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Plant species diversity and density patterns along altitude gradient covering high-altitude alpine regions of west Himalaya, India

Plant species diversity and density patterns along altitude gradient covering high-altitude... GEOLOGY, ECOLOGY, AND LANDSCAPES INWASCON https://doi.org/10.1080/24749508.2022.2163606 RESEARCH ARTICLE Plant species diversity and density patterns along altitude gradient covering high-altitude alpine regions of west Himalaya, India a a b b c K. Chandra Sekar , Neha Thapliyal , Aseesh Pandey , Bhaskar Joshi , Sandipan Mukherjee , b b b d b Puja Bhojak , Monica Bisht , Deepika Bhatt , Sourab Singh and Amit Bahukhandi a b Garhwal Regional Centre, G.B. Pant National Institute of Himalayan Environment, Srinagar, India; Centre for Biodiversity Conservation and Management, G. B. Pant National Institute of Himalayan Environment, Almora, India; Ladakh Regional Centre, G.B. Pant National Institute of Himalayan Environment, Leh, India; Centre for Land & Water Resource Management, G. B. Pant National Institute of Himalayan Environment, Almora, India ABSTRACT ARTICLE HISTORY Received 27 September 2022 Understanding species richness and diversity patterns and their governing factors in less-to- Accepted 25 December 2022 unexplored regions across Himalaya provide invaluable insights into exploring drivers which shape as well as influence plant community structures. The present investigation explores plant KEYWORDS species richness and diversity patterns across different growth forms and its association with Alpine; species richness; environmental parameters along altitudinal gradient (3200 m-4800 m) in alpine regions of west diversity; Uttarakhand; Himalaya, India. A total of 265 plant taxa were documented from study area with higher Himalaya; India proportion of herbs (212), followed by shrubs (44) and trees (9). Species richness, diversity, and density patterns were estimated for each growth form along altitude gradients using polynomial regression and an apparent monotonically decreasing trend (p < 0.05) was seen across transects, with highest values for herbs. Beta diversity, estimated for each transect, was low in Darma for herbs exhibiting high species packaging and homogenous composition, and high in Mana showing more scope for occurrence of rare/occasional herbs. Four major distinct altitudinal zones were identified for Uttarakhand alpines using cluster dendrogram, i.e., 3200 to 3500 m, 3600 to 3900 m, 4000 to 4500 m and 4600 to 4800 m with respect to their vegetation composition. NMDS of combined dataset along altitude gradient across transects also exhib- ited proximity among lower altitudesof transects with similar species composition (like Anaphalis, Danthonia, Geranium, Pedicularis, Potentilla), while high-altitude plots were scattered towards both ends of axesinhabiting specialized plant species (like Gentiana, Nardostachys, Saussurea, Sedum, Swertia). The relationship between vegetation variables (richness, diversity and density) and climate variables was modelled using Pearson’s correlation (P < 0.001) and temperature, precipitation, and solar radiation exhibited positive correlation, while windspeed showed negative correlation. Relative effect of climatic parameters on species composition, analysed by CCA, showed strongest influence of precipitation in vegetation zones with high axes correlation, followed by temperature, isothermality and wind speed, while influence solar radiation was lowest. Thus, under the current climate change scenario, any change in these factors may alter the composition of these high-altitude area and threaten the unique flora as well as the fauna dependent on it. Hence, any effort made towards conservation would eventually benefit a significant proportion of Himalayan biodiversity. Significance statement The alpine regions of Himalaya have rich plant diversity and vary along with elevational gradient. Environmental parameters have played significant role on plant distribution and diversity. Studies on relation between the plant diversity and environmental factors are limited. Keeping in view the study has been conducted to analyse the plant diversity with possible environmental parameters. The observed variation in temperature and precipitation patterns along altitude gradient significantly correlates with the species richness and diversity patterns. The linear correlation states altitude as the variable which best explains the variations in richness and diversity patterns. Introduction of habitats inhabited by variable life-forms. Recent Mountains are recognized as biologically diverse eco- decades have witnessed a noticeable increment in bio- systems supporting high proportions of flora and diversity research especially in these cold biomes at fauna and harbouring a large number of protected high elevations. Several studies have demonstrated areas (Mallen-Cooper & Pickering, 2008). Owing to characteristic relationships between altitude, species the different topography, micro and macroclimatic richness patterns and composition, varying from conditions, mountain ecosystems consist of a range hump-shaped with maximum richness in mid- CONTACT K. Chandra Sekar kcsekar1312@rediffmail.com Garhwal Regional Centre, G.B. Pant National Institute of Himalayan Environment, Srinagar, Uttarakhand, India © 2023 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group on behalf of the International Water, Air & Soil Conservation Society(INWASCON). This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. 2 K. C. SEKAR ET AL. altitudes (Sharma et al., 2014; Vetaas & Grytnes, 2002) mounting anthropogenic pressure and over exploita- to monotonically decreasing (Ghafari et al., 2018; Kala tion of natural resources have made the Himalayan & Mathur, 2002), depending on whether parts or ecosystem one of the world’s most fragile and threa- whole gradient is taken into account (Rahbek, 1995). tened ecosystems (Bisht et al., 2022; Chandra Sekar & A variety of environmental factors change over short Srivastava, 2009; Murti, 2001). Any change or distur- distances along an altitude gradient, such as tempera- bance in the biotic and abiotic components of this ture, precipitation, solar radiation, and wind speed unique ecosystem is bound to cause imbalance in the which directly or indirectly influence plant diversity ecosystems. Due to remoteness, inaccessibility and and its composition (Cui et al., 2019; Khuroo et al., harsh climatic conditions these high-altitude areas 2010; Körner, 2007; Laiolo et al., 2018). Owing to this, have largely escaped the attention of ecologists, geo- plant occurrence and composition in mountain sys- graphers and natural resource managers. It is only in tem are more influenced by environmental conditions the recent decades that investigations have been taken (Körner, 2007; Rawal et al., 2018), thus making alti- up in high-altitude regions in Western Himalaya tude a suitable gradient for studying spatial variation detailing the plant diversity and composition in diversity and vegetation composition as well as the (Acharya et al., 2011; Raina & Sharma, 2012; Rawat effects of climate change. et al., 2010, 2010; Rawat, 2005). Himalaya, the youngest mountain ecosystem, is Under the global climate change scenario, globally recognized as biodiversity hotspot and inha- a comprehensive understanding of vegetation distribu- bits a rich repository of unique biodiversity tion and dynamics at the upper elevational limit of (Carpenter, 2005; Mallen-Cooper & Pickering, 2008). vascular plants across all life forms context for manage- The vast mountain range (over 2500 km long and 80 to ment planning and development of strategies for 300 km wide), rising from low-lying plains and reach- achieving is needed along with their responses to envir- ing upto 8000 m asl, provides altitude range exhibiting onmental parameters. Further, it provides necessary wide bioclimatic regimes (Vetaas & Grytnes, 2002). sustainable management of the ecosystem (Negi et al., The two broad climate regimes of Himalaya are mon- 2018). Keeping in view the above considerations and soonal and non-monsoonal. While the great the vast gap areas, the present study aims to assess Himalayan belt occurs under the Indian monsoon diversity and distribution patterns of different life regime characterized by glacial features like lateral forms along altitude gradient in alpine regions of and median moraines, the arid mountainous tracts in Uttarakhand, west Himalaya. This study has considered the extreme north and parallel to the Himalayan range multiple representative altitude transects spread across represent the non-monsoonal regime. These high- entire alpines of Uttarakhand Himalayas for providing altitude biomes exhibit extreme climatic conditions answers to following questions: (i) how does species such as diurnal temperature fluctuations, scanty rain- richness, diversity and density patterns in various life fall, and high wind velocity and are completely snow- forms vary along altitude gradient? (ii) Does vegetation bound for months. The soil is poor in organic matter, compositiondiffer considerably with altitude as well nitrogen content, sandy or sandy-loam in texture and across regions? (iii) How climatic factors influence a pH range of 7–11 (Körner, 2007; Negi, 2002). Owing plant species abundance in high-altitudes? to these extreme conditions, the region exhibitsa wide variety of ecological traitsalong with a number of Material and methods specialized habits such as cushion forming, diminutive and bushy (Bhattarai et al., 2010; Pandey et al., 2018). The study was conducted in high-altitude (>3000 m) Alpine vegetation starts from around 3200 m with alpine regions of Uttarakhand, west Himalaya. lower elevations (3200–3500 m) forming temperate Positioned in the rain shadow of the Himalaya, the alpine forests dominated by tree species such asAbies region remains unaffected by Indian monsoons, thus pindrow, Cedrus deodara and Pinus wallichiana. representing high-altitude non-monsoonal regime Towards higher elevations, tree species are unable to and exhibit extreme climate conditions, such as diur- cope with environmental stress and grow shorter and nal fluctuations in temperatures (−45ºC to 40ºC), often in lower densities and marks the treeline above scanty and erratic rainfall (<600 mm), high wind velo- which they are unable to grow. This zone mainly city and snowfall (Chandra Sekar et al., 2020). comprises alpine scrubs dominated by bushy habit Furthermore, region consists of arid mountainous with prostrate, woody branches and long deep roots tracts constituted of sediments of Tethyan sea bed like Caragana spp., Juniperus spp. and Lonicera spp., and sandy-loam with appreciable amount of clay and followed by alpine meadows in higher elevations poor organic content. The vegetation found in the dominated by herbaceous growth form including areas can be segregated into temperate forests with forbs, sedges, tussock, and non-tussock grasses.The open canopy and stunted growth (scrubs and GEOLOGY, ECOLOGY, AND LANDSCAPES 3 Figure 1. Study area (Source: http://srtm.csi.cgiar.org) and locations of seven studied altitude transects in alpine regions of Uttarakhand. meadows) (Murti, 2001). After several field visits and Sample design and vegetation assessment consultation of relevant literatures (Aswal & Intensive survey trips were carried out during 2014 to Mehrotra, 1994; Murti, 2001; Negi, 2002; Vashistha 2018 in selected transects in order to prepare et al., 2011), a study area map was extracted and six a baseline of plant species across all life forms (i.e., altitudes transects were selected for vegetation assess- trees, shrubs and herbs). Collection of plant specimen ment Figure 1 (see S1 Table): 4 K. C. SEKAR ET AL. was done in accordance with the permission of State Species richness, S = Total number of species � � Forest Department received from Chief Wildlife Shannon diversity index, H ¼ p lnp ; and i i Warden (Letter no. 917/5-6/DDN/dated i¼1 16 September 2013) and complying with the IUCN Beta diversity (β) was calculated using Whittaker Policy Statement Involving Species at Risk of (1960) formula as given in Mena and Vázquez Extinction. To support reproducibility, voucher speci- � Domínguez (2005): β ¼ 1; where α is the mean mens for all collected specimens were deposited in the number of species per altitude belt, and s is the total herbarium of GBP National Institute of Himalayan number of species recorded across the study system (i. Environment, Kosi, Almora (Acronym- GBP). e., altitude transect). The Shannon provides the diver- Information regarding collected plants list, their vou- sity of species in a community (i.e., altitudinal basis). cher specimen (Accession no.) and collector has been For each of the transect, values of selected environ- provided in Supplementary File (S1 Table) along with mental variables (average temperature, average preci- the complete list of plants species. Specimens were pitation, average solar radiation, average wind speed, collected as per the methods given in Jain and Rao average isothermality), which represent different direct and indirect gradients important for plant dis- (1977) and identified using local and regional floras tribution (Williams et al., 2012), were obtained along (Deva & Naithani, 1986; Gaur, 1999; Naithani, 1984; selected altitude ranges from WorldClim version 2.1 Osmaston, 1927; Pusalkar & Singh, 2012) and earlier climate data for 2000–2018 [https://www.worldclim. herbarium records (i.e., Botanical Survey of India, org/data/worldclim21.html] a spatial resolution of 30s Dehradun, Kolkata; Forest Research Institute – (~1 km ). Dehradun). Selected representative transects were investigated for compositional patterns of vegetation using strati- Data processing fied random sampling. Beginning from 3200 m, each transect was divided into 100 m altitude bands upto To numerically model distribution of different habit its respective upper elevational limits to carry out along altitude gradients, quadratic models were fitted vegetation assessment along altitude gradient. between altitude and species distributions indices Within each altitude band, three random sample (richness, diversity, density). Selection of the quadratic plots of 0.25 ha (50 × 50 m ) were established. In model was made based on the model performance that each sample plot, random sub-plots were laid, i.e., was primarily evaluated by computing statistically sig- 2 2 10 (10 × 10 m ) for trees, 5 (5 × 5 m ) for shrubs and nificant r values (P < 0.05). Moreover, to assess dis- saplings, and 10 (1 × 1 m ) for herbs and seedling. tribution of sample means from observations and For tree species, individuals with <10 cm girth and model simulations, analysis of variance (ANOVA) <20 cm height were considered as seedlings, those was carried out, particularly, estimated F-value of between 10–30 cm girth and >20 cm in height as each model was compared with critical F-value. The saplings and with >30 cm girth at breast height of result section highlights those models for which 1.37 m as trees. Number of individuals of plant spe- a significantly high r value was noted at P < 0.05. cies per quadrat was taken, pooled and analysed as Subsequently, results from ANOVA for these models per standard phytosociological approaches (Gairola were elaborated. et al., 2014; Goswami & Das, 2018; Mishra, 1968; Non-metric multidimensional scaling (NMDS) and Mueller-Dombois & Ellenberg, 1974; Rawat & permutational multivariate analysis of variance Everson, 2012; Zou et al., 2013) and Important (PERMANOVA, 999 permutations) were performed Value Index was calculated as per (Floeter et al., using R-vegan package (version 2.5–3) function 2004). “metaMDS,” to assess the community composition along various altitude transects, ordination plots Total number of individuals of aspecies in all quadrats Density ¼ were produced. Abbreviated scientific names of plant Total number of quadrats studied taxa used is documented in S2 Table. In order to select an optimum dimension for NMDS analyses for both Total number of quadrats in which species occur herbs and shrubs, stress-dimension analyses were car- Frequencyð%Þ ¼ Total number of quadrats studied ried out and dimension 2 was used for both NMDS � 100 analyses of shrubs and herbs using “Bray-Curtis” dis- tance. urther, influence of selected climatic parameters Total number of individuals of aspecies in all quadrats (temperature, precipitation, solar radiation, wind Abundance ¼ Total number of quadrats in which species occur speed, isothermality) on species richness, diversity and density was evaluated using Pearson’s correlation IVI = Relative Density + Relative Frequency + coefficient method to check significant relationships Relative Abundance GEOLOGY, ECOLOGY, AND LANDSCAPES 5 among selected parameters. Further, Canonical was 1.1 for trees, 2.0 for shrubs and 1.7 for herbs, Correspondence Analysis (CCA) was carried out to while, species to family (S/F) values were 1.8 for assess relationships between environmental para- trees, 3.1 for shrubs and 5.0 for herbs. Furthermore, meters and species composition in study transects. across the altitude zones, S/G and S/F values declined The community data matrices of plant species for all monotonically from lower to higher zones for trees transects were used with the constrained matrices and shrubs. However (S/F) ratio in 3501–4000 m zone composing of environmental parameters, (i) altitude, was higher (3.9) as compared to that in 3000–3500 m (ii) temperature, (iii) precipitation, (iv) solar radiation, (3.6) for herbs Table 1. (v) isothermality, (vi) wind speed. The nature of rela- At quadrate level, species richness was maximum in tionships between soil parameters and vegetation dis- Johar, for all growth forms, ranging, while Shannon tribution was depicted through ordination diagram. diversity Index (H’) was highest in Niti-Malari for The CCA analyses were carried out using the R-Vegan tress (1–1.05), Johar for shrubs (0.45–0.91) and package (version 2.5–3) function “CCA” and results Byans 1 for herbs (0.91–1.33) (S3 Figure). Broad were analysed with respect to inertia, ranks and eigen range of compositional attributes of vegetation in values of constrained matrices and parameters. study transects depicted higher densities of trees in Byans 1 &2, i.e., 20–240 ind/ha, while shrub and herb densities were higher in Darma (1200–3000ind/ha) and Byans 1 (31000–115000ind/ha), respectively (S4 Results Figure). Beta diversity values calculated for each life Floristic diversity pool form along altitude gradient exhibited an increasing trend in all transects, thus indicating scattered and A total of 265 vascular plants belonging to 155 genera heterogeneous plant composition especially at high and 55 families were observed along the investigated altitudes. Apart from this, beta diversity contributed altitude range (i.e., 3200–4800 m) of Uttarakhand significantly to transect level species richness of Byans, alpines (S2Table). Among these, nine were gymnos- Johar, Mana and Nelang for herbs and shrubs, but very perms distributed in six genera and three families, little for trees. However, in Darma tree species change while the rest belong to angiosperms. Asteraceae was rapidly while herbs and shrubs change slowly and in the most dominant family having 30 plant taxa fol- Niti-Malari, shrub species exhibited rapid change lowed by Rosaceae (21) and Ranunculaceae (18). In Table 2. terms of growth forms, herbaceous habit (81.7%) was dominant, followed by shrubs (14.9%) and trees (3.4%). In relation to 500 m altitude zones, plant Altitudinal relationship of compositional features population was maximumin the lowest zone, i.e., 3000–3500 m (72% of total flora), followed by 3501– Considering the alpine treeline area broadly varies 4000 m (62%), 4001–4500 m (38%) and 4501–4800 m from 3000–4000 m in the west Himalayas, treeline in (9%) Table 1, thus exhibiting a continuous decline in present study was identified at 3200–3900 m with 1–3 the number of plant species with increasing altitude. tree species per plot, 3–11 shrubs per plot and 9–30 There was a more rapid decline in distribution of herbs per plot). Patterns of species richness and diver- higher level of taxa (genera and family) with altitude sity of different growth forms across altitude range- nd as compared to that of species Table 1. The species to were modelled and a 2 order polynomial regression genera (S/G) value calculated for the entire landscape showed a significant fit (p < 0.05, Figure 2), with both Table 1. Floristic diversity pool (S- Species, G- Genus, F- Family) in high-altitude alpine region of Uttarakhand along altitude gradient. Trees Shrubs Herbs Altitudes (m) S G F S/G S/F S G F S/G S/F S G F S/G S/F 3000–3500 9 8 5 1.1 1.8 38 20 13 1.9 2.9 146 97 41 1.5 3.6 3501–4000 4 4 4 1.0 1.0 24 16 11 1.5 2.2 137 89 35 1.5 3.9 4001–4500 - - - - - 10 9 7 1.1 1.4 93 66 31 1.4 3.0 4501–5000 - - - - - 2 2 2 1.0 1.0 22 19 12 1.2 1.8 Total 9 8 5 1.1 1.8 40 20 12 2.0 3.1 219 127 44 1.7 5.0 Note: S- Species; G-Genera; F- Family. Table 2. Beta diversity variations across life forms and transects. Beta diversity in different transects Life forms Byans 1 Byans 2 Darma Johar Mana Nelang Niti-Malari Trees 1.86 1.67 2.54 1.53 1.89 2.33 1.67 Shrubs 3.82 5.45 1.52 2.91 6.54 4.26 3.89 Herbs 4.33 4.14 1.38 3.55 5.17 4.36 1.94 6 K. C. SEKAR ET AL. Figure 2. Species richness, diversity and density trends along altitude gradients in different life forms [A] Trees, [B] Shrubs, [C] Herbs in alpine of Uttarakhand, west Himalaya. richness and diversity monotonically decreasing with Betula utilis, Juniperus semigloboosa and Salix denti- increasing altitude. While the decrease in tree diversity culata were present in 3600–3900 m zone. Among was steep at altitude zone 3600–3800 m with no spe- shrubs, the species of Astragalus, Berberis, cies observed above 4000 m, it was moderate for herbs Cotoneaster and Salix were dominant in lower altitude and shrubs upto the upper hard boundaries. zones, while Cassiope fastigiata, Ephedra gerardiana, Furthermore, density distribution of different growth E. intermedia, Juniperus indica and Rhododendron formsalong altitude gradientfollowed a similar anthopogon are mostly dominated in high-altitudes decreasing pattern. For different size classes (i.e., (Table 3). Moreover, types of dominant herbaceous trees, saplings, seedlings), density decreased at higher plants highly varied across altitude gradient. At lower altitudes drastically, ranging from 10–540 ind/h for altitude zones several common species such as trees, 10–80 ind/h for saplings and 100–1000 ind/h Arenaria bryophylla, Bistorta affinis, Danthonia for seedlings (Figure 2) (S2 Figure). As for shrub and cachemyriana, Geranium wallichianum, Oxyria herb densities in relation to altitude, a relatively mod- digyna, Oxytropis lapponica, Potentilla spp., erate but significant decline (p < 0.05) was seen Taraxacum officinale and Thymus linearis were pre- (Figure 2). Based on the Important Value Index sent in fairly high proportions, whereas higher alti- (IVI) of plant taxa in each altitude, dominant and tudes exhibited clear dominance of Bistorta vivipara, codominant species across three growth forms are Carex nivalis, Gentiana stipitata, Geranium wallichia- listed in Table 3. Cluster dendrogram for the study num, Juncus concinnus and Meconopsis aculeata. area identified four major altitudinal zones based on species composition, i.e., 3200 to 3500 m, 3600 to Vegetation composition across different transects 3900 m, 4000 to 4500 m and 4600 to 4800 m (Figure 3). Lowest altitude zone (3200–3500 m) inhab- NMDS of vegetation abundance and combined dataset ited tree species such as Pinus wallichiana, Abies pin- of studied transects further exhibited a strong gradient drow, Prunus cornuta and Cedrus deodara, while separating sample (plots) and species between GEOLOGY, ECOLOGY, AND LANDSCAPES 7 Table 3. Dominant and co-dominant taxa along altitude gradient in studied alpine regions of Uttarakhand, west Himalaya. Dominant species and co-dominant taxa (IVI) Elevation Trees Shrubs Herbs 3200 Pinus wallichiana (151.0), Berberis jaeschkeana (31.1), Cassiope fastigiata (24.2), Bistorta affinis (12.9), Geranium wallichianum (10.8), Abies pindrow (57.5) Juniperus communis (23.2) Danthonia cachemyriana (9.8) 3300 Pinus wallichiana (110.6), Casragana versicolor (32.4), Berberis jaeschkeana (27.4), Geranium wallichianum (11.4), Taraxacum officinale Abies pindrow (96.3) Cotoneaster microphyllus (21.7) (9.8), Bistorta affinis (9.3) 3400 Pinus wallichiana (159.5), Juniperus communis (33.0), Berberis jaeschkeana (28.5), Thymus linearis (15.5), Poa alpina (15.3), Danthonia Betula utilis (81.4) Cotoneaster microphyllus (28.5) cachemyriana (11.1) 3500 Pinus wallichiana (176.0), Caragana versicolor (37.0), Juniperus communis (34.6), Geranium wallichianum (14.1), Danthonia Betula utilis (70.1) Berberis umbellate (25.5), Juniperus indica (25.5) cachemyriana (11.0), Poa alpina (10.6) 3600 Pinus wallichiana (143.2), Juniperus indica (53.0), Cotoneaster microphyllus (29.7), Potentilla argyrophylla var. atrosanguinea (16.9), Poa Betula utilis (112.5) Caragana versicolor (26.8) alpina (11.7), Thymus linearis (10.2) 3700 Betula utilis (146.9), Salix Caragana versicolor (29.1), Juniperus indica (26.0), Potentilla argyrophylla var. atrosanguinea (18.0), denticulata (73.4) Astragalus oplites (25.4), Salix flagellaris (25.4) Taraxacum officinale (12.7), Thymus linearis (12.5) 3800 Betula utilis (225.5), Myrtama elegans (40.0), Hyssopus officinalis (3.5), Poa alpina (21.4), Thymus linearis (17.3), Oxyria Juniperus semiglobosa Astragalus oplites (28.7), Caragana versicolor (28.7) digyna (15.3) (74.5) 3900 Betula utilis (300.0) Juniperus indica (55.0), Cassiope fastigiata (39.0), Danthonia cachemyriana (15.7), Bromus japonicus Berberis jaeschkeana (34.4) (14.9), Potentilla argyrophylla (14.6) 4000 Cassiope fastigiata (53.7), Berberis jaeschkeana (34.0), Poa alpina (21.5), Oxytropis lapponica (16.8), Rhododendron anthopogon (34.0) Potentilla argyrophylla var. atrosanguinea (16.2) 4100 Cassiope fastigiata (64.9), Juniperus indica (56.7), Arenaria bryophylla (15.3), Euphrasia himalayica Caragana versicolor (39.5) (14.7), Poa alpina (14.5) 4200 Cassiope fastigiata (84.5), Rhododendron anthopogon Bistorta affinis (18.7), Bergenia stracheyi(16.4), Carex (54.8), Juniperus communis (52.1) nubigeana (14.2), Saxifraga flagellaris (14.2) 4300 Juniperus indica (117.9), Caragana versicolor (72.6), Potentilla argyrophylla (23.3), Bistorta affinis (21.9), Ephedra intermedia (72.6) Poa alpina (18.1) 4400 Juniperus indica (181.7), Ephedra gerardiana (118.3) Poa alpina (32), Geramiun wallichianum (31), Aconogonum tortusom (18) 4500 Juniperus indica (170.0), Ephedra gerardiana (130.0) Geranium wallichianum (44.6), Poa alpina (35.1), Potentilla argyrophylla var. atrosanguinea (27.9) 4600 Juniperus indica (300) Geranium wallichianum (95.2), Bistorta vivipara (24.9), Meconopsis aculeata (24.9), Poa alpina (24.9) 4700 Ephedra intermedia (300) Juncus concinnus (96.2), Carex nivalis (26.9), Gentiana stipitata (26.9), Poa alpina (26.9) 4800 Geranium wallichianum (100.0), Allardia glabra (27.8), Bromus inermis (27.8) transects as well as altitude range along the first two and higher altitude plots (4400–4800 m) in Mana axis (NMDS1 and NMDS2) (Figure 4). Separation formed distinct cluster with Darma transect showing along the first axis explained 40% of variation proximity in their species composition inhabiting while second axis explained 20%, thus underscored Cotoneaster spp., Convolvulus arvensis, Polygonum the affinity between different altitudinal vegetation polystachyum, Rosa macrophylla. Mid-altitude zones zones. Across all transects, distinct clusters denoting in Mana, i.e., 3600–3800 m, showed similarity with vegetations zones were identified characterized by spe- Nelang, while 3900–4300 m zone was scattered along cific species. Nelang transect formed a cluster towards positive end of first axisalong with species like Galium the negative end of first axis, with plotsat higher alti- asperuloides Nardostachys jatamansi, Picrorhiza kur- tude (3900–4400 m) in more proximity with each roa, Poa compressa and Saussurea simpsoniana. other inhabiting speciessuch as Aconogonum tortuo- Overall, sample plots in lower altitudes of transects sum, Melica persica, Poa tibetica, Sisymbrium brassici- were more clustered towards the middle of both axes, forme, Urtica hyperborea. Lower and mid-altitude inhabiting common plant species. Most of the sample plots of Byans 1 (3200–3700 m) and Byans 2 (3200– plots in higher altitudes occupied extreme ends (posi- 3800 m) as well as NitiMalari transect plots formed tive and negative) of both axes, thus showing increase a separate cluster towards negative end of second axis, in rarity in species composition moving from low to inhabited by similar species (such as Carex nivalis, high altitudes. Delphinium brunonianum, Primula denticulata, Rheum webbianum, U. dioca). On contrary, higher Relationship of vegetation composition with altitude plots in Byans 1 (>3800 m) & Byans 2 environmental variables (>3900 m) formed a distinct cluster towards positive end of first axis characterised by species such as Correlation analysis among climate variables and Christolea himalayensis, Leontopodium brachyactis, plant diversity indices is depicted in Figure 5 along Ranunculus palmatifidus, Saxifraga flagellaris. For with Pearson’s correlation coefficient and significance Johar, study plots were scattered across the four quad- level of relation. As established earlier also, altitude rants, thus exhibited large variations in species com- has a significant effect on density, diversity, and rich- position along altitude gradient. Lower (3200–3500 m) ness of plant species (p < 0.001), with an apparent 8 K. C. SEKAR ET AL. Figure 3. Hierarchical cluster dendrogram showing distinct altitudinal zones with respect to the species composition in alpine region of Uttarakhand, west Himalaya. decrease in density (r= -0.77), diversity (r = −0.87), Byans 1 & 2 due to the high wind speed, low tempera- and richness (r = −0.70). Similarly, there was strong ture and precipitation species like Christolea hima- negative correlation (p < 0.001) of temperature (r = layensis, Gentiana stipitata, Leontopodium brachyatis, −0.96), precipitation (r = −0.77), and solar radiation Rhodiola bupleuroides, Saussurea gossypiphora, (r = −0.71) with altitude as well, while it was positively S. simpsoniana and Sibbaldia parviflora. Contrary to significant with wind speed (r = 0.83). Apart from this, species like Anaphalis nepalensis, A. triplinivalis, altitude, temperature and precipitation also made Berberis umbellata, Carum carvi, Cotoneaster rotundi- a strong gradient influencing plant species richness folius and Ribes orientale favoured lower altitude and diversity in alpine regions Figure 5. Increase in transects like Darma and Johar with high temperature temperature as well as precipitation favoured com- and precipitation. Occupying positive end of first axis paratively more number as well as variety of species and negative end of second axis, Nelang transect in study area, which is evident with significantly posi- exhibited very little dependence on temperature, pre- tive correlation coefficient values (p <0.01). As indi- cipitation and wind speed. In context of plant species, cated by r values, solar radiation and wind speed the site thus occupies herbs like Aconogonum tortu- exhibited significant (p < 0.05) but weak correlation sum, Rumex nepalensis, Urtica hyperborea and with richness and diversity indices, thus exhibiting Verbascum thapsus. Furthermore, isothermality of little influence on presence and types of plant taxa in study site favoured growth and abundance of species alpine. like Aconitum balfourii, Gentiana argentea, The relative effect of climatic parameters on plant G. leucomelaena and Pedicularis tubiformis. species composition, analysed by CCA, showed stron- gest influence of precipitation in forming altitudinal Discussion zones having specific vegetation composition with high axes correlation (Figure 6), followed by tempera- The results of our study are based on data gathered ture, isothermality, and wind speed, while influence of during a time period from 2014 to 2018 of field work. solar radiation was lowest. Total inertia of combined A baseline floristic survey of study area (field and data set was 13.02 where constrained climatic para- literature based) documented a total of 932 plant meters explained 48% of total variances on first axis taxa belonging to 371 genera in 76 families Chandra and 30% on second axis. Among altitude transects, in Sekar et al., 2022. Hence, in comparison with previous GEOLOGY, ECOLOGY, AND LANDSCAPES 9 Figure 4. Non-metric Multi-Dimensional Scaling (NMDS) representing species composition among sample plots along altitude gradients of study transect. 10 K. C. SEKAR ET AL. Figure 5. Pearson’s Correlation between plant richness, density and diversity indices with climate variables. studies (Negi, 2002), the high-altitude alpine zones their abundance in different communitieswith (3200–4800 m) of Uttarakhand accounts for 58% extreme conditions (Abbas et al., 2019). alpine flora of west Himalayas. While comparing The patterns of S/G along altitude gradient are used with pan-Himalayas (Rana & Rawat, 2017), the study to describe speciation and diversification rates (Floeter area inhabits 10% species, 16% genera, and 32% et al., 2004). The general altitudinal decrease in S/G as families. Furthermore, 47 plants were under different well as S/F recorded in our study indicates their phy- threat categories as per IUCN and CAMP (Ved et al., logenetic dispersion towards higher altitudes. On 2003). Among these, Dactylorhiza hatagirea, comparing the overall S/G values of different life Nardostachys jatamansi and Picrorhiza kurroa were forms, the higher ratio for shrubs indicates more “Critically Endangered,” while Aconitum heterophyl- intense diversification within the genera as compared lum, Angelica glauca, Ephedra gerardiana, Fritillaria to trees and herbs. Similarly, within family herbs exhi- roylei, Saussurea obvallata and Podophyllum hexan- bit significantly more intense phylogenetic clumping drum were “Endangered.” Hence, any effort made as compared to others. The increasing values of beta towards conservation ofthis high-altitude zone would diversity calculated along altitude gradient for each eventually benefit a significant proportion of transect indicates heterogenous plant composition, Himalayan biodiversity including flora as well as thus exhibiting more scope for occurrence of rare fauna. Dominance of herbaceous growth form in taxa as compared to lower altitudes (Rawal et al., terms of richness, diversity as well as density is appar- 2018). This can be attributed to the variable levels of ent due to extreme climatic conditions in the region environmental as well as anthropogenic pressure pre- which causes poor assemblages of trees and shrubs vailing in the different study areas. Relatively high (generally high stature species). Apart from this, dom- level of disturbance and poor species pool leads to inance of families such as Asteraceae and Rosaceae can lower beta diversity in herbs, while it exhibits be attributed to their specialized morphology, and a reverse trend for trees. Among the studied transects, broad ecological amplitude enabling them to establish Darma with low β diversity exhibits high species GEOLOGY, ECOLOGY, AND LANDSCAPES 11 Figure 6. Canonical Correspondence Analysis of combined data set: ordination of sample pots and vegetation abundance constrained by their relationships to environmental variables. Significant parameters are shown as biplot vector. packaging and homogenous composition in case of season paving way for plant species tolerant to herbs, while becomes highly heterogeneous for trees extreme conditions, a phenomenon known as “ecolo- (high β diversity). Similarly, with high beta diversity of gical filtering.” These fluctuations also influence soil herbs in Mana transect there is more scope for occur- texture, nutrients and stability of substrate which in rence of rare/occasional herbaceous species, which turn are responsible for maintaining vegetation com- contributes for higher heterogeneity of species position (Sanchez-Gonzalez & Lopez-Mata, 2005). composition. Hence, forming distinct vegetation zones along alti- Several studies have demonstrated characteristic tude, with tree species occupying the lowest zone relationships between altitude and patterns of species along with woody anddeep-rooted shrubs like richness, diversity, and compositional features (such Caragana spp., Juniperus spp. and Lonicera spp., fol- as density, total basal area) along a gradient (Acharya lowed by meadows at higher elevations dominated by et al., 2011; Rawat et al., 2010; Sharma & Raina, 2013). herbaceous growth form including forbs, sedges, tus- In present study also, density and altitude exhibited sock and non-tussock grasses. highly significant relationship (p < 0.05, F>F ) in Ordination analysis (NMDS) clearly exhibited var- crit entire study zone. This signifies the strong dependence iation in vegetation composition among transects of floristic diversity and density with altitude, irrespec- (Figure 4). While temperature was the major factor tive of the growth stage or life forms in the alpine responsible for variation along altitude, among trans- region. Our results are congruent with several former ects precipitation had the strongest influence on vege- studies (Kala & Mathur, 2002; Pandey et al., 2018; tation composition, followed by temperature, wind Rawal et al., 2018; Sharma et al., 2014; Sinha et al., speed and isothermality as depicted by CCA analysis. 2018; Vetaas & Grytnes, 2002; Wang et al., 2006). Byans and Niti-Malari regions with cold, dry climate Reduction of plants in higher altitude may be attrib- and high wind speed had similar species composition. uted to eco-physiological constraints, such as extre- Nelang region formed a separate cluster due to its mely low temperature, precipitation and solar variable climate such as comparatively low tempera- radiation (Figure 5) which results in short growing ture, precipitation, isothermality and wind speed. 12 K. C. SEKAR ET AL. Darma and Johar fell in the same cluster with com- documents the plant species diversity and composition paratively warm and wet climate experiencing slower of remote areas of Himalaya which are known to be winds and optimum isothermality. natural laboratories to study the responses of plants to While comparing to earlier literature (Chandra climate change. While our study is more focussed on Sekar et al., 2015; Garbyal et al., 2007; Lohani et al., plant diversity, it can be expanded to broader aspects 2013; Mitra et al., 2017; Rawat, 2005; Uniyal et al., such as phenological and physiological studies of alpine 2002), the density of Aconitum heterophyllum, plants and their relation with climatic and edaphic Allium stracheyi, Bergenias tracheyi, Carum carvi, factors (Majeed et al., 2022) and opt more towards Hippophae salicifolia, Rhododendron anthopogon, R. plant distribution models (Bayat et al., 2021; Jahani & campanulatum, Swertia ciliata and Thymus linearis Saffariha, 2020; Jahani et al., 2021; Mosaffaei et al., was lesser, while that of Juniperus communis, 2020). J. indica, Polygonatum verticillatum density was increased. This can be attributed to the irregular har- Conclusion vesting of former species for fuel and medicinal pur- poses. The population of Betula utilis highly decreased This study is the first attempt to document and analyse in the area due to its utilization for fuelwood in higher floristic composition of high-altitude alpine regions Himalayan regions (Mitra et al., 2017). Climatic bar- panning across Uttarakhand, west Himalaya, along riers, limiting tree habits, resulting limited density altitude gradient and determine major climate drivers coupled with high extraction as fuelwood may be the influencing vegetation patterns. Hierarchical clustering reason of population reduction. Furthermore, trees as well as ordination analysis (NMDS and CCA) per- and saplings of B. utilis are available, but there are formed also showed concordant results, underscoring no seedlings recorded in 3500 m. Thus, protection of altitudinal vegetation zonation throughout study area these saplings is need of the hour, so that the popula- as well as its relationship with environmental variables. tion of B. utilis can be saved. In view of importance of Phytosociological studies undertaking multiple trans- the tree, the local considered it as “sacred” and various ects and life forms in high altitude alpine regions are conservation activities were observed. The people of very scarce especially in Indian Himalayan region. In the local areas must aware about the importance of this context, the present study also attempts to discuss plants and some good initiatives (i.e., J. semiglobosa and compare distribution patterns of vegetation across conservation in Johar) should be replicated in all other the longitudinal gradients along multiple altitudes regions, to conserve the plant diversity. Furthermore, transect. In this context, the present study gives com- density and diversity are also influenced by frequency parable ecological data sets to discuss distribution pat- of anthropogenic disturbances, i.e., road construction, tern of seven altitudinal transects. Species richness and habitation, commercial plantation and agricultural diversity along the altitude gradient in alpine region performs also results in replacement of natural vegeta- exhibited a monotonic decrease, both overall and by tion with man-made ecosystem. growth forms, with maximum values in the lower Under current global climate change scenarios, altitude zones (3200–3500 m). Beta diversity also a number of studies have projected changes in vegeta- increased with altitude thus exhibiting higher hetero- tion distribution and upward shifts in plant boundaries geneity and presence of unique species adapted to (Kuhn et al., 2016; Rai et al., 2010; Rosset et al., 2010; survive the extreme environmental conditions. Telwala et al. 2013). These shifts tend to change the Among growth forms, beta diversity was generally shape of ecosystems of high-altitude regions. However, higher for herbs and shrubs as compared to tress indi- shifting rate varies with species depending on their cating homogeneity in tree populations at high-altitude altitudinal and geographical distribution ranges, and alpines. The observed variations in temperature and their accumulative potential (Klimes et al., 2003). As precipitation patterns along altitude gradient signifi- species distribution range decrease with increasing alti- cantly correlates with the species richness and diversity tude, species with narrow amplitude of altitudinal dis- patterns, with high temperature and precipitation sup- tribution and endemics are more affected by porting plant richness and diversity. The linear correla- environmental or anthropogenic disturbances. While tion states altitude as the variable which best explains climate change (natural or anthropogenic) threatens the variations in richness and diversity patterns. high-altitude flora, it also impacts a significant portion of useful biodiversity and traditional livelihood of local people. Hence it becomes pivotal to carry out empirical Acknowledgements studies pertaining to potential extent and magnitude of The authors thankfully acknowledge the facilities provided current and future shifts in altitudinal limits of plants by Director, G.B. Pant National Institute of Himalayan in response to climate change in order to draw con- Environment (NIHE), Kosi-Katarmal, Almora for undertak- clusive conservation and management strategies ing this work. We are gratefully acknowledging the financial (Mehta et al., 2020; Pandey et al., 2019). Our study assistance received from Council of Scientific and Industrial GEOLOGY, ECOLOGY, AND LANDSCAPES 13 Research [EMR No. 38 (1346)/12/EMR-II], Kailash Sacred Upadhyay, S., & Dey, D. (2022). Influence of anthropo- Landscape Conservation and Development Initiative and genic pressure on the plant species richness and diversity National Mission on Himalayan Studies (NMHS), Ministry along the elevation gradients of Indian Himalayan of Environment, Forest and Climate Change, Government of high-altitude protected areas. Frontiers in Ecology and India (GBPNI/NMHS-2018-19/SG 11) and National Mission Evolution, 10, 751989. https://doi.org/10.3389/fevo.2022. on Sustaining the Himalayan Ecosystem (NMSHE), TF-3, 751989 DST, New Delhi. Permissions for Herbarium and library Carpenter, C. (2005). The environmental control of plant consultation received from different organizations, i.e., species density on a Himalayan elevation gradient. Botanical Survey of India, Forest Research Institute, Journal of Biogeography, 32(6), 999–1018. https://doi. Wildlife Institute of India, National Botanical Research org/10.1111/j.1365-2699.2005.01249.x Institute; Permission of State Forest Department received Chandra Sekar, K., Lalit, G., Aseesh, P., & Srivastava, S. K. from Chief Wildlife Warden (Letter no. 917/5-6/DDN/ (2015). A note on distribution of juniperus semiglobosa in dated 16 Sep. 2013); logistic and field guidance received Uttarakhand, India. Indian Journal of Forestry, 38(1), from DG, ITBP (Letter no. 2766 dated 14.08.2014) are highly 79–80. https://doi.org/10.54207/bsmps1000-2015- acknowledged. Dr. Poonam Tripathi, International Centre 0XY6MS for Integrated Mountain Development (ICIMOD), Nepal is Chandra Sekar, K., Pandey, A., Giri, L., Joshi, B. C., Bhatt, gratefully acknowledged for preparing study area map. The D., Bhojak, P., Dey, D., Thapliyal, N., Bisht, K., Bisht, M., inputs received from anonymous reviewers aided in improv- & Negi, V. S. (2022). Floristic diversity in Cold Desert ing contents of this manuscript, we highly thank them. regions of Uttarakhand Himalaya, India. Phytotaxa, 537 (1), 1–62. Chandra Sekar, K., & Srivastava, S. K. (2009). Flora of the Disclosure statement Pin Valley National Park, Himachal Pradesh. Botanical Survey of India. Ministry of Environment and Forests. No potential conflict of interest was reported by the authors. Cui, Y., Bing, H., Fang, L., Wu, Y., Yu, J., Shen, G., Jiang, M., Wang, X., & Zhang, X. (2019). Diversity patterns of the rhizosphere and bulk soil microbial communities along ORCID an altitudinal gradient in an alpine ecosystem of the east- ern Tibetan Plateau. Geoderma, 338, 118–127. https://doi. K. 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Plant species diversity and density patterns along altitude gradient covering high-altitude alpine regions of west Himalaya, India

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GEOLOGY, ECOLOGY, AND LANDSCAPES INWASCON https://doi.org/10.1080/24749508.2022.2163606 RESEARCH ARTICLE Plant species diversity and density patterns along altitude gradient covering high-altitude alpine regions of west Himalaya, India a a b b c K. Chandra Sekar , Neha Thapliyal , Aseesh Pandey , Bhaskar Joshi , Sandipan Mukherjee , b b b d b Puja Bhojak , Monica Bisht , Deepika Bhatt , Sourab Singh and Amit Bahukhandi a b Garhwal Regional Centre, G.B. Pant National Institute of Himalayan Environment, Srinagar, India; Centre for Biodiversity Conservation and Management, G. B. Pant National Institute of Himalayan Environment, Almora, India; Ladakh Regional Centre, G.B. Pant National Institute of Himalayan Environment, Leh, India; Centre for Land & Water Resource Management, G. B. Pant National Institute of Himalayan Environment, Almora, India ABSTRACT ARTICLE HISTORY Received 27 September 2022 Understanding species richness and diversity patterns and their governing factors in less-to- Accepted 25 December 2022 unexplored regions across Himalaya provide invaluable insights into exploring drivers which shape as well as influence plant community structures. The present investigation explores plant KEYWORDS species richness and diversity patterns across different growth forms and its association with Alpine; species richness; environmental parameters along altitudinal gradient (3200 m-4800 m) in alpine regions of west diversity; Uttarakhand; Himalaya, India. A total of 265 plant taxa were documented from study area with higher Himalaya; India proportion of herbs (212), followed by shrubs (44) and trees (9). Species richness, diversity, and density patterns were estimated for each growth form along altitude gradients using polynomial regression and an apparent monotonically decreasing trend (p < 0.05) was seen across transects, with highest values for herbs. Beta diversity, estimated for each transect, was low in Darma for herbs exhibiting high species packaging and homogenous composition, and high in Mana showing more scope for occurrence of rare/occasional herbs. Four major distinct altitudinal zones were identified for Uttarakhand alpines using cluster dendrogram, i.e., 3200 to 3500 m, 3600 to 3900 m, 4000 to 4500 m and 4600 to 4800 m with respect to their vegetation composition. NMDS of combined dataset along altitude gradient across transects also exhib- ited proximity among lower altitudesof transects with similar species composition (like Anaphalis, Danthonia, Geranium, Pedicularis, Potentilla), while high-altitude plots were scattered towards both ends of axesinhabiting specialized plant species (like Gentiana, Nardostachys, Saussurea, Sedum, Swertia). The relationship between vegetation variables (richness, diversity and density) and climate variables was modelled using Pearson’s correlation (P < 0.001) and temperature, precipitation, and solar radiation exhibited positive correlation, while windspeed showed negative correlation. Relative effect of climatic parameters on species composition, analysed by CCA, showed strongest influence of precipitation in vegetation zones with high axes correlation, followed by temperature, isothermality and wind speed, while influence solar radiation was lowest. Thus, under the current climate change scenario, any change in these factors may alter the composition of these high-altitude area and threaten the unique flora as well as the fauna dependent on it. Hence, any effort made towards conservation would eventually benefit a significant proportion of Himalayan biodiversity. Significance statement The alpine regions of Himalaya have rich plant diversity and vary along with elevational gradient. Environmental parameters have played significant role on plant distribution and diversity. Studies on relation between the plant diversity and environmental factors are limited. Keeping in view the study has been conducted to analyse the plant diversity with possible environmental parameters. The observed variation in temperature and precipitation patterns along altitude gradient significantly correlates with the species richness and diversity patterns. The linear correlation states altitude as the variable which best explains the variations in richness and diversity patterns. Introduction of habitats inhabited by variable life-forms. Recent Mountains are recognized as biologically diverse eco- decades have witnessed a noticeable increment in bio- systems supporting high proportions of flora and diversity research especially in these cold biomes at fauna and harbouring a large number of protected high elevations. Several studies have demonstrated areas (Mallen-Cooper & Pickering, 2008). Owing to characteristic relationships between altitude, species the different topography, micro and macroclimatic richness patterns and composition, varying from conditions, mountain ecosystems consist of a range hump-shaped with maximum richness in mid- CONTACT K. Chandra Sekar kcsekar1312@rediffmail.com Garhwal Regional Centre, G.B. Pant National Institute of Himalayan Environment, Srinagar, Uttarakhand, India © 2023 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group on behalf of the International Water, Air & Soil Conservation Society(INWASCON). This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. 2 K. C. SEKAR ET AL. altitudes (Sharma et al., 2014; Vetaas & Grytnes, 2002) mounting anthropogenic pressure and over exploita- to monotonically decreasing (Ghafari et al., 2018; Kala tion of natural resources have made the Himalayan & Mathur, 2002), depending on whether parts or ecosystem one of the world’s most fragile and threa- whole gradient is taken into account (Rahbek, 1995). tened ecosystems (Bisht et al., 2022; Chandra Sekar & A variety of environmental factors change over short Srivastava, 2009; Murti, 2001). Any change or distur- distances along an altitude gradient, such as tempera- bance in the biotic and abiotic components of this ture, precipitation, solar radiation, and wind speed unique ecosystem is bound to cause imbalance in the which directly or indirectly influence plant diversity ecosystems. Due to remoteness, inaccessibility and and its composition (Cui et al., 2019; Khuroo et al., harsh climatic conditions these high-altitude areas 2010; Körner, 2007; Laiolo et al., 2018). Owing to this, have largely escaped the attention of ecologists, geo- plant occurrence and composition in mountain sys- graphers and natural resource managers. It is only in tem are more influenced by environmental conditions the recent decades that investigations have been taken (Körner, 2007; Rawal et al., 2018), thus making alti- up in high-altitude regions in Western Himalaya tude a suitable gradient for studying spatial variation detailing the plant diversity and composition in diversity and vegetation composition as well as the (Acharya et al., 2011; Raina & Sharma, 2012; Rawat effects of climate change. et al., 2010, 2010; Rawat, 2005). Himalaya, the youngest mountain ecosystem, is Under the global climate change scenario, globally recognized as biodiversity hotspot and inha- a comprehensive understanding of vegetation distribu- bits a rich repository of unique biodiversity tion and dynamics at the upper elevational limit of (Carpenter, 2005; Mallen-Cooper & Pickering, 2008). vascular plants across all life forms context for manage- The vast mountain range (over 2500 km long and 80 to ment planning and development of strategies for 300 km wide), rising from low-lying plains and reach- achieving is needed along with their responses to envir- ing upto 8000 m asl, provides altitude range exhibiting onmental parameters. Further, it provides necessary wide bioclimatic regimes (Vetaas & Grytnes, 2002). sustainable management of the ecosystem (Negi et al., The two broad climate regimes of Himalaya are mon- 2018). Keeping in view the above considerations and soonal and non-monsoonal. While the great the vast gap areas, the present study aims to assess Himalayan belt occurs under the Indian monsoon diversity and distribution patterns of different life regime characterized by glacial features like lateral forms along altitude gradient in alpine regions of and median moraines, the arid mountainous tracts in Uttarakhand, west Himalaya. This study has considered the extreme north and parallel to the Himalayan range multiple representative altitude transects spread across represent the non-monsoonal regime. These high- entire alpines of Uttarakhand Himalayas for providing altitude biomes exhibit extreme climatic conditions answers to following questions: (i) how does species such as diurnal temperature fluctuations, scanty rain- richness, diversity and density patterns in various life fall, and high wind velocity and are completely snow- forms vary along altitude gradient? (ii) Does vegetation bound for months. The soil is poor in organic matter, compositiondiffer considerably with altitude as well nitrogen content, sandy or sandy-loam in texture and across regions? (iii) How climatic factors influence a pH range of 7–11 (Körner, 2007; Negi, 2002). Owing plant species abundance in high-altitudes? to these extreme conditions, the region exhibitsa wide variety of ecological traitsalong with a number of Material and methods specialized habits such as cushion forming, diminutive and bushy (Bhattarai et al., 2010; Pandey et al., 2018). The study was conducted in high-altitude (>3000 m) Alpine vegetation starts from around 3200 m with alpine regions of Uttarakhand, west Himalaya. lower elevations (3200–3500 m) forming temperate Positioned in the rain shadow of the Himalaya, the alpine forests dominated by tree species such asAbies region remains unaffected by Indian monsoons, thus pindrow, Cedrus deodara and Pinus wallichiana. representing high-altitude non-monsoonal regime Towards higher elevations, tree species are unable to and exhibit extreme climate conditions, such as diur- cope with environmental stress and grow shorter and nal fluctuations in temperatures (−45ºC to 40ºC), often in lower densities and marks the treeline above scanty and erratic rainfall (<600 mm), high wind velo- which they are unable to grow. This zone mainly city and snowfall (Chandra Sekar et al., 2020). comprises alpine scrubs dominated by bushy habit Furthermore, region consists of arid mountainous with prostrate, woody branches and long deep roots tracts constituted of sediments of Tethyan sea bed like Caragana spp., Juniperus spp. and Lonicera spp., and sandy-loam with appreciable amount of clay and followed by alpine meadows in higher elevations poor organic content. The vegetation found in the dominated by herbaceous growth form including areas can be segregated into temperate forests with forbs, sedges, tussock, and non-tussock grasses.The open canopy and stunted growth (scrubs and GEOLOGY, ECOLOGY, AND LANDSCAPES 3 Figure 1. Study area (Source: http://srtm.csi.cgiar.org) and locations of seven studied altitude transects in alpine regions of Uttarakhand. meadows) (Murti, 2001). After several field visits and Sample design and vegetation assessment consultation of relevant literatures (Aswal & Intensive survey trips were carried out during 2014 to Mehrotra, 1994; Murti, 2001; Negi, 2002; Vashistha 2018 in selected transects in order to prepare et al., 2011), a study area map was extracted and six a baseline of plant species across all life forms (i.e., altitudes transects were selected for vegetation assess- trees, shrubs and herbs). Collection of plant specimen ment Figure 1 (see S1 Table): 4 K. C. SEKAR ET AL. was done in accordance with the permission of State Species richness, S = Total number of species � � Forest Department received from Chief Wildlife Shannon diversity index, H ¼ p lnp ; and i i Warden (Letter no. 917/5-6/DDN/dated i¼1 16 September 2013) and complying with the IUCN Beta diversity (β) was calculated using Whittaker Policy Statement Involving Species at Risk of (1960) formula as given in Mena and Vázquez Extinction. To support reproducibility, voucher speci- � Domínguez (2005): β ¼ 1; where α is the mean mens for all collected specimens were deposited in the number of species per altitude belt, and s is the total herbarium of GBP National Institute of Himalayan number of species recorded across the study system (i. Environment, Kosi, Almora (Acronym- GBP). e., altitude transect). The Shannon provides the diver- Information regarding collected plants list, their vou- sity of species in a community (i.e., altitudinal basis). cher specimen (Accession no.) and collector has been For each of the transect, values of selected environ- provided in Supplementary File (S1 Table) along with mental variables (average temperature, average preci- the complete list of plants species. Specimens were pitation, average solar radiation, average wind speed, collected as per the methods given in Jain and Rao average isothermality), which represent different direct and indirect gradients important for plant dis- (1977) and identified using local and regional floras tribution (Williams et al., 2012), were obtained along (Deva & Naithani, 1986; Gaur, 1999; Naithani, 1984; selected altitude ranges from WorldClim version 2.1 Osmaston, 1927; Pusalkar & Singh, 2012) and earlier climate data for 2000–2018 [https://www.worldclim. herbarium records (i.e., Botanical Survey of India, org/data/worldclim21.html] a spatial resolution of 30s Dehradun, Kolkata; Forest Research Institute – (~1 km ). Dehradun). Selected representative transects were investigated for compositional patterns of vegetation using strati- Data processing fied random sampling. Beginning from 3200 m, each transect was divided into 100 m altitude bands upto To numerically model distribution of different habit its respective upper elevational limits to carry out along altitude gradients, quadratic models were fitted vegetation assessment along altitude gradient. between altitude and species distributions indices Within each altitude band, three random sample (richness, diversity, density). Selection of the quadratic plots of 0.25 ha (50 × 50 m ) were established. In model was made based on the model performance that each sample plot, random sub-plots were laid, i.e., was primarily evaluated by computing statistically sig- 2 2 10 (10 × 10 m ) for trees, 5 (5 × 5 m ) for shrubs and nificant r values (P < 0.05). Moreover, to assess dis- saplings, and 10 (1 × 1 m ) for herbs and seedling. tribution of sample means from observations and For tree species, individuals with <10 cm girth and model simulations, analysis of variance (ANOVA) <20 cm height were considered as seedlings, those was carried out, particularly, estimated F-value of between 10–30 cm girth and >20 cm in height as each model was compared with critical F-value. The saplings and with >30 cm girth at breast height of result section highlights those models for which 1.37 m as trees. Number of individuals of plant spe- a significantly high r value was noted at P < 0.05. cies per quadrat was taken, pooled and analysed as Subsequently, results from ANOVA for these models per standard phytosociological approaches (Gairola were elaborated. et al., 2014; Goswami & Das, 2018; Mishra, 1968; Non-metric multidimensional scaling (NMDS) and Mueller-Dombois & Ellenberg, 1974; Rawat & permutational multivariate analysis of variance Everson, 2012; Zou et al., 2013) and Important (PERMANOVA, 999 permutations) were performed Value Index was calculated as per (Floeter et al., using R-vegan package (version 2.5–3) function 2004). “metaMDS,” to assess the community composition along various altitude transects, ordination plots Total number of individuals of aspecies in all quadrats Density ¼ were produced. Abbreviated scientific names of plant Total number of quadrats studied taxa used is documented in S2 Table. In order to select an optimum dimension for NMDS analyses for both Total number of quadrats in which species occur herbs and shrubs, stress-dimension analyses were car- Frequencyð%Þ ¼ Total number of quadrats studied ried out and dimension 2 was used for both NMDS � 100 analyses of shrubs and herbs using “Bray-Curtis” dis- tance. urther, influence of selected climatic parameters Total number of individuals of aspecies in all quadrats (temperature, precipitation, solar radiation, wind Abundance ¼ Total number of quadrats in which species occur speed, isothermality) on species richness, diversity and density was evaluated using Pearson’s correlation IVI = Relative Density + Relative Frequency + coefficient method to check significant relationships Relative Abundance GEOLOGY, ECOLOGY, AND LANDSCAPES 5 among selected parameters. Further, Canonical was 1.1 for trees, 2.0 for shrubs and 1.7 for herbs, Correspondence Analysis (CCA) was carried out to while, species to family (S/F) values were 1.8 for assess relationships between environmental para- trees, 3.1 for shrubs and 5.0 for herbs. Furthermore, meters and species composition in study transects. across the altitude zones, S/G and S/F values declined The community data matrices of plant species for all monotonically from lower to higher zones for trees transects were used with the constrained matrices and shrubs. However (S/F) ratio in 3501–4000 m zone composing of environmental parameters, (i) altitude, was higher (3.9) as compared to that in 3000–3500 m (ii) temperature, (iii) precipitation, (iv) solar radiation, (3.6) for herbs Table 1. (v) isothermality, (vi) wind speed. The nature of rela- At quadrate level, species richness was maximum in tionships between soil parameters and vegetation dis- Johar, for all growth forms, ranging, while Shannon tribution was depicted through ordination diagram. diversity Index (H’) was highest in Niti-Malari for The CCA analyses were carried out using the R-Vegan tress (1–1.05), Johar for shrubs (0.45–0.91) and package (version 2.5–3) function “CCA” and results Byans 1 for herbs (0.91–1.33) (S3 Figure). Broad were analysed with respect to inertia, ranks and eigen range of compositional attributes of vegetation in values of constrained matrices and parameters. study transects depicted higher densities of trees in Byans 1 &2, i.e., 20–240 ind/ha, while shrub and herb densities were higher in Darma (1200–3000ind/ha) and Byans 1 (31000–115000ind/ha), respectively (S4 Results Figure). Beta diversity values calculated for each life Floristic diversity pool form along altitude gradient exhibited an increasing trend in all transects, thus indicating scattered and A total of 265 vascular plants belonging to 155 genera heterogeneous plant composition especially at high and 55 families were observed along the investigated altitudes. Apart from this, beta diversity contributed altitude range (i.e., 3200–4800 m) of Uttarakhand significantly to transect level species richness of Byans, alpines (S2Table). Among these, nine were gymnos- Johar, Mana and Nelang for herbs and shrubs, but very perms distributed in six genera and three families, little for trees. However, in Darma tree species change while the rest belong to angiosperms. Asteraceae was rapidly while herbs and shrubs change slowly and in the most dominant family having 30 plant taxa fol- Niti-Malari, shrub species exhibited rapid change lowed by Rosaceae (21) and Ranunculaceae (18). In Table 2. terms of growth forms, herbaceous habit (81.7%) was dominant, followed by shrubs (14.9%) and trees (3.4%). In relation to 500 m altitude zones, plant Altitudinal relationship of compositional features population was maximumin the lowest zone, i.e., 3000–3500 m (72% of total flora), followed by 3501– Considering the alpine treeline area broadly varies 4000 m (62%), 4001–4500 m (38%) and 4501–4800 m from 3000–4000 m in the west Himalayas, treeline in (9%) Table 1, thus exhibiting a continuous decline in present study was identified at 3200–3900 m with 1–3 the number of plant species with increasing altitude. tree species per plot, 3–11 shrubs per plot and 9–30 There was a more rapid decline in distribution of herbs per plot). Patterns of species richness and diver- higher level of taxa (genera and family) with altitude sity of different growth forms across altitude range- nd as compared to that of species Table 1. The species to were modelled and a 2 order polynomial regression genera (S/G) value calculated for the entire landscape showed a significant fit (p < 0.05, Figure 2), with both Table 1. Floristic diversity pool (S- Species, G- Genus, F- Family) in high-altitude alpine region of Uttarakhand along altitude gradient. Trees Shrubs Herbs Altitudes (m) S G F S/G S/F S G F S/G S/F S G F S/G S/F 3000–3500 9 8 5 1.1 1.8 38 20 13 1.9 2.9 146 97 41 1.5 3.6 3501–4000 4 4 4 1.0 1.0 24 16 11 1.5 2.2 137 89 35 1.5 3.9 4001–4500 - - - - - 10 9 7 1.1 1.4 93 66 31 1.4 3.0 4501–5000 - - - - - 2 2 2 1.0 1.0 22 19 12 1.2 1.8 Total 9 8 5 1.1 1.8 40 20 12 2.0 3.1 219 127 44 1.7 5.0 Note: S- Species; G-Genera; F- Family. Table 2. Beta diversity variations across life forms and transects. Beta diversity in different transects Life forms Byans 1 Byans 2 Darma Johar Mana Nelang Niti-Malari Trees 1.86 1.67 2.54 1.53 1.89 2.33 1.67 Shrubs 3.82 5.45 1.52 2.91 6.54 4.26 3.89 Herbs 4.33 4.14 1.38 3.55 5.17 4.36 1.94 6 K. C. SEKAR ET AL. Figure 2. Species richness, diversity and density trends along altitude gradients in different life forms [A] Trees, [B] Shrubs, [C] Herbs in alpine of Uttarakhand, west Himalaya. richness and diversity monotonically decreasing with Betula utilis, Juniperus semigloboosa and Salix denti- increasing altitude. While the decrease in tree diversity culata were present in 3600–3900 m zone. Among was steep at altitude zone 3600–3800 m with no spe- shrubs, the species of Astragalus, Berberis, cies observed above 4000 m, it was moderate for herbs Cotoneaster and Salix were dominant in lower altitude and shrubs upto the upper hard boundaries. zones, while Cassiope fastigiata, Ephedra gerardiana, Furthermore, density distribution of different growth E. intermedia, Juniperus indica and Rhododendron formsalong altitude gradientfollowed a similar anthopogon are mostly dominated in high-altitudes decreasing pattern. For different size classes (i.e., (Table 3). Moreover, types of dominant herbaceous trees, saplings, seedlings), density decreased at higher plants highly varied across altitude gradient. At lower altitudes drastically, ranging from 10–540 ind/h for altitude zones several common species such as trees, 10–80 ind/h for saplings and 100–1000 ind/h Arenaria bryophylla, Bistorta affinis, Danthonia for seedlings (Figure 2) (S2 Figure). As for shrub and cachemyriana, Geranium wallichianum, Oxyria herb densities in relation to altitude, a relatively mod- digyna, Oxytropis lapponica, Potentilla spp., erate but significant decline (p < 0.05) was seen Taraxacum officinale and Thymus linearis were pre- (Figure 2). Based on the Important Value Index sent in fairly high proportions, whereas higher alti- (IVI) of plant taxa in each altitude, dominant and tudes exhibited clear dominance of Bistorta vivipara, codominant species across three growth forms are Carex nivalis, Gentiana stipitata, Geranium wallichia- listed in Table 3. Cluster dendrogram for the study num, Juncus concinnus and Meconopsis aculeata. area identified four major altitudinal zones based on species composition, i.e., 3200 to 3500 m, 3600 to Vegetation composition across different transects 3900 m, 4000 to 4500 m and 4600 to 4800 m (Figure 3). Lowest altitude zone (3200–3500 m) inhab- NMDS of vegetation abundance and combined dataset ited tree species such as Pinus wallichiana, Abies pin- of studied transects further exhibited a strong gradient drow, Prunus cornuta and Cedrus deodara, while separating sample (plots) and species between GEOLOGY, ECOLOGY, AND LANDSCAPES 7 Table 3. Dominant and co-dominant taxa along altitude gradient in studied alpine regions of Uttarakhand, west Himalaya. Dominant species and co-dominant taxa (IVI) Elevation Trees Shrubs Herbs 3200 Pinus wallichiana (151.0), Berberis jaeschkeana (31.1), Cassiope fastigiata (24.2), Bistorta affinis (12.9), Geranium wallichianum (10.8), Abies pindrow (57.5) Juniperus communis (23.2) Danthonia cachemyriana (9.8) 3300 Pinus wallichiana (110.6), Casragana versicolor (32.4), Berberis jaeschkeana (27.4), Geranium wallichianum (11.4), Taraxacum officinale Abies pindrow (96.3) Cotoneaster microphyllus (21.7) (9.8), Bistorta affinis (9.3) 3400 Pinus wallichiana (159.5), Juniperus communis (33.0), Berberis jaeschkeana (28.5), Thymus linearis (15.5), Poa alpina (15.3), Danthonia Betula utilis (81.4) Cotoneaster microphyllus (28.5) cachemyriana (11.1) 3500 Pinus wallichiana (176.0), Caragana versicolor (37.0), Juniperus communis (34.6), Geranium wallichianum (14.1), Danthonia Betula utilis (70.1) Berberis umbellate (25.5), Juniperus indica (25.5) cachemyriana (11.0), Poa alpina (10.6) 3600 Pinus wallichiana (143.2), Juniperus indica (53.0), Cotoneaster microphyllus (29.7), Potentilla argyrophylla var. atrosanguinea (16.9), Poa Betula utilis (112.5) Caragana versicolor (26.8) alpina (11.7), Thymus linearis (10.2) 3700 Betula utilis (146.9), Salix Caragana versicolor (29.1), Juniperus indica (26.0), Potentilla argyrophylla var. atrosanguinea (18.0), denticulata (73.4) Astragalus oplites (25.4), Salix flagellaris (25.4) Taraxacum officinale (12.7), Thymus linearis (12.5) 3800 Betula utilis (225.5), Myrtama elegans (40.0), Hyssopus officinalis (3.5), Poa alpina (21.4), Thymus linearis (17.3), Oxyria Juniperus semiglobosa Astragalus oplites (28.7), Caragana versicolor (28.7) digyna (15.3) (74.5) 3900 Betula utilis (300.0) Juniperus indica (55.0), Cassiope fastigiata (39.0), Danthonia cachemyriana (15.7), Bromus japonicus Berberis jaeschkeana (34.4) (14.9), Potentilla argyrophylla (14.6) 4000 Cassiope fastigiata (53.7), Berberis jaeschkeana (34.0), Poa alpina (21.5), Oxytropis lapponica (16.8), Rhododendron anthopogon (34.0) Potentilla argyrophylla var. atrosanguinea (16.2) 4100 Cassiope fastigiata (64.9), Juniperus indica (56.7), Arenaria bryophylla (15.3), Euphrasia himalayica Caragana versicolor (39.5) (14.7), Poa alpina (14.5) 4200 Cassiope fastigiata (84.5), Rhododendron anthopogon Bistorta affinis (18.7), Bergenia stracheyi(16.4), Carex (54.8), Juniperus communis (52.1) nubigeana (14.2), Saxifraga flagellaris (14.2) 4300 Juniperus indica (117.9), Caragana versicolor (72.6), Potentilla argyrophylla (23.3), Bistorta affinis (21.9), Ephedra intermedia (72.6) Poa alpina (18.1) 4400 Juniperus indica (181.7), Ephedra gerardiana (118.3) Poa alpina (32), Geramiun wallichianum (31), Aconogonum tortusom (18) 4500 Juniperus indica (170.0), Ephedra gerardiana (130.0) Geranium wallichianum (44.6), Poa alpina (35.1), Potentilla argyrophylla var. atrosanguinea (27.9) 4600 Juniperus indica (300) Geranium wallichianum (95.2), Bistorta vivipara (24.9), Meconopsis aculeata (24.9), Poa alpina (24.9) 4700 Ephedra intermedia (300) Juncus concinnus (96.2), Carex nivalis (26.9), Gentiana stipitata (26.9), Poa alpina (26.9) 4800 Geranium wallichianum (100.0), Allardia glabra (27.8), Bromus inermis (27.8) transects as well as altitude range along the first two and higher altitude plots (4400–4800 m) in Mana axis (NMDS1 and NMDS2) (Figure 4). Separation formed distinct cluster with Darma transect showing along the first axis explained 40% of variation proximity in their species composition inhabiting while second axis explained 20%, thus underscored Cotoneaster spp., Convolvulus arvensis, Polygonum the affinity between different altitudinal vegetation polystachyum, Rosa macrophylla. Mid-altitude zones zones. Across all transects, distinct clusters denoting in Mana, i.e., 3600–3800 m, showed similarity with vegetations zones were identified characterized by spe- Nelang, while 3900–4300 m zone was scattered along cific species. Nelang transect formed a cluster towards positive end of first axisalong with species like Galium the negative end of first axis, with plotsat higher alti- asperuloides Nardostachys jatamansi, Picrorhiza kur- tude (3900–4400 m) in more proximity with each roa, Poa compressa and Saussurea simpsoniana. other inhabiting speciessuch as Aconogonum tortuo- Overall, sample plots in lower altitudes of transects sum, Melica persica, Poa tibetica, Sisymbrium brassici- were more clustered towards the middle of both axes, forme, Urtica hyperborea. Lower and mid-altitude inhabiting common plant species. Most of the sample plots of Byans 1 (3200–3700 m) and Byans 2 (3200– plots in higher altitudes occupied extreme ends (posi- 3800 m) as well as NitiMalari transect plots formed tive and negative) of both axes, thus showing increase a separate cluster towards negative end of second axis, in rarity in species composition moving from low to inhabited by similar species (such as Carex nivalis, high altitudes. Delphinium brunonianum, Primula denticulata, Rheum webbianum, U. dioca). On contrary, higher Relationship of vegetation composition with altitude plots in Byans 1 (>3800 m) & Byans 2 environmental variables (>3900 m) formed a distinct cluster towards positive end of first axis characterised by species such as Correlation analysis among climate variables and Christolea himalayensis, Leontopodium brachyactis, plant diversity indices is depicted in Figure 5 along Ranunculus palmatifidus, Saxifraga flagellaris. For with Pearson’s correlation coefficient and significance Johar, study plots were scattered across the four quad- level of relation. As established earlier also, altitude rants, thus exhibited large variations in species com- has a significant effect on density, diversity, and rich- position along altitude gradient. Lower (3200–3500 m) ness of plant species (p < 0.001), with an apparent 8 K. C. SEKAR ET AL. Figure 3. Hierarchical cluster dendrogram showing distinct altitudinal zones with respect to the species composition in alpine region of Uttarakhand, west Himalaya. decrease in density (r= -0.77), diversity (r = −0.87), Byans 1 & 2 due to the high wind speed, low tempera- and richness (r = −0.70). Similarly, there was strong ture and precipitation species like Christolea hima- negative correlation (p < 0.001) of temperature (r = layensis, Gentiana stipitata, Leontopodium brachyatis, −0.96), precipitation (r = −0.77), and solar radiation Rhodiola bupleuroides, Saussurea gossypiphora, (r = −0.71) with altitude as well, while it was positively S. simpsoniana and Sibbaldia parviflora. Contrary to significant with wind speed (r = 0.83). Apart from this, species like Anaphalis nepalensis, A. triplinivalis, altitude, temperature and precipitation also made Berberis umbellata, Carum carvi, Cotoneaster rotundi- a strong gradient influencing plant species richness folius and Ribes orientale favoured lower altitude and diversity in alpine regions Figure 5. Increase in transects like Darma and Johar with high temperature temperature as well as precipitation favoured com- and precipitation. Occupying positive end of first axis paratively more number as well as variety of species and negative end of second axis, Nelang transect in study area, which is evident with significantly posi- exhibited very little dependence on temperature, pre- tive correlation coefficient values (p <0.01). As indi- cipitation and wind speed. In context of plant species, cated by r values, solar radiation and wind speed the site thus occupies herbs like Aconogonum tortu- exhibited significant (p < 0.05) but weak correlation sum, Rumex nepalensis, Urtica hyperborea and with richness and diversity indices, thus exhibiting Verbascum thapsus. Furthermore, isothermality of little influence on presence and types of plant taxa in study site favoured growth and abundance of species alpine. like Aconitum balfourii, Gentiana argentea, The relative effect of climatic parameters on plant G. leucomelaena and Pedicularis tubiformis. species composition, analysed by CCA, showed stron- gest influence of precipitation in forming altitudinal Discussion zones having specific vegetation composition with high axes correlation (Figure 6), followed by tempera- The results of our study are based on data gathered ture, isothermality, and wind speed, while influence of during a time period from 2014 to 2018 of field work. solar radiation was lowest. Total inertia of combined A baseline floristic survey of study area (field and data set was 13.02 where constrained climatic para- literature based) documented a total of 932 plant meters explained 48% of total variances on first axis taxa belonging to 371 genera in 76 families Chandra and 30% on second axis. Among altitude transects, in Sekar et al., 2022. Hence, in comparison with previous GEOLOGY, ECOLOGY, AND LANDSCAPES 9 Figure 4. Non-metric Multi-Dimensional Scaling (NMDS) representing species composition among sample plots along altitude gradients of study transect. 10 K. C. SEKAR ET AL. Figure 5. Pearson’s Correlation between plant richness, density and diversity indices with climate variables. studies (Negi, 2002), the high-altitude alpine zones their abundance in different communitieswith (3200–4800 m) of Uttarakhand accounts for 58% extreme conditions (Abbas et al., 2019). alpine flora of west Himalayas. While comparing The patterns of S/G along altitude gradient are used with pan-Himalayas (Rana & Rawat, 2017), the study to describe speciation and diversification rates (Floeter area inhabits 10% species, 16% genera, and 32% et al., 2004). The general altitudinal decrease in S/G as families. Furthermore, 47 plants were under different well as S/F recorded in our study indicates their phy- threat categories as per IUCN and CAMP (Ved et al., logenetic dispersion towards higher altitudes. On 2003). Among these, Dactylorhiza hatagirea, comparing the overall S/G values of different life Nardostachys jatamansi and Picrorhiza kurroa were forms, the higher ratio for shrubs indicates more “Critically Endangered,” while Aconitum heterophyl- intense diversification within the genera as compared lum, Angelica glauca, Ephedra gerardiana, Fritillaria to trees and herbs. Similarly, within family herbs exhi- roylei, Saussurea obvallata and Podophyllum hexan- bit significantly more intense phylogenetic clumping drum were “Endangered.” Hence, any effort made as compared to others. The increasing values of beta towards conservation ofthis high-altitude zone would diversity calculated along altitude gradient for each eventually benefit a significant proportion of transect indicates heterogenous plant composition, Himalayan biodiversity including flora as well as thus exhibiting more scope for occurrence of rare fauna. Dominance of herbaceous growth form in taxa as compared to lower altitudes (Rawal et al., terms of richness, diversity as well as density is appar- 2018). This can be attributed to the variable levels of ent due to extreme climatic conditions in the region environmental as well as anthropogenic pressure pre- which causes poor assemblages of trees and shrubs vailing in the different study areas. Relatively high (generally high stature species). Apart from this, dom- level of disturbance and poor species pool leads to inance of families such as Asteraceae and Rosaceae can lower beta diversity in herbs, while it exhibits be attributed to their specialized morphology, and a reverse trend for trees. Among the studied transects, broad ecological amplitude enabling them to establish Darma with low β diversity exhibits high species GEOLOGY, ECOLOGY, AND LANDSCAPES 11 Figure 6. Canonical Correspondence Analysis of combined data set: ordination of sample pots and vegetation abundance constrained by their relationships to environmental variables. Significant parameters are shown as biplot vector. packaging and homogenous composition in case of season paving way for plant species tolerant to herbs, while becomes highly heterogeneous for trees extreme conditions, a phenomenon known as “ecolo- (high β diversity). Similarly, with high beta diversity of gical filtering.” These fluctuations also influence soil herbs in Mana transect there is more scope for occur- texture, nutrients and stability of substrate which in rence of rare/occasional herbaceous species, which turn are responsible for maintaining vegetation com- contributes for higher heterogeneity of species position (Sanchez-Gonzalez & Lopez-Mata, 2005). composition. Hence, forming distinct vegetation zones along alti- Several studies have demonstrated characteristic tude, with tree species occupying the lowest zone relationships between altitude and patterns of species along with woody anddeep-rooted shrubs like richness, diversity, and compositional features (such Caragana spp., Juniperus spp. and Lonicera spp., fol- as density, total basal area) along a gradient (Acharya lowed by meadows at higher elevations dominated by et al., 2011; Rawat et al., 2010; Sharma & Raina, 2013). herbaceous growth form including forbs, sedges, tus- In present study also, density and altitude exhibited sock and non-tussock grasses. highly significant relationship (p < 0.05, F>F ) in Ordination analysis (NMDS) clearly exhibited var- crit entire study zone. This signifies the strong dependence iation in vegetation composition among transects of floristic diversity and density with altitude, irrespec- (Figure 4). While temperature was the major factor tive of the growth stage or life forms in the alpine responsible for variation along altitude, among trans- region. Our results are congruent with several former ects precipitation had the strongest influence on vege- studies (Kala & Mathur, 2002; Pandey et al., 2018; tation composition, followed by temperature, wind Rawal et al., 2018; Sharma et al., 2014; Sinha et al., speed and isothermality as depicted by CCA analysis. 2018; Vetaas & Grytnes, 2002; Wang et al., 2006). Byans and Niti-Malari regions with cold, dry climate Reduction of plants in higher altitude may be attrib- and high wind speed had similar species composition. uted to eco-physiological constraints, such as extre- Nelang region formed a separate cluster due to its mely low temperature, precipitation and solar variable climate such as comparatively low tempera- radiation (Figure 5) which results in short growing ture, precipitation, isothermality and wind speed. 12 K. C. SEKAR ET AL. Darma and Johar fell in the same cluster with com- documents the plant species diversity and composition paratively warm and wet climate experiencing slower of remote areas of Himalaya which are known to be winds and optimum isothermality. natural laboratories to study the responses of plants to While comparing to earlier literature (Chandra climate change. While our study is more focussed on Sekar et al., 2015; Garbyal et al., 2007; Lohani et al., plant diversity, it can be expanded to broader aspects 2013; Mitra et al., 2017; Rawat, 2005; Uniyal et al., such as phenological and physiological studies of alpine 2002), the density of Aconitum heterophyllum, plants and their relation with climatic and edaphic Allium stracheyi, Bergenias tracheyi, Carum carvi, factors (Majeed et al., 2022) and opt more towards Hippophae salicifolia, Rhododendron anthopogon, R. plant distribution models (Bayat et al., 2021; Jahani & campanulatum, Swertia ciliata and Thymus linearis Saffariha, 2020; Jahani et al., 2021; Mosaffaei et al., was lesser, while that of Juniperus communis, 2020). J. indica, Polygonatum verticillatum density was increased. This can be attributed to the irregular har- Conclusion vesting of former species for fuel and medicinal pur- poses. The population of Betula utilis highly decreased This study is the first attempt to document and analyse in the area due to its utilization for fuelwood in higher floristic composition of high-altitude alpine regions Himalayan regions (Mitra et al., 2017). Climatic bar- panning across Uttarakhand, west Himalaya, along riers, limiting tree habits, resulting limited density altitude gradient and determine major climate drivers coupled with high extraction as fuelwood may be the influencing vegetation patterns. Hierarchical clustering reason of population reduction. Furthermore, trees as well as ordination analysis (NMDS and CCA) per- and saplings of B. utilis are available, but there are formed also showed concordant results, underscoring no seedlings recorded in 3500 m. Thus, protection of altitudinal vegetation zonation throughout study area these saplings is need of the hour, so that the popula- as well as its relationship with environmental variables. tion of B. utilis can be saved. In view of importance of Phytosociological studies undertaking multiple trans- the tree, the local considered it as “sacred” and various ects and life forms in high altitude alpine regions are conservation activities were observed. The people of very scarce especially in Indian Himalayan region. In the local areas must aware about the importance of this context, the present study also attempts to discuss plants and some good initiatives (i.e., J. semiglobosa and compare distribution patterns of vegetation across conservation in Johar) should be replicated in all other the longitudinal gradients along multiple altitudes regions, to conserve the plant diversity. Furthermore, transect. In this context, the present study gives com- density and diversity are also influenced by frequency parable ecological data sets to discuss distribution pat- of anthropogenic disturbances, i.e., road construction, tern of seven altitudinal transects. Species richness and habitation, commercial plantation and agricultural diversity along the altitude gradient in alpine region performs also results in replacement of natural vegeta- exhibited a monotonic decrease, both overall and by tion with man-made ecosystem. growth forms, with maximum values in the lower Under current global climate change scenarios, altitude zones (3200–3500 m). Beta diversity also a number of studies have projected changes in vegeta- increased with altitude thus exhibiting higher hetero- tion distribution and upward shifts in plant boundaries geneity and presence of unique species adapted to (Kuhn et al., 2016; Rai et al., 2010; Rosset et al., 2010; survive the extreme environmental conditions. Telwala et al. 2013). These shifts tend to change the Among growth forms, beta diversity was generally shape of ecosystems of high-altitude regions. However, higher for herbs and shrubs as compared to tress indi- shifting rate varies with species depending on their cating homogeneity in tree populations at high-altitude altitudinal and geographical distribution ranges, and alpines. The observed variations in temperature and their accumulative potential (Klimes et al., 2003). As precipitation patterns along altitude gradient signifi- species distribution range decrease with increasing alti- cantly correlates with the species richness and diversity tude, species with narrow amplitude of altitudinal dis- patterns, with high temperature and precipitation sup- tribution and endemics are more affected by porting plant richness and diversity. The linear correla- environmental or anthropogenic disturbances. While tion states altitude as the variable which best explains climate change (natural or anthropogenic) threatens the variations in richness and diversity patterns. high-altitude flora, it also impacts a significant portion of useful biodiversity and traditional livelihood of local people. Hence it becomes pivotal to carry out empirical Acknowledgements studies pertaining to potential extent and magnitude of The authors thankfully acknowledge the facilities provided current and future shifts in altitudinal limits of plants by Director, G.B. Pant National Institute of Himalayan in response to climate change in order to draw con- Environment (NIHE), Kosi-Katarmal, Almora for undertak- clusive conservation and management strategies ing this work. We are gratefully acknowledging the financial (Mehta et al., 2020; Pandey et al., 2019). Our study assistance received from Council of Scientific and Industrial GEOLOGY, ECOLOGY, AND LANDSCAPES 13 Research [EMR No. 38 (1346)/12/EMR-II], Kailash Sacred Upadhyay, S., & Dey, D. (2022). Influence of anthropo- Landscape Conservation and Development Initiative and genic pressure on the plant species richness and diversity National Mission on Himalayan Studies (NMHS), Ministry along the elevation gradients of Indian Himalayan of Environment, Forest and Climate Change, Government of high-altitude protected areas. Frontiers in Ecology and India (GBPNI/NMHS-2018-19/SG 11) and National Mission Evolution, 10, 751989. https://doi.org/10.3389/fevo.2022. on Sustaining the Himalayan Ecosystem (NMSHE), TF-3, 751989 DST, New Delhi. Permissions for Herbarium and library Carpenter, C. (2005). The environmental control of plant consultation received from different organizations, i.e., species density on a Himalayan elevation gradient. Botanical Survey of India, Forest Research Institute, Journal of Biogeography, 32(6), 999–1018. https://doi. Wildlife Institute of India, National Botanical Research org/10.1111/j.1365-2699.2005.01249.x Institute; Permission of State Forest Department received Chandra Sekar, K., Lalit, G., Aseesh, P., & Srivastava, S. K. from Chief Wildlife Warden (Letter no. 917/5-6/DDN/ (2015). A note on distribution of juniperus semiglobosa in dated 16 Sep. 2013); logistic and field guidance received Uttarakhand, India. Indian Journal of Forestry, 38(1), from DG, ITBP (Letter no. 2766 dated 14.08.2014) are highly 79–80. https://doi.org/10.54207/bsmps1000-2015- acknowledged. Dr. Poonam Tripathi, International Centre 0XY6MS for Integrated Mountain Development (ICIMOD), Nepal is Chandra Sekar, K., Pandey, A., Giri, L., Joshi, B. C., Bhatt, gratefully acknowledged for preparing study area map. The D., Bhojak, P., Dey, D., Thapliyal, N., Bisht, K., Bisht, M., inputs received from anonymous reviewers aided in improv- & Negi, V. S. (2022). Floristic diversity in Cold Desert ing contents of this manuscript, we highly thank them. regions of Uttarakhand Himalaya, India. Phytotaxa, 537 (1), 1–62. Chandra Sekar, K., & Srivastava, S. K. (2009). Flora of the Disclosure statement Pin Valley National Park, Himachal Pradesh. Botanical Survey of India. Ministry of Environment and Forests. No potential conflict of interest was reported by the authors. Cui, Y., Bing, H., Fang, L., Wu, Y., Yu, J., Shen, G., Jiang, M., Wang, X., & Zhang, X. (2019). Diversity patterns of the rhizosphere and bulk soil microbial communities along ORCID an altitudinal gradient in an alpine ecosystem of the east- ern Tibetan Plateau. Geoderma, 338, 118–127. https://doi. K. 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Journal

Geology Ecology and Landscapes – Taylor & Francis

Published: Jan 14, 2023

Keywords: Alpine; species richness; diversity; Uttarakhand; Himalaya; India

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