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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 13 Vol.:(0123456789) Tropical Ecology 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 13 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 Tropical Ecology 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 13 Tropical Ecology 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 13 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%. Tropical Ecology 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 13 Tropical Ecology 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 13 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. Tropical Ecology 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 13 Tropical Ecology 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. 13 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. Tropical Ecology 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 13 Tropical Ecology 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 13 (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 Tropical Ecology 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 Tropical Ecology Abbas Z, Alam J, Muhammad S, Bussmann R, Khan SM, Hussain M (2019) Phyto-cultural diversity of the Shigar valley (Central Karakorum) Baltistan, Northern Pakistan. Ethnobot Res Appl 18:31. https://doi.org/10.32859/era.18.31.1-18 Abbas Z, Khan SM, Alam J, Ullah Z, Abideen Z (2020) Plant communities and anthropo-natural threats in the Shigar Valley, (Central Karakorum) Baltistan-Pakistan. Pak J Bot 53(3):987–994 Abdullah KSM, Pieroni A, Haq UZ, Ahmad Z (2020) Mazri (Nannorrhops ritchiana (Griff) Aitch.): a remarkable source of manufacturing traditional handicrafts, goods and utensils in Pakistan. J Ethnobiol Ethnomed 16:45. https://doi.org/10.1186/s13002-02000394-0 Ahmad Z, Khan SM, Ali S, Rahman I, Ara H, Noreen I, Khan A (2016a) Indicator species analyses of weed communities of maize crop in district Mardan, Pakistan. Pak J Weed Sci Res 22(2):227–238 Ahmad Z, Khan SM, Abd_Allah EF, Alqarawi AA, Hashem A (2016b) Weed species composition and distribution pattern in the maize crop under the influence of edaphic factors and farming practices: a case study from Mardan, Pakistan. Saudi J Biol Sci 23(6):741–748 Ahmad Z, Khan SM, Ali MI, Fatima N, Ali S (2019) Pollution indicandum and marble waste polluted ecosystem; role of selected indicator plants in phytoremediation and determination of pollution zones. J Clean Prod 236:117709 Ahmed M, Husain T, Sheikh A, Hussain SS, Siddiqui MF (2006) Phytosociology and structure of Himalayan forests from different climatic zones of Pakistan. Pak J Bot 38(2):361 Ali S, Nasir J (1990) Flora of Pakistan. Shamim Printing Press, Karachi Ali S, Qaiser M (1995) 2006, Flora of Pakistan. Agricultural Research Council, Islamabad Ali H, Qaiser M (2010a) Contribution to the red list of Pakistan. A case study of Astragalus gahiratensis Ali (Fabaceae-Papilionoideae). Pak J Bot 42(3):1523–1528 Ali H, Qaiser M (2010b) Contribution to the Red List of Pakistan: a case study of Gaillonia chitralensis (Rubiaceae). Pak J Bot 42:205–212 Ali A, Badshah L, Hussain F (2018) Vegetation structure and threats to montane temperate ecosystems in Hindukush Range, Swat, Pakistan. Appl Ecol Environ Res 16(4):4789–4811 Anwar S, Khan SM, Ahmad Z, Ullah Z, Iqbal M (2019) Floristic composition and ecological gradient analyses of the Liakot Forests in the Kalam region of District Swat, Pakistan. J For Res 30(4):1407–1416 Bano S, Khan SM, Alam J, Alqarawi AA, Abd_Allah EF, Ahmad Z, Rahman IU, Ahmad H, Aldubise A, Hashem A (2018) Ecofloristic studies of native plants of the Beer Hills along the Indus River in the districts Haripur and Abbottabad, Pakistan. Saudi J Biol Sci 25(4):801–810 Bergeron SP, Bradley RL, Munson A, Parsons W (2013) Physicochemical and functional characteristics of soil charcoal produced at five different temperatures. Soil Biol Biochem 58:140–146 Borcard D, Legendre P, Drapeau P (1992) Partialling out the spatial component of ecological variation. Ecology 73(3):1045–1055 Braak CJ, Prentice IC (1988) A theory of gradient analysis. Advances in ecological research, 18th edn. Elsevier, New York, pp 271–317 Brown L, Bezuidenhout H (2005) The vegetation of the farms Ingleside and Welgedacht of the Mountain Zebra National Park, Eastern Cape. Koedoe 48(2):23–42 Cook ER, Kairiukstis LA (2013) Methods of dendrochronology: applications in the environmental sciences. Springer, Berlin Tropical Ecology Curtis JT, McIntosh RP (1950) The interrelations of certain analytic and synthetic phytosociological characters. Ecology 31(3):434–455 Dhyani A, Jain R, Pandey A (2019) Contribution of root-associated microbial communities on soil quality of oak and pine forests in the Himalayan ecosystem. Trop Ecol 60(2):271–280 Dufrêne M, Legendre P (1997) Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol Monogr 67(3):345–366 Durrani M (2000) Ecological evaluation of some rangeland plants of Harboi Hills, Kalat, Balochistan. Ph. D, Thesis University of Peshawar, Pakistan Fujiwara K (1987) Aims and methods of phytosociology or “vegetation science”. Plant ecology and taxonomy to the memory of Dr Satoshi Nakanishi. The Kobe Geobotanical Society, Kobe, pp. 607–628 Gaston KJ (2000) Global patterns in biodiversity. Nature 405(6738):220–227 Gee GW, Bauder JW (1986) Particle-size analysis: methods of soil analysis: part 1—physical and mineralogical methods (methods of soil). Geoarchaeology 5(1):383–411 Hair J, Black W, Babin B, Anderson R, Tatham R (2006) Multivariate data analysis, 6th edn. Pearson Prentice Hall, Upper Saddle River Haq UZ, Khan SM, Ahmad Z, Abdullah SSA, Mustafa G, Razzaq A, Manan F, Ullah A, Hussain M (2020) An evaluation of conservation status and ecological zonation of Alnus Nitida; a monophyletic species of the Sino-Japanese Region. J Anim Plant Sci 30(5):1224–1235 Hazrat A, Wahab M (2011) Threatened native plants of Dir Kohistan valley, Khyber Pukhtunkhwa, Pakistan. FUUAST J Biol 1:35–38 Hazrat A, Shah J, Ali M, Iqbal I (2007) Medicinal value of Ranunculaceae of Dir valley. Pak J Bot 39(4):1037 Hussain I, Olson KR, Ebelhar SA (1999) Long-term tillage effects on soil chemical properties and organic matter fractions. Soil Sci Soc Am J 63(5):1335–1341 International Union for Conservation of Nature (IUCN) International Union for Conservation of Nature, and Natural Resources, Species Survival Commission (2001) IUCN red list categories and criteria. IUCN, Gland Iqbal M, Khan S, Khan MA, Ur Rahman I, Abbas Z (2015) Exploration and inventorying of weeds in wheat crop of the district Malakand, Pakistan. Pak J Weed Sci Res 21(3):435–452 Iqbal M, Khan SM, Khan MA, Ahmad Z, Ahmad H (2018) A novel approach to phytosociological classification of weeds flora of an agro-ecological system through cluster, two way cluster and indicator species analyses. Ecol Ind 84:590–606 Ismail AH, Lim CC, Omar WMW (2019) Evaluation of spatial and temporal variations in zooplankton community structure with reference to water quality in Teluk Bahang Reservoir, Malaysia. Trop Ecol 60(2):186–198 Jackson M (1963) Interlayering of expansible layer silicates in soils by chemical weathering. Clays Clay Miner 11(1):29–46 Jan A, Ali SI (2009) Conservation status of Astragalus gilgitensis Ali (Fabaceae): a critically endangered species in the Gilgit District, Pakistan. Phyton (Horn) 48(2):211–223 Kamran S, Khan SM, Ahmad Z, Rahman AU, Iqbal M, Manan F, Haq ZU, Ullah S (2019) The role of graveyards in species conservation and beta diversity: a vegetation appraisal of sacred habitats from Bannu, Pakistan. J For Res 30(4):1147–1158 Kent M (2011) Vegetation description and data analysis: a practical approach. Wiley, Hoboken Khadanga SS, Jayakumar S (2020) Tree biomass and carbon stock: understanding the role of species richness, elevation, and disturbance. Trop Ecol:1–14 Khan SM, Ahmad H (2015) Species diversity and use patterns of the alpine flora with special reference to climate change in the Naran, Pakistan. Climate change impacts on high-altitude ecosystems. Springer, Berlin, pp 155–175 Khan SM, Harper D, Page S, Ahmad H (2011) Species and community diversity of vascular flora along environmental gradient in Naran Valley: a multivariate approach through indicator species analysis. Pak J Bot 43(5):2337–2346 Khan SM, Page S, Ahmad H, Shaheen H, Harper D (2012a) Vegetation dynamics in the Western Himalayas, diversity indices and climate change. Sci Technol Dev 31(3):232–243 Khan W, Ahmad H, Haq F, Islam M, Bibi F (2012b) Present status of moist temperate vegetation of Thandiani forests district Abbottabad Pakistan. Int J Biosci 2(10):80–88 Khan SM, Page SE, Ahmad H, Harper DM (2013) Sustainable utilization and conservation of plant biodiversity in montane ecosystems: the western Himalayas as a case study. Ann Bot 112(3):479–501 Khan SM, Page S, Ahmad H, Harper D (2014) Ethno-ecological importance of plant biodiversity in mountain ecosystems with special emphasis on indicator species of a Himalayan Valley in the northern Pakistan. Ecol Ind 37:175–185 Khan T, Khan IA, Rehman A, Ahmed N (2016a) Conservation status evaluation of Berberis species across the Karakoram Mountain Ranges, Pakistan using IUCN red list categories and criteria. J For Res 27(6):1385–1390 Khan W, Khan SM, Ahmad H, Ahmad Z, Page S (2016b) Vegetation mapping and multivariate approach to indicator species of a forest ecosystem: a case study from the Thandiani sub Forests Division (TsFD) in the Western Himalayas. Ecol Indic 71:336–351 Khan W, Khan SM, Ahmad H, Shakeel A, Page S (2017) Ecological gradient analyses of plant associations in the Thandiani forests of the Western Himalayas, Pakistan. Turk J Bot 41(3):253–264 Lepš J, Šmilauer P (2003) Multivariate analysis of ecological data using CANOCO. Cambridge University Press, Cambridge Malik Z (2005) Comparative study of the vegetation of Ganga Chotti and Bedori Hill Dist. Bagh Azad Jammu and Kashmir. Ph. D. thesis University of Peshawar, Peshawar. Mehmood A, Khan SM, Shah AH, Shah AH, Ahmad H (2015) First floristic exploration of the District Torghar, Khyber Pakhtunkhwa, Pakistan. Pak J Bot 47:57–70 Moore P, Chapman S (1986) Chemical analysis. Methods in plant ecology. Blackwell Scientific Publications, Oxford, pp 315–317 Muhammad I (2003) Conservation of indigenous medicinal plants and their traditional knowledge found in moist temperate Himalayas Pakistan. Quaid-i-Azam University, Islamabad Nasir Y (1991) Threatened plants of Pakistan. Plant life of South Asia. Royal Botanic Garden, Edinburgh, pp 229–234 Nazir A, Malik RN, Ajaib M (2012) Phytosociological studies of the vegetation of Sarsawa Hills District Kotli, Azad Jammu & Kashmir. Biologia (Pakistan) 58(1 and 2):123–133 Nelson DW, Sommers LE (1996) Total carbon, organic carbon, and organic matter. Methods soil anal. Part 3 Chem methods 5:961–1010 Noureen S, Arshad M, Mahmood K, Ashraf MY (2008) Improvement in fertility of nutritionally poor sandy soils of Cholistan desert, Pakistan by Calligonum polygonoides Linn. Pak J Bot 40(1):265–274 Pennings SC, Silliman BR (2005) Linking biogeography and community ecology: latitudinal variation in plant–herbivore interaction strength. Ecology 86(9):2310–2319 Perveen A, Hussain MI (2007) Plant biodiversity and phytosociological attributes of Gorakh Hill (Khirthar Range). Pak J Bot 39(3):691–698 Rahman AU, Khan SM, Saqib Z, Ullah Z, Ahmad Z, Ekercin S, Mumtaz AS, Ahmad H (2020) Diversity and abundance of climbers in relation to their hosts and elevation in the monsoon forests of Murree in the Himalayas. Pak J Bot 52(2):601–612 13 Tropical Ecology Ramzan M, Uddin S, Shah S, Ahmad M, Ali S, Ahmad B, Khan W, Ud Din S (2016) Tillage and mulching effect on weed dynamics and yield components of maize crop in district Peshawar under semi arid environment. Pak J Weed Sci Res 22(1):95–102 Ravindranath NH, Ostwald M (2007) Carbon inventory methods: handbook for greenhouse gas inventory, carbon mitigation and roundwood production projects, 29th edn. Springer, Berlin Sakya S, Bania A (1998) Natural vegetation of Chandragiri region. Ecoprint 5:51–55 Sarir M, Durrani M, Mian IA (2006) Effect of the source and rate of humic acid on phosphorus transformations. J Agric Biol Sci 1(1):29–31 Shaheen H, Shinwari ZK (2012) Phyto diversity and endemic richness of Karambar lake vegetation from Chitral, Hindukush-Himalayas. Pak J Bot 44(1):17–21 Shaheen H, Khan SM, Harper DM, Ullah Z, Allem Qureshi R (2011) Species diversity, community structure, and distribution patterns in western Himalayan alpine pastures of Kashmir, Pakistan. Mt Res Dev 31(2):153–159 Shaheen H, Awan SN, Aziz S (2017) Distribution pattern, conservation status, and associated flora of the genus Juniperus in subalpine pastures of the Kashmir Himalayas. Mt Res Dev 37(4):487–493 Shank R, Noorie E (1950) Microclimate vegetation in a small valley in eastern Tennessee. Ecology 11:531–539 Sheikh K, Ahmad T, Khan MA (2002) Use, exploitation and prospects for conservation: people and plant biodiversity of Naltar Valley, northwestern Karakorums, Pakistan. Biodivers Conserv 11(4):715–742 Shinwari Z, Shah M, Awan R (1996) The ethnobotany of Kharan district, Baluchistan. In: Proceedings of First Train. workshop Ethnob. Appl. Conserv., PARC, Islamabad, pp 124–132 Soltanpour P (1991) Determination of nutrient availability and elemental toxicity by AB-DTPA soil test and ICPS. Advances in soil science. Springer, Berlin, pp 165–190 13 Stewart RR (1972) Flora of West Pakistan: an annotated catalogue of the vascular plants of West Pakistan and Kashmir. Map Geog 6, Rawalpindi, p 1028 Takhtajan A (1970) Origin and dispersal of flowering plants. Springer, Leningrad Takhtajan A, Crovello TJ, Cronquist A (1986) Floristic regions of the world, 544th edn. University of California press, Berkeley ter Braak CJ, Barendregt LG (1986) Weighted averaging of species indicator values: its efficiency in environmental calibration. Math Biosci 78(1):57–72 Ullah A, Rashid A (2014) Conservation status of threatened medicinal plants of Mankial Valley Hindukush Range, Pakistan. Int J Biodivers Conserv 6(1):59–70 Ullah Z, Ahmad M, Sher H, Shaheen H, Khan SM (2015) Phytogeographic analysis and diversity of grasses and sedges (Poales) of northern Pakistan. Pak J Bot 47:93–104 Walter KS, Gillett HJ (1998) 1997 IUCN red list of threatened plants. IUCN, Gland Wana D (2002) Plant communities and diversity along altitudinal gradients from Lake Abaya to Chencha Highlands. MA Thesis. AAU Wilson MJ, Bayley SE (2012) Use of single versus multiple biotic communities as indicators of biological integrity in northern prairie wetlands. Ecol Ind 20:187–195 Yimer F (2007) Soil properties in relation to topographic aspects, vegetation communities and land use in the south-eastern highlands of Ethiopia Uppsala: Sveriges lantbruksuniv. Acta Universitatis Agr Sueciae 45:1652–6880 Zeb A, Iqbal Z, Khan SM, Rahman IU, Haq F, Afzal A, Ijaz F (2020) Species diversity, biological spectrum and phenological behaviour of vegetation of Biha Valley (Swat). Pakistan Acta Ecol Sinica 40(3):190–196