Environmental determinants of plant

associations and evaluation of the

conservation status of Parrotiopsis
jacquemontiana in Dir, the Hindu Kush
Range of Mountains
Fazal Manan, Shujaul Mulk Khan,
Zeeshan Ahmad, Saqib Kamran, Zahoor
Ul Haq, Fatima Abid, Majid Iqbal &
Abdullah
Tropical Ecology

ISSN 0564-3295

Trop Ecol
DOI 10.1007/s42965-020-00109-2

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Tropical Ecology International Society
https://doi.org/10.1007/s42965-020-00109-2 for Tropical Ecology

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   · Zeeshan Ahmad1 · Saqib Kamran1 · Zahoor Ul Haq1 · Fatima Abid1 ·
Majid Iqbal1 · Abdullah1

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 Eura-
sia 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 meth-
ods 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. Par-
rotiopsis 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 effects of environmental gradients on vegetation or whole

Plant species represent the hierarchical expression of vegeta-
tion 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 commu-
nities/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
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 com-
plexity (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 clas-
sification 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 fac-
tors (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; Rah-
man et al. 2020).
Poorly planned urbanization, deforestation and overex-
ploitation of natural resources are occurring at an alarm-
ing 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 wild-
life. A total of 1572 genera and 5521 species of flower-
ing 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 threat-
ened flowering plants were 580–650 (i.e. 12% of the total
flora). However, this estimation was not based on any cri-
teria. 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 Hima-
layan 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 popula-
tion size, extent of occurrence and area of occupancy was
not obtained).
Parrotiopsis jacquemontiana is an important endemic
plant of the Western Himalayan floristic province (Takhta-
jan 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). Geo-
graphically, 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 ­k m 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-felspar-
biotite, rocks of diorite group and a chain of aplite-peg-
matite-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 Par-
rotiopsis 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 estab-
lished 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 alco-
hol 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 speci-
mens 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. Graz-
ing 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). ecological amplitude and calculated historical distribution
Samples were placed in polythene bags and tagged with per- was compared with IUCN criteria for threatened categories
manent markers. Larger soil particles were removed by siev- Version 3.1 (IUCN, 2001) for evaluation of the conservation
ing and the remaining samples were shade dried (Gee and status of P. jacquemontiana. The plant species were then cat-
Bauder 1986; Khan et al. 2017). Physicochemical analysis egorized into threatened categories (IUCN 2001).
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 Results
through filter paper. For determination of pH the electrodes
of pH meter was dipped into the prepared sample and reading A total of 142 plant species were collected belonging to 62
was noted directly (Jackson 1963). An electrical conductivity families, including 21 (14.8%) trees, 24 (16.9%) shrubs,
(EC) meter was used for determination of EC (Jackson 1963; and 97 (68.3%) herbs. The most dominant families were
Hussain et al. 1999; Wilson and Bayley 2012). Soil texture Poaceae, Rosaceae, Labiatae and Asteraceae with 12, 11,
was analyzed via hydrometer (Sarir et al. 2006; Bergeron et al. 8 and 6 species, respectively.
2013). Organic matter concentration was determined by stand-
ardized solution of ­FeSO4 and ­K2Cr2O7 (Nelson and Sommers
Cluster analysis and two way cluster analysis
1996), whereas for the determination of P and K (Soltanpour
1991) method was used for each sample.
Cluster analysis classified all transects along with associ-
ated quadrats into four major associations (Fig. 2). Two
way cluster analysis further realized distribution of plant
Data analyses
species at each station that can be clearly seen in two main
branches of cluster dendrogram (Fig.  3). The detailed
All vegetation and environmental variables data were ana-
descriptions of each association are as follows:-
lyzed to assess the relationships between plant species com-
position and ecological factors using PCORD (version 5)
Taxus baccata, Viburnum grandiflorum and Pteridium
and CANOCO (version 4.5). PCORD was used to classify
aquilinum plant association
plants into different associations through cluster analysis
(CA), two way cluster analysis (TWCA) and its associated
Cluster and two way cluster analysis clustered a total of 24
indicators via indicator species analysis (ISA) (Lepš and
quadrats/stations in this association. The association name
Šmilauer 2003). Monte Carlo tests were used for statistical
was given based on indicator species analysis (ISA). The top
significance after determination of Indicator Values of each
three indicator species of this association were Taxus baccata,
indicator. A threshold level of Indicator value of 25% with
Viburnum grandiflorum and Pteridium aquilinum (Table 1;
95% significance (P value ≤ 0.05) was considered as cut-off
Fig. 4). These were indicators of higher phosphorous con-
for identifying indicators and identified indicator species
centration and steep slope environmental variables (Appen-
(IS) were used for naming the plant associations (Braak and
dix Table 2). Dominant trees with highest Importance Values
Prentice 1988; Dufrêne and Legendre 1997). Species area
Index (IVI) of this association were Pinus wallichiana, Cedrus
curves (SAC) were constructed using Sorenson distance
deodara and Quercus baloot, while the rare tree species were
measure to assess sample size adequacy. Canonical corre-
Acer caesium, Ficus carica and Pinus roxburghii with mini-
spondence analysis (CCA) was used for ordination analysis
mum IVI in the region. Dominant shrubs were Sarcococca
in estimate effects of environmental variables on plant spe-
saligna and Wikstroemia canescens, however, Indigofera
cies composition, distribution pattern and abundance.
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
IUCN Red list Criteria
with high IVI values while Polygonatum cirrhifolium, Alliaria
petiolata and Artemisia vulgaris were the rare herbs of the
Different human and natural factors were recorded with refer-
region (Appendix Table 2).IV = Indicator Value
ence to their impact on P. jacquemontiana population dynam-
The plants of this association were associated with
ics while interviewing local inhabitants in the study area,
clay loam soil and grew on north and northwest aspects
as guided by the IUCN criterion (A–E). Google Earth Path
at altitudes of 1758–2239 m. Electrical conductivity (EC)
software (V 1.4.6) was used for the calculation of extent of
ranged from 0.44 to 0.79 d/Sm, pH 6.39–6.42, organic
occurrence (EOO) and area of occupancy (AOO) available
mater 0.6–0.75%, phosphorus (P) 2.4–9.2 mg/kg, potas-
on the IUCN official website (Khan et al. 2016a). The dis-
sium (K) 100–220 mg/kg and soil saturation 41–68%.
tribution and number of plants scored with reference to their

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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 macro-
phylla, 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 cotini-
folium 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 scabiosi-
folia 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 Brachypo-
dium 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 associa- The dominant shrubs were Parrotiopsis jacquemontiana,
tion and determinant environmental variables Isodon rugosus and Jasminum humile with high IVI values,
S. no Indicator species Variables IV P* value whereas rare shrubs included Sorbaria tomentosa, Berberis
lycium and Spiraea species. The herb layer was dominated
Association-01 total number of quadrates ( plots) is 24
by Origanum vulgare, Rostraria cristata and Viola odorata
 1 Taxus baccata L Phosphorous 16.7 0.0526
with high IVI values while Bupleurum falcatum, Fallopia
 2 Viburnum grandiflorum Wall Phosphorous 19 0.0328
convolvulus and Herniaria glabra were the rare herbs of the
 3 Pteridium aquilinum L Slope 12 0.0282
region with low IVI values.
Association-02 total number of quadrates ( plots) is 29
The soil state of this association revealed EC from 0.62 to
 1 Ailanthus altissima Potassium (K) 46 0.031
0.49 d/Sm, pH 6.19–7.2, OM 0.6–1.3%, P 1.8–7.8 mg/kg, K
 2 Rosa webbiana Aspect 16.7 0.0136
100–220 mg/kg and saturation 49–70%, while its altitudinal
 3 Arenaria leptoclados Saturation 69.7 0.0054
range is 1564–2000 m.
Association-03 total number of quadrates ( plots) is 42
 1 Quercus dilatata Slope 9.4 0.0340
Environmental gradient
 2 Cotoneaster nummularia Stumps 31.1 0.0272
 3 Brachypodium sylvaticum Saturation 22.9 0.0486
Detrended correspondence analysis (DCA) and canonical
Association-04 total number of quadrates ( plots) is 11
correspondence analysis (CCA)
 1 Olea ferruginea pH 21 0.01
 2 Myrsine Africana Potassium 30.8 0.0584
Detrended correspondence analysis (DCA) was performed
 3 Prunella vulgaris pH 30.9 0.0438
to clarify distribution of 142 species and 106 quadrats along
IV Indicator Value the axes of detrended gradient analysis. Association 1st is
mostly present at the top of plot. Association 3rd is at bot-
tom while association 4th is present in between 1st and 3rd
Olea Ferruginea, Myrsine Africana And Prunella Vulgaris associations. Association 2nd lies at the left side of the plot
Plant association (Fig. 8). In various plant species ordination each green pyra-
mid in the figure show a plant species, while the distance
A total of 11 stations clustered this association. The top three between them show similarity and differences index. The
indicators of this association were Olea ferruginea, Myrsine CCA (bi-plot diagram) of first quadrant (on top left) indi-
africana and Prunella vulgaris (Table 1; Fig. 7). These spe- cated most of the plants were gathered under the influence
cies were the indicator of lower pH and higher potassium of slope, stumps, potassium, pH and electrical conductivity
concentration environmental variables. The dominant tree (EC). While in 2nd quadrant (on bottom left) most of the
species with highest IVI values included Quercus baloot, environmental variables clustered around texture and satu-
Quercus semecarpifolia and Cornus macrophylla, while ration of soil. Furthermore on the 3rd quadrant most of the
the rare tree species were Acer caesium with minimum IVI. 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 lepto-
clados 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.
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. jacque-
montiana is 2218 ­km2 (< 5000 ­km2), while AOO is 210
­km2 (< 500 k­ m2) 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 B1 = extent of occurrence, B2 = area of occupancy, b = con-

Population dynamics and conservation status of P. jacque-
montiana are expressed below using hierarchical alphanu-
meric 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 reduc-
tion were not ceased based on (a) = direct observation; (c) = a
decline in area of occupancy, extent of occurrence and qual-
ity of habitat; (d) = actual or potential levels of exploita-
tion, B = geographic range in the form of EOO and AOO’
tinuing 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 vari-
ables. 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 veg-
etation patterns and species composition. Our results show simi-
larities 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 Paki-
stan (Stewart 1972; Ali and Qaiser 1995). Khan et al. (2014) also
found that Poaceae and Asteraceae were the dominant plant fami-
lies 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 dis-
trict 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 fami-
lies. 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; Dur-
rani 2000; Malik 2005; Mehmood et al. 2015).
Plant species distribution, association formation and their
indicator species are typically attributed to edaphic and envi-
ronmental 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 associa-
tion are strongly correlated with high phosphorous concentra-
tions and steep slopes. Ailanthus altissima, Rosa webbiana and
Arenaria leptoclados are the top indicator species of 2nd asso-
ciation. 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 satura-
tion 40–70% at altitude between 1556–2313 m. The Indicator
Species Analysis shows that the indicator species of this associa-
tion 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 associa-
tion 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 vari-
ables. 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 envi-
ronmental gradients. Furthermore, the vegetation of Cholistan
desert was explored by Noureen et al. (2008) on the basis of envi-
ronmental variables. Whereas, Yimer (2007) stated that soil inter-
rupt 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 forma-
tion. 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 popu-
lation and geographic range i.e., EOO = 2218 k­ m2 (< 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. jacque-
montiana 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 potas-
sium concentrations, elevation, aspect, slope, lower pH, electri-
cal 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 indica-
tors 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


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 maxi-
mum observed indicator value (IV) of species (P ≤ 0.05) (top indicators are showed in bold font)
Botanical names Aspect Potassium Phosphorus pH Saturation Slope Numbers of Stump

13
Max IV P* Max IV P* Max IV P* Max IV P* Max IV P* Max IV P* Max IV P*

Achyranthes aspera Linn 4 8.3 0.112 180 5.3 0.612 5 9.1 0.262 7 33.3 0.025 62 50 0.104 3 1.7 1.000 0 2.2 1.000
Acrachne racemosa (Heyne ex Roem. and Schult.) 4 8.3 0.117 180 5.3 0.614 5 9.1 0.252 5 25 0.068 50 22.3 0.336 2 2.2 0.421 0 2.2 1.000
Adiantum caudatum 4 4.6 0.885 180 4.4 0.810 1 18.4 0.037 6 7.1 1.000 72 13.3 0.563 3 4.6 0.657 1 6.7 0.529
Adiantum venustum 1 9.7 0.537 240 17.1 0.239 3 7.8 0.432 6 9.1 0.974 65 11.5 0.661 3 10.2 0.133 3 4.4 0.895
Agrostis gigantea Roth, Tent 1 6.5 0.614 180 13.9 0.225 6 4.4 0.754 6 6.1 1.000 49 21.3 0.326 2 5 0.445 1 8 0.334
Ajuga bracteosa Wall 1 1.1 1.000 200 6.2 0.273 1 7.7 0.370 6 1 1.000 70 8.3 0.792 2 2.2 0.425 2 5.9 0.258
Alliaria petiolata (M. Bieb.) 1 1.1 1.000 160 5.9 0.426 6 11.1 0.143 6 1 1.000 50 11.1 0.538 3 1.7 1.000 3 12.5 0.105
Androsace rotundifolia Hardwicke 1 2.2 1.000 160 2.8 0.927 7 12.6 0.094 6 2 1.000 50 5.6 0.925 2 1.1 1.000 1 1.6 1.000
Arabidopsis wallichii (Hook. f. and Thoms.) 1 3.2 1.000 100 38.7 0.073 7 28.1 0.007 6 3 1.000 66 8 0.803 2 6.7 0.074 0 6.7 0.435
Arenaria leptoclados (Reichb.) 4 41.7 0.000 140 11.6 0.246 3 11.1 0.119 5 19.9 0.251 42 69.7 0.005 3 5 0.373 0 11.1 0.212
Arenaria orbiculata Royle ex Edgew. and Hook.f 4 8.3 0.117 180 5.3 0.614 5 9.1 0.252 5 25 0.068 46 19.6 0.275 2 2.2 0.421 0 2.2 1.000
Arisaema flavum (Forsk.) 1 4.3 0.916 220 5.9 0.516 6 17 0.035 6 4 1.000 65 25 0.244 3 6.7 0.137 3 3.9 0.661
Arisaema jacquemontii Blume, Rumphia 1 6.5 0.675 180 9 0.407 7 8.4 0.343 6 6.1 1.000 47 20.6 0.285 3 10 0.067 7 19.3 0.077
Artemisia vulgaris Linnaeus 1 4.3 0.839 120 2 0.977 4 3 0.939 6 4 1.000 60 6 0.877 2 2.3 1.000 1 4.4 0.558
Arthraxon prionodes (Steud.) 1 1.1 1.000 200 6.2 0.272 1 7.7 0.371 6 1 1.000 71 50 0.108 2 2.2 0.427 1 3.1 0.574
Asplenium adiantum-nigrum 1 9.6 0.637 140 9 0.551 5 15.6 0.082 6 12.1 0.911 58 8.3 0.839 3 7.7 0.494 2 5.3 0.851
Asplenium ceterach 1 3.2 1.000 200 3.2 0.668 5 5.3 0.550 6 3 1.000 58 28.3 0.222 2 3.2 0.466 0 2.6 0.867
Asplenium trichomanes 4 5.8 0.687 100 21.6 0.187 5 6.2 0.569 5 14.4 0.395 43 13.6 0.550 3 7.6 0.283 2 7.7 0.473
Asplenium tricorn 1 1.1 1.000 200 6.2 0.273 4 5.9 0.546 6 1 1.000 50 11.1 0.563 2 2.2 0.432 7 33.3 0.029
Athyrium sp. 1 1.1 1.000 220 11.1 0.112 3 4.3 0.760 6 1 1.000 65 8.3 0.790 2 2.2 0.415 0 2.2 1.000
Bergenia ciliata (Haw.) 1 1.1 1.000 140 5 0.795 2 4 1.000 6 1 1.000 70 8.3 0.792 3 1.7 1.000 0 2.2 1.000
Author's personal copy

Bistorta amplexicaulis (D. Don) 1 19.4 0.204 240 15.5 0.300 4 17.5 0.081 6 18.2 0.651 62 10.4 0.708 2 8.7 0.972 3 4 1.000
Brachypodium sylvaticum (Huds.) P.Beauv 1 22.4 0.518 200 15.2 0.410 1 22.9 0.049 6 35.4 0.278 71 28.1 0.095 2 25 0.150 3 12.5 0.759
Bromus japonicus Thunb 1 1.1 1.000 180 5.3 0.622 1 7.7 0.380 6 1 1.000 70 8.3 0.792 2 2.2 0.429 1 3.1 0.568
Bupleurum falcatum Linn 1 7.5 0.595 200 6.3 0.618 2 10 0.279 6 7.1 1.000 50 10.5 0.723 3 9.4 0.113 7 14 0.171
Bupleurum lanceolatum Wall 1 2.2 1.000 200 3.9 0.473 2 2.3 1.000 6 2 1.000 65 4.6 1.000 3 3.3 0.515 1 2.1 0.870
Cannabis sativa Linn 4 41.7 0.002 120 7.2 0.397 5 5.2 0.597 7 26.4 0.099 42 26.6 0.215 3 8.3 0.085 0 8.1 0.330
Carex sp. 4 16.7 0.279 180 17.5 0.285 6 4.5 0.962 5 24.6 0.205 49 16.6 0.346 2 7.9 0.951 1 10.7 0.513
Chenopodium album Linnaeus 4 24.4 0.003 180 3.7 0.828 5 9.8 0.268 5 12.7 0.291 62 12.5 0.576 3 6.2 0.236 2 3.6 0.813
Chenopodium botrys L 4 8.3 0.116 120 4.8 1.000 4 5.9 0.550 6 1 1.000 70 8.3 0.785 3 1.7 1.000 0 2.2 1.000
Chenopodium murale L 4 24.7 0.001 180 2.8 0.892 7 4 0.785 5 45.1 0.009 42 44.8 0.039 2 6.2 0.184 0 5.3 0.521
Conyza canadensis 1 2.2 1.000 240 45.5 0.029 4 3.5 0.817 6 2 1.000 68 13.9 0.499 2 4.4 0.178 0 4.4 0.496
Cymbopogon jwarancusa (Jones) 1 3.2 1.000 200 3.2 0.676 6 6 0.435 6 3 1.000 55 6 0.854 3 5 0.262 0 2.8 0.884
Cymbopogon pospischilii (K. Schum.) 4 4.9 0.782 100 31.3 0.056 7 6.1 0.529 6 5.1 1.000 49 5 0.955 3 3.9 0.566 2 4.3 0.740
Tropical Ecology

Cynoglossum glochidiatum Wall. ex Benth 1 2.2 1.000 220 5.7 0.361 7 14.7 0.059 6 2 1.000 48 17.9 0.378 3 3.3 0.499 2 4.9 0.394
 Table 2  (continued)
Botanical names 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*
Tropical Ecology

Cynoglossum lanceolatum Forssk 1 1.1 1.000 200 6.2 0.273 4 5.9 0.546 6 1 1.000 50 11.1 0.563 2 2.2 0.432 7 33.3 0.029
Cystopteris fragilis (L.) Bernh 1 2.2 1.000 180 10.5 0.155 4 5 0.637 6 2 1.000 55 9 0.673 2 2 0.766 3 12.1 0.121
Desmostachya bipinnata (Linn.) 1 1.1 1.000 160 5.9 0.437 2 4 1.000 6 1 1.000 60 4.8 1.000 2 2.2 0.432 2 5.9 0.265
Dioscorea deltoidea Wall 1 3.2 1.000 160 9.3 0.193 5 5.9 0.461 6 3 1.000 50 6.5 0.851 3 5 0.259 0 6.7 0.424
Dryopteris stewartii Fraser-Jenk 1 4.3 0.918 220 5.5 0.546 6 5.9 0.455 6 4 1.000 58 7.9 0.839 3 4.5 0.394 3 6.8 0.462
Duchesnea indica (Andrews) 1 1.1 1.000 180 5.3 0.628 6 11.1 0.143 6 1 1.000 55 11.1 0.559 2 2.2 0.432 2 5.9 0.271
Dysphania botrys (L.) 1 1.1 1.000 200 6.2 0.273 4 5.9 0.546 6 1 1.000 50 11.1 0.563 2 2.2 0.432 7 33.3 0.029
Euphorbia indica Lam 1 3.2 1.000 120 2.4 0.956 5 6.1 0.462 6 3 1.000 70 9.9 0.681 3 2.6 0.619 0 3.4 0.705
Euphorbia wallichii Hook 1 3.2 1.000 200 3.8 0.658 2 5.1 0.680 6 3 1.000 60 2.5 1.000 3 5 0.263 1 5.5 0.497
Fallopia convolvulus (L.) 1 1.1 1.000 140 5 0.791 2 4 1.000 7 33.3 0.030 50 11.1 0.555 3 1.7 1.000 0 2.2 1.000
Fragaria vesca L 1 27.5 0.265 160 12.1 0.618 2 8.8 0.887 6 37.4 0.251 71 17 0.303 3 18.9 0.803 2 11.6 0.824
Galium aparine L 4 3.6 1.000 140 6.1 0.597 5 12.7 0.128 6 6.1 1.000 60 3.9 0.990 3 10 0.056 0 2.7 0.985
Galium tricornutum Dandy 1 3.2 1.000 120 8.1 0.252 5 5.1 0.597 6 3 1.000 60 6.9 0.829 3 5 0.255 0 2.5 0.901
Geranium wallichianum D. Don ex sweet 1 3.2 1.000 180 9.7 0.197 6 7.3 0.331 6 3 1.000 55 10.4 0.561 2 1.4 1.000 2 4.2 0.589
Girardinia diversifolia (Forssk.) 1 3.2 1.000 140 7.2 0.274 6 5.8 0.426 6 3 1.000 50 5.1 0.916 3 5 0.251 0 2.8 0.856
Girardinia palmata (Forssk.) Gaudich 1 1.1 1.000 120 4.8 1.000 4 5.9 0.534 6 1 1.000 62 50 0.090 3 1.7 1.000 0 2.2 1.000
Herniaria glabra L 1 2.2 1.000 180 2.7 0.926 6 8.2 0.260 7 32.4 0.054 50 11.1 0.555 3 3.3 0.515 2 4.3 0.636
Impatiens brachycentra Kar. and Kir 1 13.7 0.749 100 7.6 0.893 6 11 0.392 5 20.9 0.526 43 28.9 0.074 3 20.2 0.095 3 9.7 0.753
Lactuca dissecta D.Don 4 40.4 0.000 140 6.6 0.600 3 2.5 0.988 5 45 0.030 42 34.3 0.046 2 4 0.947 0 10.2 0.374
Lamium album L 4 8.3 0.103 140 5 0.795 3 4.3 0.762 6 1 1.000 70 8.3 0.802 3 1.7 1.000 0 2.2 1.000
Lecanthus peduncularis (Royle) Wedd 1 1.1 1.000 180 5.3 0.621 7 16.7 0.053 6 1 1.000 48 25 0.218 3 1.7 1.000 2 5.9 0.279
Lespedeza juncea (Linn.) Pers 1 3.2 1.000 220 19.3 0.151 1 11.8 0.153 6 4 1.000 72 4.8 0.976 2 8.9 0.034 1 4.5 0.720
Author's personal copy

Melissa officinalis L 1 1.1 1.000 160 5.9 0.437 2 4 1.000 6 1 1.000 60 4.8 1.000 2 2.2 0.432 2 5.9 0.265
Micromeria biflora Benth 1 2.2 1.000 200 12.5 0.101 1 5.1 0.618 6 2 1.000 72 14.1 0.497 2 4.4 0.179 3 10.6 0.155
Oenothera javanica 1 1.1 1.000 180 5.3 0.605 4 5.9 0.539 6 1 1.000 55 11.1 0.545 2 2.2 0.448 3 12.5 0.114
Onychium contiguum C.Hope 1 1.1 1.000 140 5 0.808 2 4 1.000 6 1 1.000 58 33.3 0.125 2 2.2 0.429 0 2.2 1.000
Oplismenus hirtellus (L.) P.Beauv 1 2.2 1.000 220 10.9 0.154 6 10.8 0.176 6 2 1.000 65 7.2 0.856 3 1.6 1.000 1 3 0.698
Oplismenus undulatifolius (Ard.) 1 3.2 1.000 220 8.1 0.307 3 5.3 0.597 6 3 1.000 49 5.8 0.911 2 1.4 1.000 0 2.7 0.823
Origanum vulgare L 1 13.3 0.597 100 21.5 0.159 7 14.5 0.146 7 27.4 0.212 49 13.2 0.464 2 17.6 0.123 1 10.5 0.617
Oxalis corniculata L 4 31.2 0.007 240 14.5 0.351 6 10 0.334 5 13.4 0.904 42 26 0.125 2 10.9 0.319 2 8.7 0.628
Perilla frutescens (L.) Britton 1 13.2 0.428 140 3.9 0.995 6 5.4 0.881 6 16.2 0.746 49 13.3 0.506 3 12.5 0.273 3 18.7 0.137
Pimpinella diversifolia DC., Prodr 1 5.4 0.702 200 13.9 0.211 1 18.5 0.033 6 5.1 1.000 70 15.2 0.483 2 7.9 0.104 3 13.1 0.198
Piptatherum laterale (Regel) Nevski 4 60.6 0.007 240 15.8 0.582 4 23.4 0.077 5 56.8 0.078 47 35.4 0.004 2 28.9 0.953 7 44.2 0.018
Plantago lanceolata L 4 7.8 0.117 220 2.5 0.961 5 6.3 0.418 5 24.2 0.117 65 5.8 0.864 2 6.7 0.073 0 3.7 0.685
Plantago major L 4 8.3 0.117 180 5.3 0.614 5 9.1 0.252 5 25 0.068 68 13.2 0.463 2 2.2 0.421 0 2.2 1.000

13


Table 2  (continued)
Botanical names Aspect Potassium Phosphorus pH Saturation Slope Numbers of Stump
P* P* P* P* P* P* P*

13
Max IV Max IV Max IV Max IV Max IV Max IV Max IV

Polygala abyssinica R.Br. ex Fresen 1 3.2 1.000 220 6.5 0.408 7 7.2 0.309 6 3 1.000 72 13.5 0.527 2 6.7 0.070 2 3.3 0.727
Polygonatum cirrhifolium (Wall.) Royle 1 2.2 1.000 140 3.8 0.805 2 3.1 0.905 6 2 1.000 49 6.1 0.864 3 3.3 0.501 2 2.5 0.828
Polygonatum multiflorum (L.) All 1 2.2 1.000 160 3.8 0.520 5 5.3 0.608 6 2 1.000 60 4.8 1.000 3 3.3 0.509 3 9.5 0.194
Polygonatum odoratum (Mill.) Druce 1 1.1 1.000 200 6.2 0.282 1 7.7 0.372 6 1 1.000 71 50 0.104 3 1.7 1.000 3 12.5 0.105
Polygonatum verticillatum (L.) All 1 1.1 1.000 160 5.9 0.426 6 11.1 0.143 6 1 1.000 50 11.1 0.538 3 1.7 1.000 3 12.5 0.105
Polygonum paronychioides C.A. Mey 4 6.2 0.307 120 4.5 0.717 4 14.1 0.062 6 4 1.000 50 5.1 0.915 2 2.4 0.867 7 30.3 0.006
Potentilla anserina L 4 8.3 0.110 140 5 0.798 3 4.3 0.763 6 1 1.000 55 11.1 0.548 3 1.7 1.000 0 2.2 1.000
Prunella vulgaris L 1 5.4 0.857 220 12.3 0.259 7 5.9 0.546 7 30.9 0.044 58 12.1 0.583 2 3.7 0.811 3 4.8 0.771
Pteridium aquilinum (L.) Kuhn 1 7.5 0.586 160 10.2 0.377 7 7.1 0.461 6 7.1 1.000 47 20.5 0.347 3 11.7 0.028 7 14.2 0.193
Ranunculus sceleratus L 1 4.3 0.921 180 14 0.179 7 28.4 0.003 6 4 1.000 48 34.6 0.111 3 6.7 0.142 2 8.1 0.337
Rostraria cristata (L.) Tzvelev 4 5 0.876 180 4.1 0.751 2 6.4 0.478 7 30.6 0.043 50 6.1 0.905 3 6.7 0.220 0 3.9 0.894
Rosularia rosulata (Edgew.) 4 16.7 0.011 140 2.8 0.894 4 3.2 0.892 6 2 1.000 55 6.8 0.808 3 3.3 0.509 0 4.4 0.493
Rubia cordifolia L 1 1.1 1.000 220 11.1 0.124 6 11.1 0.148 6 1 1.000 65 8.3 0.792 3 1.7 1.000 1 3.1 0.570
Rumex dentatus L 1 2.2 1.000 180 10.5 0.172 6 7.4 0.399 5 22.3 0.123 55 8.3 0.747 2 1.8 0.747 2 11.8 0.127
Scrophularia scabiosifolia Benth 4 8.3 0.117 160 5.9 0.436 4 5.9 0.538 6 1 1.000 55 11.1 0.548 3 1.7 1.000 1 3.1 0.572
Stellaria media (L.) Vill 1 5.4 0.856 160 6.4 0.416 4 7.5 0.348 6 5.1 1.000 68 12.9 0.569 2 4.9 0.311 7 48.5 0.003
Tagetes minuta L 1 4.3 0.839 240 21.3 0.111 1 4.7 0.717 6 4 1.000 72 10.2 0.687 2 8.9 0.034 0 8.9 0.298
Taraxacum officinale (L.) 1 8.6 0.571 180 14.7 0.303 1 5.7 0.624 6 8.1 1.000 47 19.1 0.321 2 14.2 0.005 7 21.1 0.075
Thalictrum foliolosum DC 1 1.1 1.000 180 5.3 0.615 7 16.7 0.052 6 1 1.000 48 25 0.205 3 1.7 1.000 1 3.1 0.570
Trifolium repens L 1 1.1 1.000 200 6.2 0.267 6 11.1 0.142 6 1 1.000 66 20 0.307 2 2.2 0.422 1 3.1 0.567
Trigonella sp. 1 1.1 1.000 160 5.9 0.431 2 4 1.000 6 1 1.000 68 25 0.210
Urtica dioica L 4 8.3 0.116 120 4.8 1.000 4 5.9 0.550 6 1 1.000 70 8.3 0.785 3 1.7 1.000 0 2.2 1.000
Author's personal copy

Valeriana jatamansi Jones 1 8.6 0.573 240 34.8 0.040 5 4.3 0.826 6 8.1 1.000 66 8.5 0.807 2 4.1 0.907 3 6.1 0.638
Verbascum thapsus L 1 22.6 0.160 240 6.9 0.886 7 12.5 0.238 6 21.2 0.535 48 19 0.198 2 15.8 0.176 1 22.7 0.105
Viola odorata L 4 53.9 0.019 180 14.2 0.676 7 16.2 0.386 5 27 0.786 47 13.2 0.470 3 50.5 0.001 7 39.8 0.024
Viola rupestris F.W.Schmidt 1 1.1 1.000 200 6.2 0.276 1 7.7 0.372 6 1 1.000 70 8.3 0.786 2 2.2 0.438 1 3.1 0.566
Sida cordata (Burm.f.) Borss.Waalk 1 1.1 1.000 160 5.9 0.437 2 4 1.000 6 1 1.000 60 4.8 1.000 2 2.2 0.432 2 5.9 0.265
Berberis lycium Royle in Trans. Linn 1 11.8 0.400 220 27.9 0.053 5 4.9 0.797 6 11.1 0.979 66 8.9 0.841 3 6.3 0.757 0 7.3 0.668
Clematis grata Wall 1 2.2 1.000 140 10 0.178 2 8 0.333 6 2 1.000 55 7.4 0.776 3 3.3 0.503 0 4.4 0.614
Cotoneaster microphyllus 4 39 0.017 100 22.3 0.135 7 18.2 0.080 5 34.9 0.191 49 7.7 0.899 2 19.1 0.264 0 13.6 0.466
Cotoneaster nummularia 1 2.2 1.000 180 3.6 0.770 4 3.9 0.750 6 2 1.000 65 6 0.868 2 4.4 0.184 7 31.1 0.027
Desmodium elegans DC 1 1.1 1.000 160 5.9 0.434 2 4 1.000 6 1 1.000 72 20 0.310 3 1.7 1.000 1 3.1 0.582
Hedera nepalensis K. Koch, Hort. Dendrol 4 4.5 1.000 120 4 0.880 2 3.4 0.941 6 9.1 1.000 62 17.4 0.361 3 7.3 0.316 3 12.1 0.315
Indigofera heterantha Wall 4 53.1 0.043 200 20.5 0.268 1 21.3 0.103 5 43.9 0.254 72 17.3 0.116 2 44.6 0.055 7 18.4 0.785
Isodon rugosus (Wall. ex Benth.) 1 20.4 0.183 100 12.6 0.398 7 15.6 0.125 5 18.1 0.636 71 6.6 0.952 3 12.4 0.461 2 8.6 0.762
Tropical Ecology
 Table 2  (continued)
Botanical names 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*
Tropical Ecology

Jasminum humile Linn 1 14 0.336 100 27.1 0.122 7 14.8 0.102 6 13.1 0.879 49 9.5 0.764 2 7.9 0.637 0 9.4 0.556
Lonicera quinquelocularis Hardwicke 1 7.5 0.604 180 4.3 0.798 1 10.6 0.181 6 7.1 1.000 72 27.3 0.122 2 6.1 0.379 1 10.9 0.302
Myrsine africana Linn 4 4.3 0.950 100 30.8 0.058 7 11.3 0.190 6 8.1 1.000 49 4.9 0.971 2 5 0.722 0 10.4 0.325
Onopordum acanthium L 1 1.1 1.000 140 5 0.803 3 4.3 0.756 6 1 1.000 55 11.1 0.557 3 1.7 1.000 2 5.9 0.275
Parrotiopsis jacquemontiana (Dcne.) 1 67 0.001 240 17.2 0.490 7 24.2 0.011 6 41.1 0.489 71 11.9 0.295 2 59.5 0.005 3 22.8 0.845
Prunus tomentosa Thunb 4 23.6 0.005 160 3.8 0.811 2 3.8 0.815 6 5.1 1.000 49 6.4 0.901 2 7.8 0.127 7 13.3 0.169
Rosa brunonii Lindl 1 3.2 1.000 240 41.8 0.016 4 2.4 1.000 6 3 1.000 62 37.5 0.094 3 2 1.000 0 2.6 1.000
Rubus niveus Thunb 1 1.1 1.000 160 5.9 0.440 2 4 1.000 6 1 1.000 68 25 0.198 2 2.2 0.426 0 2.2 1.000
Sarcococca saligna (D.Don) 1 7.5 0.581 140 7.5 0.480 2 9.4 0.259 6 7.1 1.000 58 16.2 0.403 3 4.2 0.791 0 10.9 0.275
Sorbaria tomentosa (Lindl.) Rehder 1 4.3 0.918 180 2.8 0.843 4 16.8 0.041 6 4 1.000 62 34.9 0.101 3 3.8 0.470 3 7.3 0.392
Spiraea sp. 1 3.2 1.000 120 7.6 0.423 3 6.1 0.522 6 3 1.000 60 6.5 0.871 2 3.9 0.468 0 3.5 0.749
Strobilanthes glutinosus (Nees) Bremek 1 2.2 1.000 120 4 1.000 5 8.8 0.239 6 2 1.000 60 4.2 0.944 3 1.5 1.000 1 3 0.725
Rosa webbiana 4 16.7 0.013 160 3.6 0.559 4 3.8 0.789 6 2 1.000 42 38.3 0.152 2 1.1 1.000 1 2.1 0.865
Viburnum cotinifolium D. Don 1 11.8 0.402 180 5.8 0.784 6 9.5 0.313 6 11.1 1.000 47 18.3 0.325 2 5.3 0.977 1 7.8 0.630
Viburnum grandiflorum Wall. ex DC 1 10.8 0.459 240 22.1 0.205 7 19 0.033 6 10.1 1.000 48 15.8 0.424 3 6.4 0.634 7 12.2 0.338
Wikstroemia canescens Wall. ex Meisn 1 18.3 0.227 220 5.1 0.963 5 8.4 0.527 6 17.2 0.704 47 10.4 0.711 3 18.9 0.031 7 9.4 0.634
Acer caesium Wall 4 6 0.386 120 4.6 0.705 6 4.1 0.683 7 18.4 0.238 50 3.5 0.990 3 3.5 0.632 3 7.5 0.436
Ailanthus altissima (Mill.) 1 2.2 1.000 240 45.5 0.031 4 3.4 0.892 6 2 1.000 48 18.7 0.337 2 4.4 0.179 1 1.8 1.000
Cedrus deodara (Roxb. ex D. Don) 4 17.9 0.072 160 22.2 0.189 7 17.6 0.073 5 26.8 0.136 47 28.8 0.114 3 10.7 0.184 7 22.6 0.069
Celtis australis Willd 1 1.1 1.000 140 5 0.801 1 7.7 0.369 6 1 1.000 70 8.3 0.779 2 2.2 0.448 1 3.1 0.572
Cornus macrophylla Wall 1 8.6 0.595 180 14.3 0.323 1 3.8 0.888 6 8.1 1.000 58 17.2 0.368 2 6.2 0.439 1 5 0.754
Diospyros kaki Linn 1 1.1 1.000 200 6.2 0.272 1 7.7 0.371 6 1 1.000 71 50 0.108 2 2.2 0.427 1 3.1 0.574
Author's personal copy

Diospyros lotus Linn 1 3.2 1.000 200 10.1 0.167 1 13.6 0.053 6 3 1.000 71 35.3 0.133 2 6.7 0.071 2 3.4 0.722
Ficus carica L 1 1.1 1.000 160 5.9 0.426 6 11.1 0.143 6 1 1.000 50 11.1 0.538 3 1.7 1.000 3 12.5 0.105
Juglans regia Linn 4 7.4 0.212 200 3.5 0.499 1 4.9 0.693 6 2 1.000 71 40.9 0.134 2 1.3 1.000 1 1.8 1.000
Olea ferruginea Royle 1 1.1 1.000 160 5.9 0.439 5 9.1 0.252 6 1 1.000 60 4.8 1.000 3 1.7 1.000 0 2.2 1.000
Pinus roxburghii Sarg 1 4.3 1.000 220 5.9 0.438 6 6.3 0.427 6 4 1.000 65 3.2 0.990 3 6.7 0.136 1 3.7 0.782
Pinus wallichiana A.B.Jacks 1 25.8 0.106 160 14.6 0.377 7 13.3 0.267 6 24.2 0.454 71 15 0.445 3 27.7 0.004 3 37.6 0.037
Platanus orientalis L 4 8.3 0.110 140 5 0.798 3 4.3 0.763 6 1 1.000 55 11.1 0.548 3 1.7 1.000 0 2.2 1.000
Populus ciliata Wall. ex Royle 1 1.1 1.000 180 5.3 0.615 7 16.7 0.052 6 1 1.000 48 25 0.205 3 1.7 1.000 1 3.1 0.570
Quercus baloot Griff 1 13.6 0.972 140 14.9 0.376 1 21 0.046 7 23.4 0.486 72 13.1 0.446 3 14.8 0.917 1 10.7 0.790
Quercus dilatata Royle, Illust 1 10.5 0.817 160 13.1 0.394 5 14.6 0.146 5 8.5 1.000 70 7.5 0.892 2 9.4 0.034 1 9.3 0.692
Quercus incana 1 3.2 1.000 120 2.5 0.985 6 8.7 0.229 6 3 1.000 58 16.7 0.419 3 2.7 0.620 1 2 0.952
Quercus semecarpifolia Sm 1 18.3 0.228 200 13.4 0.385 4 24 0.020 6 17.2 0.676 72 5.5 0.978 2 20.5 0.011 7 43.7 0.020
Rhamnus virgatus Roxb 1 1.1 1.000 120 4.8 1.000 2 4 1.000 6 1 1.000 60 4.8 1.000 3 1.7 1.000 0 2.2 1.000

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Tropical Ecology

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