounded by high mountains. The Koh-i-Hindu
Raj mountain range spreads from north to south on the
western border and separates district Dir from Afghanistan
and Chitral. Floristically, it lies in the Western Himalayan
Province of Irano-Turanian region (Takhtajan 1970). Geographically, District Swat lies in the east, Chitral on the
north, Afghanistan on the west and Malakand agency on
the south (Hazrat et al. 2007). Their total areas comprise
5284 km 2 with 450.8 persons per kilometer population
density (census report of 2017) (Fig. 1). Metamorphic and
Igneous rocks dominate the study region which comprises
four main types of rock i.e., Amphibolite, Quartz-felsparbiotite, rocks of diorite group and a chain of aplite-pegmatite-dykes. The well foliated amphibolite has gneissic
and quartzite interlayers and additions of granitic rocks as
well. Flora of the district Dir consists on various kinds of
roses, Bougain viIlea, Jasmine, Geranium, Gul-e-dawoodi,
Gul-e-khairo, Gardinia, Pollen Zira, Banafsha, Gangora,
Berg-e-Sumbol, Antedolt, Sarlobal and Zahar Morha (Haq
et al. 2020).
Vegetation sampling
A total of 21 altitudinal transects (varied in length from
250–800 m) were established where the species of Parrotiopsis jacquemontiana were present at a distance of
3–5 km. Along each transect 3–7 quadrates were laid
down depending upon the length of each transect. The
sampling began at the lowest elevation to the mountain
peak (1540–2313 m). A total of 318 quadrats were established having 100 × 100 m, 25 × 25 m and 1 × 1 m size,
106 each for trees, shrubs and herbs, respectively (Moore
and Chapman 1986; Bano et al. 2018; Anwar et al. 2019).
For each transect absolute and relative density, cover and
frequency of each species, along with their Importance
Values Index (IVI) were calculated through formulae
designed by Curtis and McIntosh (1950) and Kamran
et al. (2019). Plant specimens were collected, labeled
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Fig. 1 GIS generated map of
the study area representing 21
transects with special reference
to elevation zones
with tags, placed between newspapers, pressed with plant
presser, poisoned using mercuric chloride plus ethyl alcohol solution and mounted on standard herbarium sheets
(Iqbal et al. 2015; Khan and Ahmad 2015; Ahmad et al.
2016a). All the specimens were identified with the help
of Flora of Pakistan and other available literature (Ali
and Nasir 1990; Khan et al. 2016b). The plant specimens were deposited in the Herbarium of Quaid-i-Azam
University Islamabad (ISL), Pakistan. Global Positioning
System (GPS) locations were recorded for each quadrat.
Aspect of the mountain i.e., south (S) and north (N) were
determined with the help of a digital compass. The soil
depth was estimated with an iron rod of 2 m length. Grazing pressure was estimated by classes 1–3 (low to high)
by observing recent signs and intensity of grazing effect.
Soil analyses
Soil samples were collected from three random points within
each quadrat up to a depth of 0.03 m and mixed thoroughly to
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make a composite sample (Ravindranath and Ostwald 2007).
Samples were placed in polythene bags and tagged with permanent markers. Larger soil particles were removed by sieving and the remaining samples were shade dried (Gee and
Bauder 1986; Khan et al. 2017). Physicochemical analysis
i.e., soil texture, pH and electrical conductivity (EC), organic
matters, phosphorus (P) and potassium (K) of each sample
was determined. Soil suspension was prepared and filtered
through filter paper. For determination of pH the electrodes
of pH meter was dipped into the prepared sample and reading
was noted directly (Jackson 1963). An electrical conductivity
(EC) meter was used for determination of EC (Jackson 1963;
Hussain et al. 1999; Wilson and Bayley 2012). Soil texture
was analyzed via hydrometer (Sarir et al. 2006; Bergeron et al.
2013). Organic matter concentration was determined by standardized solution of FeSO4 and K2Cr2O7 (Nelson and Sommers
1996), whereas for the determination of P and K (Soltanpour
1991) method was used for each sample.
Data analyses
All vegetation and environmental variables data were analyzed to assess the relationships between plant species composition and ecological factors using PCORD (version 5)
and CANOCO (version 4.5). PCORD was used to classify
plants into different associations through cluster analysis
(CA), two way cluster analysis (TWCA) and its associated
indicators via indicator species analysis (ISA) (Lepš and
Šmilauer 2003). Monte Carlo tests were used for statistical
significance after determination of Indicator Values of each
indicator. A threshold level of Indicator value of 25% with
95% significance (P value ≤ 0.05) was considered as cut-off
for identifying indicators and identified indicator species
(IS) were used for naming the plant associations (Braak and
Prentice 1988; Dufrêne and Legendre 1997). Species area
curves (SAC) were constructed using Sorenson distance
measure to assess sample size adequacy. Canonical correspondence analysis (CCA) was used for ordination analysis
in estimate effects of environmental variables on plant species composition, distribution pattern and abundance.
IUCN Red list Criteria
Different human and natural factors were recorded with reference to their impact on P. jacquemontiana population dynamics while interviewing local inhabitants in the study area,
as guided by the IUCN criterion (A–E). Google Earth Path
software (V 1.4.6) was used for the calculation of extent of
occurrence (EOO) and area of occupancy (AOO) available
on the IUCN official website (Khan et al. 2016a). The distribution and number of plants scored with reference to their
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ecological amplitude and calculated historical distribution
was compared with IUCN criteria for threatened categories
Version 3.1 (IUCN, 2001) for evaluation of the conservation
status of P. jacquemontiana. The plant species were then categorized into threatened categories (IUCN 2001).
Results
A total of 142 plant species were collected belonging to 62
families, including 21 (14.8%) trees, 24 (16.9%) shrubs,
and 97 (68.3%) herbs. The most dominant families were
Poaceae, Rosaceae, Labiatae and Asteraceae with 12, 11,
8 and 6 species, respectively.
Cluster analysis and two way cluster analysis
Cluster analysis classified all transects along with associated quadrats into four major associations (Fig. 2). Two
way cluster analysis further realized distribution of plant
species at each station that can be clearly seen in two main
branches of cluster dendrogram (Fig. 3). The detailed
descriptions of each association are as follows:Taxus baccata, Viburnum grandiflorum and Pteridium
aquilinum plant association
Cluster and two way cluster analysis clustered a total of 24
quadrats/stations in this association. The association name
was given based on indicator species analysis (ISA). The top
three indicator species of this association were Taxus baccata,
Viburnum grandiflorum and Pteridium aquilinum (Table 1;
Fig. 4). These were indicators of higher phosphorous concentration and steep slope environmental variables (Appendix Table 2). Dominant trees with highest Importance Values
Index (IVI) of this association were Pinus wallichiana, Cedrus
deodara and Quercus baloot, while the rare tree species were
Acer caesium, Ficus carica and Pinus roxburghii with minimum IVI in the region. Dominant shrubs were Sarcococca
saligna and Wikstroemia canescens, however, Indigofera
heterantha, Hedera nepalensis and Jasminum humile were
rare shrubs of this association. The herbs layer was dominated
by Viola odorata, Piptatherum laterale and Perilla frutescens
with high IVI values while Polygonatum cirrhifolium, Alliaria
petiolata and Artemisia vulgaris were the rare herbs of the
region (Appendix Table 2).IV = Indicator Value
The plants of this association were associated with
clay loam soil and grew on north and northwest aspects
at altitudes of 1758–2239 m. Electrical conductivity (EC)
ranged from 0.44 to 0.79 d/Sm, pH 6.39–6.42, organic
mater 0.6–0.75%, phosphorus (P) 2.4–9.2 mg/kg, potassium (K) 100–220 mg/kg and soil saturation 41–68%.
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Fig. 2 Cluster dendrogram of 106 studied quadrats along 21 transects establishing four major plant associations in the rgion
Ailanthus altissima, Rosa webbiana and Arenaria leptoclados
plant association
A total of 29 quadrats clustered this plant association. The
top 3 indicator species of this association were Ailanthus
altissima, Rosa webbiana and Arenaria leptoclados under
the influence of higher potassium concentration, west
aspects and low saturation in the region (Table 1; Fig. 5).
Quercus semecarpifolia, Cedrus deodara and Quercus
dilatata were the dominant species while Cornus macrophylla, Salix tetrasperma and Acer caesium were the rare
tree species of this association. The dominant shrubs were
Parrotiopsis jacquemontiana, Indigofera heterantha and
Cotoneaster microphyllus with high IVI values, whereas
rare shrubs included Sarcococca saligna, Viburnum cotinifolium and Lonicera quinquelocularis. The herbs layer was
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Fig. 3 Two way cluster analysis of PCORD software showing distribution pattern of 142 plant species among 106 quadrats establishing four
associations. The black dots illustrate the presence while white dots indicated absence of species in the region
dominated by Viola odorata, Fragaria vesca and Oxalis
corniculata while Trigonella sp., Scrophularia scabiosifolia and Oplismenus hirtellus were the rare herbs of the
association.
The plants of this association were associated with clay
loam soil and mostly occurred on northern aspects within
the altitudinal range of 1556–2313 m. EC ranged between
0.36–1.06 d/Sm, pH 5.25–7.68, OM 0.6–1.35%, P 1.5–8.1 mg/
kg, K 120–240 mg/kg and soil saturation 40–70%.
Quercus dilatata–Cotoneaster nummularia–Brachypodium
sylvaticum plant Association
A total of 42 stations comprised this plant association.
Quercus dilatata, Cotoneaster nummularia and Brachypodium sylvaticum were the top three indicators. These were
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the indicators of steep slope, higher number of stumps and
higher saturation environmental factors (Table 1; Fig. 6).
Quercus semecarpifolia, Pinus wallichiana and Quercus
baloot were the dominant Cedrus deodara, Celtis australis
and Platanus orientalis were rare tree species of the region.
The dominant shrubs were Parrotiopsis jacquemontiana,
Indigofera heterantha and Wikstroemia canescens, whereas
rare shrubs included Desmodium elegans, Onopordum
acanthium and Rubus niveus with minimum IVI values.
The herbs layer was dominated by Agrostis gigantean,
Viola odorata and Bistorta amplexicaulis while Micromeria
biflora, Rubia cordifolia and Ajuga bracteosa were the rare
herbs in region. The soil state of this association revealed
EC from 0.43 to 0.89 d/Sm, pH 6.22–6.75, OM 0.5–1.35%,
P 1.1–7.5 mg/kg, K 120–240 mg/kg and saturation 47–80%,
while its altitudinal range is 1543–2170 m.
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Table 1 Detail information of top indicator species in all the association and determinant environmental variables
S. no Indicator species
Variables
IV
Association-01 total number of quadrates ( plots) is 24
1
Taxus baccata L
Phosphorous 16.7
2
Viburnum grandiflorum Wall Phosphorous 19
3
Pteridium aquilinum L
Slope
12
Association-02 total number of quadrates ( plots) is 29
1
Ailanthus altissima
Potassium (K) 46
2
Rosa webbiana
Aspect
16.7
3
Arenaria leptoclados
Saturation
69.7
Association-03 total number of quadrates ( plots) is 42
1
Quercus dilatata
Slope
9.4
2
Cotoneaster nummularia
Stumps
31.1
3
Brachypodium sylvaticum
Saturation
22.9
Association-04 total number of quadrates ( plots) is 11
1
Olea ferruginea
pH
21
2
Myrsine Africana
Potassium
30.8
3
Prunella vulgaris
pH
30.9
P* value
0.0526
0.0328
0.0282
0.031
0.0136
0.0054
0.0340
0.0272
0.0486
0.01
0.0584
0.0438
IV Indicator Value
Olea Ferruginea, Myrsine Africana And Prunella Vulgaris
Plant association
A total of 11 stations clustered this association. The top three
indicators of this association were Olea ferruginea, Myrsine
africana and Prunella vulgaris (Table 1; Fig. 7). These species were the indicator of lower pH and higher potassium
concentration environmental variables. The dominant tree
species with highest IVI values included Quercus baloot,
Quercus semecarpifolia and Cornus macrophylla, while
the rare tree species were Acer caesium with minimum IVI.
The dominant shrubs were Parrotiopsis jacquemontiana,
Isodon rugosus and Jasminum humile with high IVI values,
whereas rare shrubs included Sorbaria tomentosa, Berberis
lycium and Spiraea species. The herb layer was dominated
by Origanum vulgare, Rostraria cristata and Viola odorata
with high IVI values while Bupleurum falcatum, Fallopia
convolvulus and Herniaria glabra were the rare herbs of the
region with low IVI values.
The soil state of this association revealed EC from 0.62 to
0.49 d/Sm, pH 6.19–7.2, OM 0.6–1.3%, P 1.8–7.8 mg/kg, K
100–220 mg/kg and saturation 49–70%, while its altitudinal
range is 1564–2000 m.
Environmental gradient
Detrended correspondence analysis (DCA) and canonical
correspondence analysis (CCA)
Detrended correspondence analysis (DCA) was performed
to clarify distribution of 142 species and 106 quadrats along
the axes of detrended gradient analysis. Association 1st is
mostly present at the top of plot. Association 3rd is at bottom while association 4th is present in between 1st and 3rd
associations. Association 2nd lies at the left side of the plot
(Fig. 8). In various plant species ordination each green pyramid in the figure show a plant species, while the distance
between them show similarity and differences index. The
CCA (bi-plot diagram) of first quadrant (on top left) indicated most of the plants were gathered under the influence
of slope, stumps, potassium, pH and electrical conductivity
(EC). While in 2nd quadrant (on bottom left) most of the
environmental variables clustered around texture and saturation of soil. Furthermore on the 3rd quadrant most of the
plants are assembled under the influence of organic matter
Fig. 4 Data attribute plots (left to right) of Indicator species of the 1st association i.e., Taxus baccata, Viburnum grandiflorum and Pteridium
aquilinum in relation to measured environmental variables
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Fig. 5 Data attribute plots of Indicator species of the second association (left to right) Ailanthus altissima, Rosa webbiana and Arenaria leptoclados under the impact of environmental variables
Fig. 6 Data attribute plots (left to right) of Quercus dilatata, Cotoneaster nummularia and Brachypodium sylvaticum (Indicator species of the
3rd association)
while on 4th quadrant the plants are mostly assembled under
the influence of altitude, aspect, phosphorous and grazing
pressure (Fig. 9).
Conservation status of Parrotiopsis jacquemontiana
Altitudinal range
Altitudinal range of P. jacquemontiana is between 1556 and
2313 m. This species is absent from south facing slopes,
rarely distributed in north-west and north-east side while on
north facing slopes it grow abundantly.
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Population estimation and geographic range
More than 75% respondents indicated that over the last
three generations, the population of P. jacquemontiana had
considerably declined (> 65%) in the wild habitat. Besides
other harmful factors, annual use of this species as fuel
is greater than its total germination. Therefore, its habitat
range has shrunk over the years. The EOO of P. jacquemontiana is 2218 km2 (< 5000 km2), while AOO is 210
km2 (< 500 km2) which was calculated using Google Earth
Path software.
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Fig. 7 Data attribute plots (left to right) of Indicator species i.e., Olea ferruginea, Myrsine africana and Prunella vulgaris in relation to different
variables
Fig. 8 Detrended Correspondence Analysis (DCA) plot showing: a distributions of 142 plant species and b four plant associations in the study
area
Status summary
Population dynamics and conservation status of P. jacquemontiana are expressed below using hierarchical alphanumeric numbering system of the said criteria and sub criteria
is as follows: ‘EN A2acd; B1B2bc (i, ii, iii)’ where ‘EN’
refers to endangered; A2 = population reduction observed,
estimated, inferred in the past where the causes of the reduction were not ceased based on (a) = direct observation; (c) = a
decline in area of occupancy, extent of occurrence and quality of habitat; (d) = actual or potential levels of exploitation, B = geographic range in the form of EOO and AOO’
B1 = extent of occurrence, B2 = area of occupancy, b = continuing decline and c = extreme fluctuation in (1) extent of
occurrence, (2) area of occupancy, (3) quality of habitat loss.
Discussion
We found a total of 142 plant species that belong to 62 families,
of which the most common were Poaceae, Rosaceae, Labiatae
and Asteraceae in Dir (upper and lower) districts. There were
21 trees (14.8%), 24 shrubs (16.9%) and 97 herbaceous (68.3%)
species. Herbaceous species had the highest cover in our study
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Fig. 9 Canonical Correspondence Analysis (CCA) diagram showing the species distribution under the influence of various environmental variables. K Potassium, EC electrical conductivity, Satu saturation, O.M organic matter, P phosphorous, GP grazing pressure
area with fewer shrubs and trees. Physiographic factors like slope,
aspect, altitudinal range and different edaphic factors affect vegetation patterns and species composition. Our results show similarities with the vegetation communities of Western Himalayan
parts of Malakand, Kashmir, Hazara, and Gilgit Baltistan regions
of Pakistan where Asteraceae, Lamiaceae, Poaceae and Rosaceae
were the most dominant families (Khan et al. 2011; Shaheen et al.
2011; Abbas et al. 2020). Similarly Asteraceae and Labiatae were
well-established and well-represented families in flora of Pakistan (Stewart 1972; Ali and Qaiser 1995). Khan et al. (2014) also
found that Poaceae and Asteraceae were the dominant plant families of Shahbaz Garhi, District Mardan, Pakistan, which strongly
support our findings. Similarly, Nazir et al (2012) reported that
Poaceae was the dominant family at the Sarsawa Hills of district Kotli Kashmir. Similar to our results Perveen and Hussain
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(2007) determined species density, frequency and cover of the
Gorakh Hills and described 74 plant species belonging 34 families. Plants play an important role in the local economy of these
regions where they provide services as fruit, food, fire wood,
timber wood, medicines, forage, etc. (Shinwari et al. 1996; Durrani 2000; Malik 2005; Mehmood et al. 2015).
Plant species distribution, association formation and their
indicator species are typically attributed to edaphic and environmental variables. Four plant associations were identified in
the study area. The Taxus baccata, Viburnum grandiflorum and
Pteridium aquilinum association characterized clay loam soil
on north and northwest aspects at altitudes between 1758 and
2239 m. The soil EC ranges between 0.44 and 0.79 d/Sm, pH
6.39–6.42, OM 0.6–0.75%, P 2.4–9.2 mg/kg, K 100–220 mg/kg,
saturation 41–68%, showing that this association was found in
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slightly acidic substrate with low organic matter. The Indicator
Species Analysis shows that the indicator species of this association are strongly correlated with high phosphorous concentrations and steep slopes. Ailanthus altissima, Rosa webbiana and
Arenaria leptoclados are the top indicator species of 2nd association. The edaphic factors of 2nd association show that these
plants can grow in soils with EC 0.36–1.06 d/Sm, pH 5.25–7.68,
OM 0.6–1.35%, P 1.5–8.1 mg/kg, K 120–240 mg/kg and saturation 40–70% at altitude between 1556–2313 m. The Indicator
Species Analysis shows that the indicator species of this association is strongly correlated with higher potassium concentrations,
west aspects and low saturation of the soil. Quercus dilatata,
Cotoneaster nummularia and Brachypodium sylvaticum are the
indicator species of 3rd association which are strongly correlated
with higher number of stumps and higher saturation of the soil.
The edaphic variable ranges between EC 0.43–0.89 d/Sm, pH
6.22–6.75, OM 0.5–1.35%, P 1.1–7.5 mg/kg, K 120–240 mg/
kg and saturation 47–80%. The altitudinal range of this association is 1543–2170 m. In 4th association the indicator species are
Olea ferruginea, Myrsine africana and Prunella vulgaris which
are strongly correlated with high pH value and high potassium
concentration. The edaphic factor ranges EC 0.39–0.9 d/Sm, pH
6.19–7.2, OM 0.6–1.3%, P 1.8–7.8 mg/kg, K 100–220 mg/kg
and saturation 49–70%. The altitudinal range is 1564–2002 m.
Other authors also reported various assemblages/associations
of plant species in relation to environmental variables (Wana
2002; Shaheen et al. 2011; Khan et al. 2012a; Nazir et al. 2012;
Shaheen et al. 2011; Ahmad et al. 2016b).
Similarly Bano et al. (2018) reported four plant communities/
associations during the eco-floristic study of Beer Hills along
the Indus River, Pakistan, which strongly support our findings.
Ahmed et al. (2006); Ramzan et al. (2016) described 24 plant
communities/associations and four monophonic definite forest
vegetation and labeled the IVI values and species composition
while studying phytosociological analysis of Himalayan forests
of Pakistan. Khan et al. (2012b) also applied similar techniques
for proper record of plant species. Borcard et al. (1992) used
canonical correspondence analysis to categorize different variables. Fourteen communities/associations were determined by
Brown and Bezuidenhout (2005) in the study of the Mountain
Zebra National park, South Africa. Overall soil pH of the study
area ranges between 5.2 and 7.7; electrical conductivity 0.4–5.0;
organic matter concentration from 0.5 to 1.35%, phosphorous
was 1.1–9.2 ppm, potassium ranges 100–240 ppm. Similarly
Khan et al. (2016b), (2017) and Shaheen et al. (2011) also found
out various plant communities or associations in relation to environmental gradients. Furthermore, the vegetation of Cholistan
desert was explored by Noureen et al. (2008) on the basis of environmental variables. Whereas, Yimer (2007) stated that soil interrupt the structure of the plant associations and amount of plant
development, ground cover, ability of natural regeneration and
extra critical influences. In our study area higher grazing pressure
was observed at lower altitude. Pennings and Silliman (2005)
reported that grazing pressure was high at lower elevation which
supports our findings. Sakya and Bania (1998) suggested that
elevation plays a key role in the community/association formation. Shank and Noorie (1950) found that atmospheric pressure
and temperature changed with increase in altitude. They also find
out that soil pH, soil nutrients, soil moisture and biotic influences
also take part in the development of plant assemblages.
Our study reveals that the population of P. jacquemontiana
has declined by more than 65% in the study area. The main
causes of reduction in its population include fuel wood, fodder
and agricultural tool handles. Due to continuous decline in population and geographic range i.e., EOO = 2218 km2 (< 5000 km2)
and AOO = 210 km2 (< 500 km2) this species falls under the
category and criteria of endangered species ‘EN A2acd; B1B2
bc (i, ii, iii). The only conservation study of P. jacquemontiana
was carried out by Ullah and Rashid (2014) in Mankial Valley
Pakistan which found that this species occurs at an altitude of
1764 m; the population of this species has reduced by more
than 80% in the Mankial Valley due to extensive use as fodder
and fuel wood. Due to this decrease in population of P. jacquemontiana it falls under the category and criteria of A (a, c) of
the endangered species which strongly supports our findings.
Some of the studies on uses of plants and the resultant human
pressures, for example a study on Mazri Palm (Abdullah et al.
2020) and another on Karakorum Range of Mountains (Abbas
et. al. 2019) adovocate about the importantce of documentation
about such species and regions and can further be strengthened
as well as coordinated at academic, research, and policy levels.
Conclusions
Cluster, two-way cluster and indicator species analyses were used
to investigate vegetation patterns and the environmental factors
that affect them. We found that that high phosphorous and potassium concentrations, elevation, aspect, slope, lower pH, electrical conductivity and soil texture are the significant environmental
variables that play an important role in determining vegetation
structure and the formation of plant associations and its indicators in our study area. The endemic Parrotiopsis jacquemontiana
is an endangered species in the study area which faces a high
risk of extinction. The main causes of its population decline and
habitat loss include its use for fuel, fodder, and making handles
for agricultural tools.
Compliance with ethical standards
Conflict of interest All the authors declared that they have no conflict
of interest.
Appendix
See Table 2.
13
13
Table 2 Results of indicator species analysis (ISA), showing all the plant species in relation to various edaphic and topographic variables, based on Monte Carlo test of significance for maximum observed indicator value (IV) of species (P ≤ 0.05) (top indicators are showed in bold font)
Botanical names
Potassium
Phosphorus
pH
Saturation
Slope
Numbers of Stump
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
4
4
4
1
1
1
1
1
1
4
4
1
1
1
1
1
1
4
1
1
1
1
1
1
1
1
4
4
4
4
4
1
1
4
1
0.112
0.117
0.885
0.537
0.614
1.000
1.000
1.000
1.000
0.000
0.117
0.916
0.675
0.839
1.000
0.637
1.000
0.687
1.000
1.000
1.000
0.204
0.518
1.000
0.595
1.000
0.002
0.279
0.003
0.116
0.001
1.000
1.000
0.782
1.000
180
180
180
240
180
200
160
160
100
140
180
220
180
120
200
140
200
100
200
220
140
240
200
180
200
200
120
180
180
120
180
240
200
100
220
0.612
0.614
0.810
0.239
0.225
0.273
0.426
0.927
0.073
0.246
0.614
0.516
0.407
0.977
0.272
0.551
0.668
0.187
0.273
0.112
0.795
0.300
0.410
0.622
0.618
0.473
0.397
0.285
0.828
1.000
0.892
0.029
0.676
0.056
0.361
5
5
1
3
6
1
6
7
7
3
5
6
7
4
1
5
5
5
4
3
2
4
1
1
2
2
5
6
5
4
7
4
6
7
7
0.262
0.252
0.037
0.432
0.754
0.370
0.143
0.094
0.007
0.119
0.252
0.035
0.343
0.939
0.371
0.082
0.550
0.569
0.546
0.760
1.000
0.081
0.049
0.380
0.279
1.000
0.597
0.962
0.268
0.550
0.785
0.817
0.435
0.529
0.059
7
5
6
6
6
6
6
6
6
5
5
6
6
6
6
6
6
5
6
6
6
6
6
6
6
6
7
5
5
6
5
6
6
6
6
0.025
0.068
1.000
0.974
1.000
1.000
1.000
1.000
1.000
0.251
0.068
1.000
1.000
1.000
1.000
0.911
1.000
0.395
1.000
1.000
1.000
0.651
0.278
1.000
1.000
1.000
0.099
0.205
0.291
1.000
0.009
1.000
1.000
1.000
1.000
62
50
72
65
49
70
50
50
66
42
46
65
47
60
71
58
58
43
50
65
70
62
71
70
50
65
42
49
62
70
42
68
55
49
48
0.104
0.336
0.563
0.661
0.326
0.792
0.538
0.925
0.803
0.005
0.275
0.244
0.285
0.877
0.108
0.839
0.222
0.550
0.563
0.790
0.792
0.708
0.095
0.792
0.723
1.000
0.215
0.346
0.576
0.785
0.039
0.499
0.854
0.955
0.378
3
2
3
3
2
2
3
2
2
3
2
3
3
2
2
3
2
3
2
2
3
2
2
2
3
3
3
2
3
3
2
2
3
3
3
1.000
0.421
0.657
0.133
0.445
0.425
1.000
1.000
0.074
0.373
0.421
0.137
0.067
1.000
0.427
0.494
0.466
0.283
0.432
0.415
1.000
0.972
0.150
0.429
0.113
0.515
0.085
0.951
0.236
1.000
0.184
0.178
0.262
0.566
0.499
0
0
1
3
1
2
3
1
0
0
0
3
7
1
1
2
0
2
7
0
0
3
3
1
7
1
0
1
2
0
0
0
0
2
2
1.000
1.000
0.529
0.895
0.334
0.258
0.105
1.000
0.435
0.212
1.000
0.661
0.077
0.558
0.574
0.851
0.867
0.473
0.029
1.000
1.000
1.000
0.759
0.568
0.171
0.870
0.330
0.513
0.813
1.000
0.521
0.496
0.884
0.740
0.394
8.3
8.3
4.6
9.7
6.5
1.1
1.1
2.2
3.2
41.7
8.3
4.3
6.5
4.3
1.1
9.6
3.2
5.8
1.1
1.1
1.1
19.4
22.4
1.1
7.5
2.2
41.7
16.7
24.4
8.3
24.7
2.2
3.2
4.9
2.2
5.3
5.3
4.4
17.1
13.9
6.2
5.9
2.8
38.7
11.6
5.3
5.9
9
2
6.2
9
3.2
21.6
6.2
11.1
5
15.5
15.2
5.3
6.3
3.9
7.2
17.5
3.7
4.8
2.8
45.5
3.2
31.3
5.7
9.1
9.1
18.4
7.8
4.4
7.7
11.1
12.6
28.1
11.1
9.1
17
8.4
3
7.7
15.6
5.3
6.2
5.9
4.3
4
17.5
22.9
7.7
10
2.3
5.2
4.5
9.8
5.9
4
3.5
6
6.1
14.7
33.3
25
7.1
9.1
6.1
1
1
2
3
19.9
25
4
6.1
4
1
12.1
3
14.4
1
1
1
18.2
35.4
1
7.1
2
26.4
24.6
12.7
1
45.1
2
3
5.1
2
50
22.3
13.3
11.5
21.3
8.3
11.1
5.6
8
69.7
19.6
25
20.6
6
50
8.3
28.3
13.6
11.1
8.3
8.3
10.4
28.1
8.3
10.5
4.6
26.6
16.6
12.5
8.3
44.8
13.9
6
5
17.9
1.7
2.2
4.6
10.2
5
2.2
1.7
1.1
6.7
5
2.2
6.7
10
2.3
2.2
7.7
3.2
7.6
2.2
2.2
1.7
8.7
25
2.2
9.4
3.3
8.3
7.9
6.2
1.7
6.2
4.4
5
3.9
3.3
2.2
2.2
6.7
4.4
8
5.9
12.5
1.6
6.7
11.1
2.2
3.9
19.3
4.4
3.1
5.3
2.6
7.7
33.3
2.2
2.2
4
12.5
3.1
14
2.1
8.1
10.7
3.6
2.2
5.3
4.4
2.8
4.3
4.9
Tropical Ecology
Achyranthes aspera Linn
Acrachne racemosa (Heyne ex Roem. and Schult.)
Adiantum caudatum
Adiantum venustum
Agrostis gigantea Roth, Tent
Ajuga bracteosa Wall
Alliaria petiolata (M. Bieb.)
Androsace rotundifolia Hardwicke
Arabidopsis wallichii (Hook. f. and Thoms.)
Arenaria leptoclados (Reichb.)
Arenaria orbiculata Royle ex Edgew. and Hook.f
Arisaema flavum (Forsk.)
Arisaema jacquemontii Blume, Rumphia
Artemisia vulgaris Linnaeus
Arthraxon prionodes (Steud.)
Asplenium adiantum-nigrum
Asplenium ceterach
Asplenium trichomanes
Asplenium tricorn
Athyrium sp.
Bergenia ciliata (Haw.)
Bistorta amplexicaulis (D. Don)
Brachypodium sylvaticum (Huds.) P.Beauv
Bromus japonicus Thunb
Bupleurum falcatum Linn
Bupleurum lanceolatum Wall
Cannabis sativa Linn
Carex sp.
Chenopodium album Linnaeus
Chenopodium botrys L
Chenopodium murale L
Conyza canadensis
Cymbopogon jwarancusa (Jones)
Cymbopogon pospischilii (K. Schum.)
Cynoglossum glochidiatum Wall. ex Benth
Aspect
Botanical names
13
Cynoglossum lanceolatum Forssk
Cystopteris fragilis (L.) Bernh
Desmostachya bipinnata (Linn.)
Dioscorea deltoidea Wall
Dryopteris stewartii Fraser-Jenk
Duchesnea indica (Andrews)
Dysphania botrys (L.)
Euphorbia indica Lam
Euphorbia wallichii Hook
Fallopia convolvulus (L.)
Fragaria vesca L
Galium aparine L
Galium tricornutum Dandy
Geranium wallichianum D. Don ex sweet
Girardinia diversifolia (Forssk.)
Girardinia palmata (Forssk.) Gaudich
Herniaria glabra L
Impatiens brachycentra Kar. and Kir
Lactuca dissecta D.Don
Lamium album L
Lecanthus peduncularis (Royle) Wedd
Lespedeza juncea (Linn.) Pers
Melissa officinalis L
Micromeria biflora Benth
Oenothera javanica
Onychium contiguum C.Hope
Oplismenus hirtellus (L.) P.Beauv
Oplismenus undulatifolius (Ard.)
Origanum vulgare L
Oxalis corniculata L
Perilla frutescens (L.) Britton
Pimpinella diversifolia DC., Prodr
Piptatherum laterale (Regel) Nevski
Plantago lanceolata L
Plantago major L
Aspect
Potassium
Phosphorus
pH
Saturation
Slope
Numbers of Stump
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
1
1
1
1
1
1
1
1
1
1
1
4
1
1
1
1
1
1
4
4
1
1
1
1
1
1
1
1
1
4
1
1
4
4
4
1.000
1.000
1.000
1.000
0.918
1.000
1.000
1.000
1.000
1.000
0.265
1.000
1.000
1.000
1.000
1.000
1.000
0.749
0.000
0.103
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.597
0.007
0.428
0.702
0.007
0.117
0.117
200
180
160
160
220
180
200
120
200
140
160
140
120
180
140
120
180
100
140
140
180
220
160
200
180
140
220
220
100
240
140
200
240
220
180
0.273
0.155
0.437
0.193
0.546
0.628
0.273
0.956
0.658
0.791
0.618
0.597
0.252
0.197
0.274
1.000
0.926
0.893
0.600
0.795
0.621
0.151
0.437
0.101
0.605
0.808
0.154
0.307
0.159
0.351
0.995
0.211
0.582
0.961
0.614
4
4
2
5
6
6
4
5
2
2
2
5
5
6
6
4
6
6
3
3
7
1
2
1
4
2
6
3
7
6
6
1
4
5
5
0.546
0.637
1.000
0.461
0.455
0.143
0.546
0.462
0.680
1.000
0.887
0.128
0.597
0.331
0.426
0.534
0.260
0.392
0.988
0.762
0.053
0.153
1.000
0.618
0.539
1.000
0.176
0.597
0.146
0.334
0.881
0.033
0.077
0.418
0.252
6
6
6
6
6
6
6
6
6
7
6
6
6
6
6
6
7
5
5
6
6
6
6
6
6
6
6
6
7
5
6
6
5
5
5
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.030
0.251
1.000
1.000
1.000
1.000
1.000
0.054
0.526
0.030
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.212
0.904
0.746
1.000
0.078
0.117
0.068
50
55
60
50
58
55
50
70
60
50
71
60
60
55
50
62
50
43
42
70
48
72
60
72
55
58
65
49
49
42
49
70
47
65
68
0.563
0.673
1.000
0.851
0.839
0.559
0.563
0.681
1.000
0.555
0.303
0.990
0.829
0.561
0.916
0.090
0.555
0.074
0.046
0.802
0.218
0.976
1.000
0.497
0.545
0.125
0.856
0.911
0.464
0.125
0.506
0.483
0.004
0.864
0.463
2
2
2
3
3
2
2
3
3
3
3
3
3
2
3
3
3
3
2
3
3
2
2
2
2
2
3
2
2
2
3
2
2
2
2
0.432
0.766
0.432
0.259
0.394
0.432
0.432
0.619
0.263
1.000
0.803
0.056
0.255
1.000
0.251
1.000
0.515
0.095
0.947
1.000
1.000
0.034
0.432
0.179
0.448
0.429
1.000
1.000
0.123
0.319
0.273
0.104
0.953
0.073
0.421
7
3
2
0
3
2
7
0
1
0
2
0
0
2
0
0
2
3
0
0
2
1
2
3
3
0
1
0
1
2
3
3
7
0
0
0.029
0.121
0.265
0.424
0.462
0.271
0.029
0.705
0.497
1.000
0.824
0.985
0.901
0.589
0.856
1.000
0.636
0.753
0.374
1.000
0.279
0.720
0.265
0.155
0.114
1.000
0.698
0.823
0.617
0.628
0.137
0.198
0.018
0.685
1.000
1.1
2.2
1.1
3.2
4.3
1.1
1.1
3.2
3.2
1.1
27.5
3.6
3.2
3.2
3.2
1.1
2.2
13.7
40.4
8.3
1.1
3.2
1.1
2.2
1.1
1.1
2.2
3.2
13.3
31.2
13.2
5.4
60.6
7.8
8.3
6.2
10.5
5.9
9.3
5.5
5.3
6.2
2.4
3.8
5
12.1
6.1
8.1
9.7
7.2
4.8
2.7
7.6
6.6
5
5.3
19.3
5.9
12.5
5.3
5
10.9
8.1
21.5
14.5
3.9
13.9
15.8
2.5
5.3
5.9
5
4
5.9
5.9
11.1
5.9
6.1
5.1
4
8.8
12.7
5.1
7.3
5.8
5.9
8.2
11
2.5
4.3
16.7
11.8
4
5.1
5.9
4
10.8
5.3
14.5
10
5.4
18.5
23.4
6.3
9.1
1
2
1
3
4
1
1
3
3
33.3
37.4
6.1
3
3
3
1
32.4
20.9
45
1
1
4
1
2
1
1
2
3
27.4
13.4
16.2
5.1
56.8
24.2
25
11.1
9
4.8
6.5
7.9
11.1
11.1
9.9
2.5
11.1
17
3.9
6.9
10.4
5.1
50
11.1
28.9
34.3
8.3
25
4.8
4.8
14.1
11.1
33.3
7.2
5.8
13.2
26
13.3
15.2
35.4
5.8
13.2
2.2
2
2.2
5
4.5
2.2
2.2
2.6
5
1.7
18.9
10
5
1.4
5
1.7
3.3
20.2
4
1.7
1.7
8.9
2.2
4.4
2.2
2.2
1.6
1.4
17.6
10.9
12.5
7.9
28.9
6.7
2.2
33.3
12.1
5.9
6.7
6.8
5.9
33.3
3.4
5.5
2.2
11.6
2.7
2.5
4.2
2.8
2.2
4.3
9.7
10.2
2.2
5.9
4.5
5.9
10.6
12.5
2.2
3
2.7
10.5
8.7
18.7
13.1
44.2
3.7
2.2
Tropical Ecology
Table 2 (continued)
13
Table 2 (continued)
Botanical names
Potassium
Phosphorus
pH
Saturation
Slope
Numbers of Stump
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
1
1
1
1
1
4
4
1
1
1
4
4
1
1
4
1
1
1
1
1
1
4
1
1
4
1
1
1
1
4
1
1
4
4
1
1.000
1.000
1.000
1.000
1.000
0.307
0.110
0.857
0.586
0.921
0.876
0.011
1.000
1.000
0.117
0.856
0.839
0.571
1.000
1.000
1.000
0.116
0.573
0.160
0.019
1.000
1.000
0.400
1.000
0.017
1.000
1.000
1.000
0.043
0.183
220
140
160
200
160
120
140
220
160
180
180
140
220
180
160
160
240
180
180
200
160
120
240
240
180
200
160
220
140
100
180
160
120
200
100
0.408
0.805
0.520
0.282
0.426
0.717
0.798
0.259
0.377
0.179
0.751
0.894
0.124
0.172
0.436
0.416
0.111
0.303
0.615
0.267
0.431
1.000
0.040
0.886
0.676
0.276
0.437
0.053
0.178
0.135
0.770
0.434
0.880
0.268
0.398
7
2
5
1
6
4
3
7
7
7
2
4
6
6
4
4
1
1
7
6
2
4
5
7
7
1
2
5
2
7
4
2
2
1
7
0.309
0.905
0.608
0.372
0.143
0.062
0.763
0.546
0.461
0.003
0.478
0.892
0.148
0.399
0.538
0.348
0.717
0.624
0.052
0.142
1.000
0.550
0.826
0.238
0.386
0.372
1.000
0.797
0.333
0.080
0.750
1.000
0.941
0.103
0.125
6
6
6
6
6
6
6
7
6
6
7
6
6
5
6
6
6
6
6
6
6
6
6
6
5
6
6
6
6
5
6
6
6
5
5
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.044
1.000
1.000
0.043
1.000
1.000
0.123
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.535
0.786
1.000
1.000
0.979
1.000
0.191
1.000
1.000
1.000
0.254
0.636
72
49
60
71
50
50
55
58
47
48
50
55
65
55
55
68
72
47
48
66
68
70
66
48
47
70
60
66
55
49
65
72
62
72
71
0.527
0.864
1.000
0.104
0.538
0.915
0.548
0.583
0.347
0.111
0.905
0.808
0.792
0.747
0.548
0.569
0.687
0.321
0.205
0.307
0.210
0.785
0.807
0.198
0.470
0.786
1.000
0.841
0.776
0.899
0.868
0.310
0.361
0.116
0.952
2
3
3
3
3
2
3
2
3
3
3
3
3
2
3
2
2
2
3
2
6.7
3.3
3.3
1.7
1.7
2.4
1.7
3.7
11.7
6.7
6.7
3.3
1.7
1.8
1.7
4.9
8.9
14.2
1.7
2.2
0.070
0.501
0.509
1.000
1.000
0.867
1.000
0.811
0.028
0.142
0.220
0.509
1.000
0.747
1.000
0.311
0.034
0.005
1.000
0.422
2
2
3
3
3
7
0
3
7
2
0
0
1
2
1
7
0
7
1
1
3.3
2.5
9.5
12.5
12.5
30.3
2.2
4.8
14.2
8.1
3.9
4.4
3.1
11.8
3.1
48.5
8.9
21.1
3.1
3.1
0.727
0.828
0.194
0.105
0.105
0.006
1.000
0.771
0.193
0.337
0.894
0.493
0.570
0.127
0.572
0.003
0.298
0.075
0.570
0.567
3
2
2
3
2
2
3
3
2
2
3
3
2
3
1.7
4.1
15.8
50.5
2.2
2.2
6.3
3.3
19.1
4.4
1.7
7.3
44.6
12.4
1.000
0.907
0.176
0.001
0.438
0.432
0.757
0.503
0.264
0.184
1.000
0.316
0.055
0.461
0
3
1
7
1
2
0
0
0
7
1
3
7
2
2.2
6.1
22.7
39.8
3.1
5.9
7.3
4.4
13.6
31.1
3.1
12.1
18.4
8.6
1.000
0.638
0.105
0.024
0.566
0.265
0.668
0.614
0.466
0.027
0.582
0.315
0.785
0.762
3.2
2.2
2.2
1.1
1.1
6.2
8.3
5.4
7.5
4.3
5
16.7
1.1
2.2
8.3
5.4
4.3
8.6
1.1
1.1
1.1
8.3
8.6
22.6
53.9
1.1
1.1
11.8
2.2
39
2.2
1.1
4.5
53.1
20.4
6.5
3.8
3.8
6.2
5.9
4.5
5
12.3
10.2
14
4.1
2.8
11.1
10.5
5.9
6.4
21.3
14.7
5.3
6.2
5.9
4.8
34.8
6.9
14.2
6.2
5.9
27.9
10
22.3
3.6
5.9
4
20.5
12.6
7.2
3.1
5.3
7.7
11.1
14.1
4.3
5.9
7.1
28.4
6.4
3.2
11.1
7.4
5.9
7.5
4.7
5.7
16.7
11.1
4
5.9
4.3
12.5
16.2
7.7
4
4.9
8
18.2
3.9
4
3.4
21.3
15.6
3
2
2
1
1
4
1
30.9
7.1
4
30.6
2
1
22.3
1
5.1
4
8.1
1
1
1
1
8.1
21.2
27
1
1
11.1
2
34.9
2
1
9.1
43.9
18.1
13.5
6.1
4.8
50
11.1
5.1
11.1
12.1
20.5
34.6
6.1
6.8
8.3
8.3
11.1
12.9
10.2
19.1
25
20
25
8.3
8.5
19
13.2
8.3
4.8
8.9
7.4
7.7
6
20
17.4
17.3
6.6
Tropical Ecology
Polygala abyssinica R.Br. ex Fresen
Polygonatum cirrhifolium (Wall.) Royle
Polygonatum multiflorum (L.) All
Polygonatum odoratum (Mill.) Druce
Polygonatum verticillatum (L.) All
Polygonum paronychioides C.A. Mey
Potentilla anserina L
Prunella vulgaris L
Pteridium aquilinum (L.) Kuhn
Ranunculus sceleratus L
Rostraria cristata (L.) Tzvelev
Rosularia rosulata (Edgew.)
Rubia cordifolia L
Rumex dentatus L
Scrophularia scabiosifolia Benth
Stellaria media (L.) Vill
Tagetes minuta L
Taraxacum officinale (L.)
Thalictrum foliolosum DC
Trifolium repens L
Trigonella sp.
Urtica dioica L
Valeriana jatamansi Jones
Verbascum thapsus L
Viola odorata L
Viola rupestris F.W.Schmidt
Sida cordata (Burm.f.) Borss.Waalk
Berberis lycium Royle in Trans. Linn
Clematis grata Wall
Cotoneaster microphyllus
Cotoneaster nummularia
Desmodium elegans DC
Hedera nepalensis K. Koch, Hort. Dendrol
Indigofera heterantha Wall
Isodon rugosus (Wall. ex Benth.)
Aspect
Botanical names
13
Jasminum humile Linn
Lonicera quinquelocularis Hardwicke
Myrsine africana Linn
Onopordum acanthium L
Parrotiopsis jacquemontiana (Dcne.)
Prunus tomentosa Thunb
Rosa brunonii Lindl
Rubus niveus Thunb
Sarcococca saligna (D.Don)
Sorbaria tomentosa (Lindl.) Rehder
Spiraea sp.
Strobilanthes glutinosus (Nees) Bremek
Rosa webbiana
Viburnum cotinifolium D. Don
Viburnum grandiflorum Wall. ex DC
Wikstroemia canescens Wall. ex Meisn
Acer caesium Wall
Ailanthus altissima (Mill.)
Cedrus deodara (Roxb. ex D. Don)
Celtis australis Willd
Cornus macrophylla Wall
Diospyros kaki Linn
Diospyros lotus Linn
Ficus carica L
Juglans regia Linn
Olea ferruginea Royle
Pinus roxburghii Sarg
Pinus wallichiana A.B.Jacks
Platanus orientalis L
Populus ciliata Wall. ex Royle
Quercus baloot Griff
Quercus dilatata Royle, Illust
Quercus incana
Quercus semecarpifolia Sm
Rhamnus virgatus Roxb
Aspect
Potassium
Phosphorus
pH
Saturation
Slope
Numbers of Stump
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
Max IV
P*
1
1
4
1
1
4
1
1
1
1
1
1
4
1
1
1
4
1
4
1
1
1
1
1
4
1
1
1
4
1
1
1
1
1
1
0.336
0.604
0.950
1.000
0.001
0.005
1.000
1.000
0.581
0.918
1.000
1.000
0.013
0.402
0.459
0.227
0.386
1.000
0.072
1.000
0.595
1.000
1.000
1.000
0.212
1.000
1.000
0.106
0.110
1.000
0.972
0.817
1.000
0.228
1.000
100
180
100
140
240
160
240
160
140
180
120
120
160
180
240
220
120
240
160
140
180
200
200
160
200
160
220
160
140
180
140
160
120
200
120
0.122
0.798
0.058
0.803
0.490
0.811
0.016
0.440
0.480
0.843
0.423
1.000
0.559
0.784
0.205
0.963
0.705
0.031
0.189
0.801
0.323
0.272
0.167
0.426
0.499
0.439
0.438
0.377
0.798
0.615
0.376
0.394
0.985
0.385
1.000
7
1
7
3
7
2
4
2
2
4
3
5
4
6
7
5
6
4
7
1
1
1
1
6
1
5
6
7
3
7
1
5
6
4
2
0.102
0.181
0.190
0.756
0.011
0.815
1.000
1.000
0.259
0.041
0.522
0.239
0.789
0.313
0.033
0.527
0.683
0.892
0.073
0.369
0.888
0.371
0.053
0.143
0.693
0.252
0.427
0.267
0.763
0.052
0.046
0.146
0.229
0.020
1.000
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
7
6
5
6
6
6
6
6
6
6
6
6
6
6
7
5
6
6
6
0.879
1.000
1.000
1.000
0.489
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.704
0.238
1.000
0.136
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.454
1.000
1.000
0.486
1.000
1.000
0.676
1.000
49
72
49
55
71
49
62
68
58
62
60
60
42
47
48
47
50
48
47
70
58
71
71
50
71
60
65
71
55
48
72
70
58
72
60
0.764
0.122
0.971
0.557
0.295
0.901
0.094
0.198
0.403
0.101
0.871
0.944
0.152
0.325
0.424
0.711
0.990
0.337
0.114
0.779
0.368
0.108
0.133
0.538
0.134
1.000
0.990
0.445
0.548
0.205
0.446
0.892
0.419
0.978
1.000
2
2
2
3
2
2
3
2
3
3
2
3
2
2
3
3
3
2
3
2
2
2
2
3
2
3
3
3
3
3
3
2
3
2
3
0.637
0.379
0.722
1.000
0.005
0.127
1.000
0.426
0.791
0.470
0.468
1.000
1.000
0.977
0.634
0.031
0.632
0.179
0.184
0.448
0.439
0.427
0.071
1.000
1.000
1.000
0.136
0.004
1.000
1.000
0.917
0.034
0.620
0.011
1.000
0
1
0
2
3
7
0
0
0
3
0
1
1
1
7
7
3
1
7
1
1
1
2
3
1
0
1
3
0
1
1
1
1
7
0
0.556
0.302
0.325
0.275
0.845
0.169
1.000
1.000
0.275
0.392
0.749
0.725
0.865
0.630
0.338
0.634
0.436
1.000
0.069
0.572
0.754
0.574
0.722
0.105
1.000
1.000
0.782
0.037
1.000
0.570
0.790
0.692
0.952
0.020
1.000
14
7.5
4.3
1.1
67
23.6
3.2
1.1
7.5
4.3
3.2
2.2
16.7
11.8
10.8
18.3
6
2.2
17.9
1.1
8.6
1.1
3.2
1.1
7.4
1.1
4.3
25.8
8.3
1.1
13.6
10.5
3.2
18.3
1.1
27.1
4.3
30.8
5
17.2
3.8
41.8
5.9
7.5
2.8
7.6
4
3.6
5.8
22.1
5.1
4.6
45.5
22.2
5
14.3
6.2
10.1
5.9
3.5
5.9
5.9
14.6
5
5.3
14.9
13.1
2.5
13.4
4.8
14.8
10.6
11.3
4.3
24.2
3.8
2.4
4
9.4
16.8
6.1
8.8
3.8
9.5
19
8.4
4.1
3.4
17.6
7.7
3.8
7.7
13.6
11.1
4.9
9.1
6.3
13.3
4.3
16.7
21
14.6
8.7
24
4
13.1
7.1
8.1
1
41.1
5.1
3
1
7.1
4
3
2
2
11.1
10.1
17.2
18.4
2
26.8
1
8.1
1
3
1
2
1
4
24.2
1
1
23.4
8.5
3
17.2
1
9.5
27.3
4.9
11.1
11.9
6.4
37.5
25
16.2
34.9
6.5
4.2
38.3
18.3
15.8
10.4
3.5
18.7
28.8
8.3
17.2
50
35.3
11.1
40.9
4.8
3.2
15
11.1
25
13.1
7.5
16.7
5.5
4.8
7.9
6.1
5
1.7
59.5
7.8
2
2.2
4.2
3.8
3.9
1.5
1.1
5.3
6.4
18.9
3.5
4.4
10.7
2.2
6.2
2.2
6.7
1.7
1.3
1.7
6.7
27.7
1.7
1.7
14.8
9.4
2.7
20.5
1.7
9.4
10.9
10.4
5.9
22.8
13.3
2.6
2.2
10.9
7.3
3.5
3
2.1
7.8
12.2
9.4
7.5
1.8
22.6
3.1
5
3.1
3.4
12.5
1.8
2.2
3.7
37.6
2.2
3.1
10.7
9.3
2
43.7
2.2
Tropical Ecology
Table 2 (continued)
13
0.800 4
0.621 7
5.9 0.540 6
16.7 0.053 6
1
1
1.000 48
1.000 48
25
25
0.208 2
0.218 3
2.2
1.7
0.425 0
1.000 2
2.2
5.9
1.000
0.279
References
P* probability value, Max maximum group value, IV indicator value
5
5.3
1.000 140
1.000 180
1.1
1.1
1
1
Salix tetrasperma Roxb
Taxus baccata L
P*
Max IV
Max IV
Max IV
Max IV
Max IV
Max IV
Max IV
Botanical names
Table 2 (continued)
P*
Potassium
Aspect
P*
Phosphorus
P*
pH
P*
Saturation
P*
Slope
P*
Numbers of Stump
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