Environmental Research 111 (2011) 967977

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Environmental Research
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Altitudinal distribution patterns of the native and alien woody ora in Kashmir Himalaya, India
Anzar A. Khuroo a, Ewald Weber b,n, A.H. Malik a, Zafar A. Reshi c, G.H. Dar a
a

Centre for Biodiversity and Taxonomy, Department of Botany, University of Kashmir, Srinagar-06, Jammu and Kashmir, India Institute of Biochemistry and Biology, Biodiversity Research, University of Potsdam, Maulbeerallee 1, 14469 Potsdam, Germany c Biological Invasions Laboratory, Department of Botany, University of Kashmir, Srinagar-06, Jammu and Kashmir, India
b

a r t i c l e i n f o
Available online 23 July 2011 Keywords: Alien species Elevation Floristics Plant invasions Species richness

abstract
Background: Many studies have shown that alien species richness pattern follows that of native species richness patterns along environmental gradients, without taking the specic composition of the two groups into account. Objectives: To compare species richness patterns of native and alien woody plants along an altitudinal gradient in Kashmir Himalaya, India, and to analyse the specic composition, e.g. proportion of life forms. Methods: Analysis of secondary data from published oristic inventories. The gradient (5004800 m asl) was split into 100 m bands and presence/absence data for each species were obtained, for each band. Results: Species richness of both native and alien species followed a hump-shaped distribution. Alien species richness dropped faster above 2000 m asl than the native did. The ratio of trees to shrubs decreased monotonically along the gradient in native species, but showed a peak at c. 2500 m asl in alien species. Alien species owered in average earlier than native species. Conclusions: The change of species richness of native and alien species along altitude is similar, but the proportion of life forms is not. Most likely both climatic and socio-economic factors affect alien species richness and its specic composition in the Kashmir Himalaya. & 2011 Published by Elsevier Inc.

1. Introduction Plant invasions in mountainous regions have recently attracted attention (Dietz, 2005; Pauchard et al., 2009; Khuroo et al., 2009; Jakobs et al., 2010). There are many reasons which make plant invasions into high-altitude mountain regions an issue of global concern. Mountains are generally biologically diverse, support a high proportion of endemic species and harbour most of the worlds protected areas (Kollmair et al., 2005). However, in recent times mountain areas are experiencing increased levels of humanmediated disturbances such as urbanisation, deforestation, grazing, and other impacts that result in disturbed habitats and affect s-Bravo et al., those forces governing biodiversity gradients (Nogue 2008). These factors, in conjunction with increasing population density and transport networks at higher altitudes facilitate the introduction and invasion of alien species (Price, 2006). Invasive alien species already present in a mountain region, but yet

Corresponding author. E-mail addresses: anzarak@gmail.com (A.A. Khuroo), ewweber@uni-potsdam.de (E. Weber), ecoakhtar@gmail.com (A.H. Malik), zreshi@yahoo.com (Z.A. Reshi), profdar99@gmail.com (G.H. Dar). 0013-9351/$ - see front matter & 2011 Published by Elsevier Inc. doi:10.1016/j.envres.2011.05.006

conned to lower altitudes, could potentially expand their altitudinal ranges in future as a result of climate change and adaptation to higher altitudes (Walther, 2007), but also simply due to reaching new sites by dispersal (Becker et al., 2005). As the number of alien species increases at higher altitudes, threats and impacts on native biodiversity, including endemic species might also increase. Plant invasions at higher altitudes thus have direct implications for biodiversity conservation and habitat management in mountain ecosystems. The most apparent pattern along altitudinal gradients is the decrease in species numbers with increasing altitude. A pertinent question for understanding plant invasions in mountain areas relates to the change of species richness along that gradient, e.g. how the change in alien species compares to that of native species. If the underlying mechanisms differ between aliens and natives, one might expect that the species richnessaltitude relationship also differs between these groups. The question of alien vs. native species richness patterns along environmental gradients has mainly been addressed by studying latitudinal or productivity gradients (Lonsdale, 1999; Levine and DAntonio, 1999). These studies found that at broad geographic scales, a positive relationship is more frequent (e.g. Stohlgren et al., 1999), and that at smaller scales, positive or negative relationships are apparent (Shea and Chesson, 2002).

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In the case of altitudinal gradients, biogeographers have demonstrated characteristic relationships between native species richness and altitude. These vary from hump-shaped relationships with maximum species richness at mid-altitudes (e.g. Rahbek, 1995; Vetaas and Grytnes, 2002; Carpenter, 2005) to monotonically decreasing relationships (e.g. Korner, 2004; Mallen-Cooper and Pickering, 2008). The shape of these relationships depends largely on the scale and whether parts or the whole gradient has been taken s-Bravo et al., 2008). Generally, a number of into account (Nogue environmental factors change with increasing altitude, some of which are directly determined by altitude, e.g. average temperature, whilst others are not directly related to altitude, e.g. precipitation (Korner, 2007), thus making species richnessaltitude relationships difcult to interpret. valo et al., 2005; Daehler, 2005; McDougall Recent studies (Are et al., 2005; Khuroo et al., 2007; Tassin et al., 2007) have shown that mountain regions also harbour many naturalised alien plant species and that these species can ascend to high altitudes. Some workers have addressed the alien-native species richness question along altitudinal gradients and found a linear decrease in both alien and native species richness along altitude (Pauchard and Alaback, 2004; Becker et al., 2005; Bear et al., 2006; Mallen-Cooper and Pickering, 2008). However, these few studies did consider a rather short altitudinal gradient. A hump-shaped distribution of alien plants along altitude was found in the Tenerife, Canary Islands (Haider et al., 2010) and Hawaii (Jakobs et al., 2010). Peaks of alien species richness were also evident in ve out of eight data sets analysed by Alexander et al. (2011) and covering several mountain ranges of the world. These analyses focused on species numbers but did not investigate how the quality of the species pool, e.g. the relative proportion of life forms, changes with altitude in native and alien species, respectively. In addition, there are yet few studies comparing both alien and native species richness patterns along altitude. Since altitudinal gradients exhibit strong environmental changes over short distances, plant occurrence is less constrained by dispersal than by environmental conditions. Such gradients are suitable for asking how native and alien species richness, and their specic composition, respond to these changes. Here, we analysed altitudinal distributions of the woody ora of Kashmir Himalaya, India. The region harbours a rich ora, including endemics, and offers a large altitudinal gradient (5004800 m asl). We compared native and alien woody species with regard to their altitudinal distribution patterns, life forms and time of owering. Specically, we asked the following questions: (1) Does species richness of woody taxa change in different ways along altitude in native and alien species, respectively? (2) Are there differences in the proportions of life forms and in owering time between alien and native species, and are these affected by altitude?, and (3) Can differences between native and alien species be attributed to human impacts such as land use patterns and population density?

Table 1 Main vegetation zones of Kashmir Himalaya. From Dar et al. (2002). Altitudes (m) 5001200 12001800 18002800 28003400 34004600 Zone Subtropical Temperate Montane Subalpine Alpine Characteristic vegetation Semi-evergreen monsoon forests Mixed forests with deciduous broad leaved and coniferous trees Coniferous forest Scrub vegetation Alpine grasslands

Greater Himalayan mountain ranges prevent rain clouds from reaching into the interior valleys, Ladakh receives just about 100 mm rain per year. Human pressure is intense in the region, urbanisation and timber extraction currently being the most serious threats to natural habitats (Oza, 2003). Population density in Jammu and Kashmir provinces is high at low altitudes, exceeding 500 persons/km2 in some districts (Anonymous,1995, 2009). 2.2. Data sources We screened the available oristic literature to compile a list of native and alien woody plant species growing in Kashmir Himalaya. Although the rst written oristic records date back to 1822 (Khuroo et al., 2010), a comprehensive and recent treatment of the ora of the region is not yet available (Dar et al., 2002). Therefore, our primary sources included both recent and older local publications (see Appendix A). The species list obtained was authenticated by consultating about 5.000 herbarium specimens collected over the last 50 years, including those by the authors of the present study, which are deposited in the Kashmir University Herbarium (KASH). We then complemented the nal species list of the woody ora by incorporating eld observations and records from oristic surveys undertaken over the last decades by the authors of the present study. Species status (alien and native, respectively) was taken from the original sources, if available, and checked with the global distribution as inferred from the Germplasm Resources Information Network (www.ars-grin.gov). Of the 511 woody species identied, 311 were native and 200 alien species (see Appendix B and Khuroo et al., 2010). Data on altitudinal distribution (minimum and maximum elevation) and owering time (months in which a species is in ower) were obtained from the primary sources as outlined above. The data were complemented with altitude records on the herbarium specimens, and by eld notes from the surveys by the present authors. If different sources gave different altitudes for a species, we assumed that the species is found within that range of altitudes. 2.3. Data analyses For each altitudinal band of 100 m, species presence or absence was recorded. This data set served as the basis for altitude related analyses. Some additional variates were computed, such as altitudinal range (difference between maximum and minimum altitude), altitudinal midpoint (average of maximum and minimum altitude), length of owering (difference between month of owering end and month of owering start), and owering peak (average of month of owering end and month of owering start). Species were allocated to various categories describing the species functional types: life form (tree, shrub, subshrub, climber) and leang phenology (deciduous, evergreen). To obtain a general picture how the life form distribution changes with increasing altitude, life form distributions were plotted for each vegetation zone and compared. In order to relate the results to variation in socio-economic factors and land use along altitude, indirect measures of population density, percent forest cover, and percent agriculture cover were used. The data for these measures were retrieved from reports published by local governmental agencies (Anonymous, 1995, 2009). These data were available at the district level only (14 districts for the study area) and it was not possible to establish a link between these and our species richness data. We used the population and land use data to qualitatively describe human impacts along altitude. Data were analysed with the SPSS statistical program version 18.0 (SPSS Inc.). Median-tests and t-tests were used to test for differences in continuous traits between alien and native species.

2. Methods 2.1. Study area The Kashmir Himalaya is situated in the northwestern part of the Himalayas and lies between 321170 and 371200 North, and between 731250 and 801300 East. The region covers an area of c. 222 235 km2 and is divided into three provinces: Jammu, Kashmir and Ladakh. This Himalayan region is endowed with a high species richness (Dar et al., 2002). The three provinces differ in the plant cover and climate (Rodgers and Panwar, 1988; Singh et al., 1998), resulting in characteristic vegetation zones (Table 1). Whereas Jammu has a subtropicaltemperate climate, that of Kashmir is temperate, and Ladakh is a cold-arid region. Jammu province receives an annual precipitation of up to 1700 mm due to the summer monsoon rains; and Kashmir has an annual precipitation of about 1050 mm, mostly as winter snow. Because the

3. Results 3.1. Patterns of species richness Species richness followed an unimodal distribution in relation to altitude and the highest species richness occurred between

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Fig. 1. Species richness of native and alien woody plants along altitude within Kashmir Himalaya, India. (a) Species richness, expressed as number of species per 100 m altitudinal band and (b) proportion of alien species.

Fig. 2. Lifeform distribution of alien (a) and native (b) woody plants at each of ve vegetation zones in Kashmir Himalaya, India. sub subtropical monsoon forest zone, tem temperate mixed forest zone, mon montane coniferous forest zone, sal subalpine scrub zone, and alp alpine grassland zone.

1000 and 2000 m asl (Fig. 1). This pattern was evident in both native and alien species, although the shape of the curve differed between alien and native species. Whereas alien species richness rapidly decreased above 1600 m asl, native species richness decreased much slower (Fig. 1). As a result, the proportion of alien species was high up to c. 2000 m asl but decreased rapidly with further increase in altitude (Fig. 1).

3.3. Altitudinal distribution limits and ranges Alien and native species signicantly differed in their altitudinal limits and ranges (Table 2). Alien species did not reach such high altitudes as did native species. The maximum altitudes of alien species were 3300 m asl (Solanum pseudocapsicum), a shrub, while that of native species were 4800 m asl (Lonicera semenovii), and 4600 m asl (Lonicera spinosa), shrubs as well. Other alien species reaching 3000 m asl or more included the trees Juglans regia, Malus baccata, Morus alba, Populus alba, P. euphratica, P. nigra, Prunus armeniaca, Pyrus communis, Salix alba, S. babylonica, and S. viminalis. Differences between minimum and maximum altitude (altitudinal range) showed great variation among both native (100 2300 m asl) and alien (1002100 m asl) ora, alien species had in average a lower altitudinal range than native species (Table 2). Maximum altitude and altitudinal range signicantly correlated both in alien and native species (Fig. 4), the slope of aliens being higher than that of natives (aliens: s 0.615 7 0.042; natives: s 0.342 7 0.023). 3.4. Flowering phenology Flowering phenology, expressed as number of months during which a species remains in ower, varied greatly and ranged from

3.2. Life form distributions The life form distributions among vegetation zones differed between alien and native species (Fig. 2). In the temperate and montane zones, native species consisted of more shrub than tree species, the opposite trend was apparent in alien species. Climbers were mostly present in the subtropical and temperate zones, with no clear difference between alien and native species. In native species, the ratio of tree to shrub species decreased monotonically along the altitudinal gradient with an initial sharp drop (Fig. 3). Alien species tree to shrub ratios showed a distinct pattern with a peak at approximately 2500 m asl. Alien and native species also differed along the altitudinal gradient with regard to the ratio of evergreen to deciduous species (Fig. 3), with evergreen species dropping above c. 1500 m asl in alien species.

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Fig. 3. Trees to shrubs ratio (a) and evergreen to deciduous ratio (b) in relation to altitude in Kashmir Himalaya, India.

Fig. 4. Relationship between maximum altitude of a species and its altitudinal range (difference between maximum and minimum altitudes) for alien and native woody plants in Kashmir Himalaya, India. Lines represent linear regressions.

Table 2 Differences between native and alien woody plant species in Kashmir Himalaya with regard to biogeographic and ecological characters. Differences were evaluated with t-tests or median tests. Character Flower start Flower end Flowering period Flowering peak Lower altitude limit Upper altitude limit Altitudinal range Mid-altitude
a

of peak owering along vegetation zones (subtropical, temperate, montane, subalpine, alpine) were April, May, May, and April in the case of alien species (no species at the alpine zone), and May, May, June, June, and July in the case of native species, respectively.

Native species 5.23 6.60 2.0a 5.90 1584.0 2381.4 700a 1982.7

Alien species 4.72 6.02 2.0a 5.34 1352.4 1888.6 400a 1620.5

F-value (P) 11.56 ( o 0.001) 14.45 ( o 0.001) 0.895a 14.25 ( o 0.001) 23.37 ( o 0.001) 61.07 ( o 0.001) o 0.001a 47.50 ( o 0.001)

4. Discussion 4.1. Unimodal distribution of species richness Both native and alien species richness followed a hump-shaped or unimodal distribution pattern along the altitudinal gradient. Such unimodal relationships have often been found in various biota for the s-Bravo et al., gradients that span a large extent (Rahbek, 1995; Nogue 2008). Our results are strikingly similar to species richness patterns in ferns along an altitudinal gradient in Central Himalaya (Bhattarai et al., 2004), to woody plant species richness in the Western Himalaya (Oommen and Shanker, 2005), and to the vascular ora of Mt. Kinabalu in Borneo (Grytnes and Beaman, 2006). Native species richness along environmental gradients has developed over long periods of time and are inuenced by area, climate, productivity and topography (see Oommen and Shanker, 2005), and evolutionary factors, e.g. adaptation and speciation (Roy and Goldberg, 2007). Since the time of human settlement, native species richness is also affected by land-use and habitat alterations. In contrast to native species, diversity gradients of

Entries are medians and P refers to Chisquare values of median tests.

2 to 5 months in aliens and from 2 to 7 months in native species. Flowering characters differed signicantly between alien and native species but not the length of the owering period (Table 2). Numbers of owering woody species per month peaked from April to July along vegetation zones (Fig. 5). Peak owering month of aliens was generally earlier than that of native species, and that difference became increasingly apparent at higher vegetation zones (Fig. 5). The increasing difference was due to an increased later month of peak ower in the case of native species, and a fairly constant month of peak ower in alien species. The months

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Fig. 5. Phenological patterns in alien and native woody plants of Kashmir Himalaya, India. For each vegetation zone, percent of owering species among months are presented. Closed circles subtropical, open circles temperate, triangles montane, crosses subalpine, and quadrates alpine. Note that there are no alien species in the alpine zone.

species was much narrower than that of native species, with a particularly strong decline of alien species richness above 1700 m. Alien species had a lower median of altitudinal ranges than native species, thus the strong drop is due to lower altitudinal distribution limits in alien species. The largest difference between native and alien species was apparent with regard to their life form representation, especially the ratio of tree to shrub species. The monotonic decline in the case of native species is expected and reects climatic constraints on tree species, but alien species contrast by exhibiting a peak at c. 2500 m asl. This peak most likely reects the inuence of human activities. Although we do not have land use or population data at the level of altitudinal bands, indirect indications support this connotation. First, human population density is highest between 1000 and 2000 m asl in the study area (Gupta, 1978; Nandy et al., 2000). Second, taking percent agricultural area and percent forest cover at the district level shows a peak within that band (Fig. 6). Over the decades, extensive tree plantation programs have been carried out by the State government between 2000 and 3000 m asl. Robinia pseudoacacia, for example, has been introduced around 1890 in North West Himalaya between 1000 and 3000 m (Troup, 1912; Muthoo and Kango, 1965). As this altitudinal band corresponds to the montane vegetation zone, characterised by coniferous forests, natural conditions for tree growth are optimum. Alien shrub species, in contrast, are mainly conned to areas of human settlements and consist of species introduced as ornamentals. Above 3000 m, climatic conditions become limiting to tree growth, reducing the tree to shrub species ratio. The ratio of deciduous to evergreen species is rather obscure but again, native species differed markedly from alien species. The rather clear peak in the case of alien species could simply reect the choice of tree species for planting. That alien species respond in different ways to environmental variation than native species do, has been shown for the Alps (Marini et al., 2009). These authors have found that variation in alien plant species richness was largely explained by temperature and human population density. Native species richness was less strongly associated with these factors but was strongly associated with the presence of calcareous bedrock. In the present study, we did consider woody species and not herbaceous species. Trees and shrubs that became naturalised have mostly been intentionally introduced as ornamentals, erosion control or plantation trees (Cronk and Fuller, 1995). If all alien plant species would be considered, including accidentally introduced agricultural

alien species have emerged within a short period of time and are the result of human activities, e.g. introducing species and altering habitats. Discerning the factors leading to an observed species richness pattern along altitude is not easy as factors are often intercorrelated (Korner, 2007). Explanations for a mid-altitude peak of species richness are varied. Some authors argue that it may be caused by dispersal constraints at both ends of the altitudinal gradient, where midaltitudinal habitats have a higher diaspore input than areas close to the endpoints of the gradient (Grytnes, 2003). Mass effect, a zone of overlap between lowland and mountain species (Shmida and Wilson, 1985; Rahbek, 1995; Lomolino, 2001) and the extent s-Bravo et al., 2008) also affect patterns of human impacts (Nogue of species richness along altitudes. Furthermore, the fraction of land area generally decreases with increasing altitude (Korner, 2007), and any pattern of species richness along altitudinal gradients is inuenced by speciesarea relationships as well. 4.2. Differences between alien and native species Despite the overall similarity, the shape of the distribution curve differed between natives and aliens. The curve of alien
Fig. 6. Relationship between lowest altitude and percent agricultural area (black circles) and percent forest cover (white circles) for 14 districts in Kashmir Himalaya, India.

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weeds, the pattern might look different. The role of economic uses and land use patterns in explaining distribution patterns of alien plants in the study area merits further investigations. 4.3. Flowering phenology The alien woody ora differed from the native woody ora in most characters associated with owering phenology. The only exemption was length of owering period. On average, alien species started to ower earlier than native species, a difference that became increasingly apparent with increasing altitude. Flowering phenology is a conservative trait (Ausin et al., 2005), and that alien plants ower earlier than native plants has been demonstrated in many cases (Cadotte and Lovett-Doust, 2001; Lake and Leishman, 2004; Godoy et al., 2009). In the present study, owering phenology of alien species could reect the choice of suitable species for planting, and that alien species originate from areas of lower altitudes within their native ranges. 4.4. Altitudinal ranges Our results demonstrate that alien species can reach considerable altitudes in high mountain ranges, which has been shown for other mountain areas as well (e.g. Weber et al., 2008; Alexander et al., 2009). Alien species reaching more than 3000 m asl in Kashmir Himalaya were mostly of Asian origin. An exception was S. pseudocapsicum, whose native range encompasses the New World, Australia, and the Pacic. Mallen-Cooper and Pickering (2008) found that alien plants reached the same altitudes as native plants in the Australian Alps, although their gradient reached from 540 to 2020 m only. In the case of alien species, altitudinal ranges are related to the species ecology but also to introduction time. For example, the shrub Amorpha fruticosa has a low altitudinal range (100 m, the shrub is found from 1600 to 1700 m asl) because it represents a recent introduction and is conned to urban centres. Alexander et al. (2011) suggested that alien plants reaching high altitudes are species with broad climatic tolerances, exhibiting large altitudinal ranges, and beginning their spread from points of introduction at low elevations. According to these authors, the drop in alien species numbers along altitude represents an ecological ltering, e.g. the number of species with a narrow altitudinal range decreases with increasing altitude. The spatial resolution of our data is not high enough to test this hypothesis, but there is some indirect evidence supporting ecological ltering. First, the slope of the regression between maximum altitude and altitudinal range was steeper in alien than in native species, indicating a stronger relationship between a species altitudinal range and the highest altitude it reaches. Second, peak owering (the month with most species in ower) markedly differed between native and alien species. Peak owering was rather homogenous in alien species among vegetation zones, whereas in native species, the position of peak owering shifted from spring to summer with increasing altitude. This indicates that alien species are not adapted to high altitudes and their presence is likely the result of a broad climatic tolerance. 4.5. Implications for biodiversity conservation High mountain areas are often considered to be at low risk of being invaded by alien species (Millenium Ecosystem Assessment, 2003). This might change in future as human caused disturbances increase (McDougall et al., 2011), resulting in an inux of new species (Khuroo et al., 2010). The presence of alien plants at high altitudes in Kashmir Himalaya highlights the potential for increasing plant invasions, and species with a large altitudinal range are likely to reach new sites (Becker et al., 2005). Here,

assessments of ecological impacts imposed by the alien species seem to be necessary (Khuroo et al., 2008). Human pressure on natural ecosystems and associated habitat alterations are intense in this Himalayan region (Oza, 2003), asking for measures towards mitigating the effects of alien species.

Acknowledgments AAK and AHM are thankful to the Head, Department of Botany, University of Kashmir, Srinagar, for providing necessary facility during the course of present study. The supporting staff at the Centre for Biodiversity and Taxonomy, University of Kashmir is acknowledged for their help in the eld and herbarium. We thank Eric Lind for uselful comments on earlier versions of the manuscript, as well as two anonymous reviewers.

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Maheshwari, J.K., 1962. Studies on the naturalised ora of India. In: Maheswari, P., Johri, B.M., Vasil, I. K. (Eds.). Proceedings of the Summer School of Botany. Sree Sarawati Press, Clacutta, pp. 156170. Maheshwari, J.K., 1979. Alien ora of India. In: Khoshoo, T.N., Nair, P.K.K. (Eds.). Progress in plant research. Today and Tomorrow Printers & Publishers, New Delhi, pp. 219228. Naqshi, R.A., Dar, G.H., Javeid, G.N., Kachroo, P., 1988. Malvaceae of Jammu and Kashmir State, India. Ann. M. Bot. Gard. 75, 14991524. Pandey, D.S., 2000. Exotics. In: Flora of India (Introductory Volume, Part II), Singh, N.P., Singh, D.K., Hajra, P.K., Sharma, B.D. (Eds.). Botanical Survey of India. Kolkata, India, pp. 266301. Polunin, O., Stainton, A., 1986. Flowers of the Himalaya. Oxford University Press, New Delhi, India. Singh, G., Misri, B., 1974. Some exotic ornamentals of Kashmir. Indian J. Hort. 31, 834. Singh, G., Kachroo, P., 1983. Exotic trees and shrubs of Kashmir. Indian For. 109, 6076. Singh, G., Kachroo, P., 1987. Forest ora of Srinagar and its neighbourhood. Bishen Singh Mahendra Pal Singh, Dehradun, India. Sharma, B.M., Kachroo, P., 1981. Plants of Jammu and plants of neighbourhood. Bishen Singh Mahendra Pal Singh, Dehradun, India. Sharma, B.M., Jamwal, P.S., 1988. Flora of upper Lidder Valleys of Kashmir Himalaya. Scientic Publishers, Jodhpur, India. Stewart, R.R., 1972. An annotated catalogue of the vascular plants of West Pakistan and Kashmir. Fakhri Press, Karachi, Pakistan. Swami, A., Gupta, R.K., 1998. Flora of Udhampur. Bishen Singh Mahendra Pal Singh, Dehradun, India.

Table B1 (continued ) Alien species Caesalpinia bonduc Campsis grandiora Campsis radicans Cassia stula Castanea sativa Casuarina equisetifolia Catalpa bignonioides Celtis australis Celtis eriocarpa Cercis siliquastrum Chaenomeles speciosa Cissampelos pareira Citrus reticulata Citrus sinensis Clematis gouriana Clerodendrum chinense Cordia dichotoma Crataeva adansonii Croton bonplandianus Cryptolepis buchananii Cryptomeria japonica Cupressus arizonica Cupressus cashmeriana Cupressus sempervirens Cupressus torulosa Cycas revoluta Cydonia oblonga Deutzia compacta Diospyros kaki Diospyros montana Dodonaea viscosa Ehretia acuminata var. serrata Elaeagnus angustifolia Eriobotrya japonica Eucalyptus alba Euonymus echinatus Euonymus hamiltonianus Euonymus japonicus Fatsia japonica Ficus carica Ficus hispida Ficus virens Forsythia viridissima Ginkgo biloba Gmelina arborea Hedera nepalensis Hibiscus hirtus Hibiscus rosa-sinensis Hibiscus syriacus Hibiscus vitifolius Hiptage benghalensis Holboellia latifolia Hymenodictyon orixense Indigofera tinctoria Ipomaea carnea subsp. stulosa Jasminum humile Jasminum rottlerianum Jatropha curcas Juglans nigra Juglans regia Kerria japonica Koelreuteria paniculata Laburnum anagyroides Lagerstroemia indica Lantana camara Laurus nobilis Lavandula angustifolia Lespedeza cuneata Lettsomia thomsoni Ligustrum lucidum Ligustrum ovalifolium Ligustrum vulgare Lonicera japonica Lonicera microphylla Magnolia grandiora Magnolia kobus Life form ES DC DC DT DT ET DT DT DT DS DS DCL ET ET DCL DS DT DT DSS DCL ET ET ET ES ET ES DT DS DT DT ES DT DT ET DT EC ES ES ES DT ET DT DS DT DT ECL ES ES ES DS ECL EC DT DS DS DS DS DS DT DT DS DT DT DT ES ET DS DSS DS DT DS DS EC DS ET DT Flowering 89 67 67 35 56 123 57 35 34 57 46 48 45 45 45 34 56 45 67 34 57 67 67 67 37 46 46 45 45 24 57 67 46 46 56 46 57 79 34 68 36 34 78 45 89 48 89 910 79 23 46 89 810 45 46 34 45 34 45 45 56 46 45 23 56 56 79 89 45 45 46 78 67 35 34 Altitudes (m) 10001600 16001700 16001700 12001500 16001800 10001300 17001800 13002200 10001300 16001700 16001700 14001900 9001500 11001500 14001800 10001300 7001500 12001700 7001500 7001500 16001700 16001900 16001700 14001900 16002800 10001400 16001800 16001800 17001900 9001400 8001400 5001700 12001900 11001800 8001600 15002400 16002700 10001700 16001800 18002100 12001800 9001400 15001700 15001700 9001300 10003000 8001100 14002200 16001800 8001900 9001200 10001400 8001200 9001600 9001900 15003000 9001500 8001500 16001700 16003000 16001800 16001700 16002100 15001700 10001500 16001700 16001700 10001900 12001500 16002000 16001700 16001700 13002000 28003200 12001600 16001700

Appendix B. List of alien and native woody species of Kashmir Himalaya, India. Species names follow GRIN Taxonomy Database (www.ars-grin.gov). Growth form (life forms): DCL deciduous climber, DS deciduous shrub, DSS deciduous subshrub, DT deciduous tree, ECL evergreen climber, ES evergreen shrub, ESS evergreen subshrub, and ET evergreen tree. Flowering periods are given in calendar months, altitudes in metres above sea level. See Table B1 here.
Table B1 Alien species Abutilon bidentatum Acacia farnesiana Acacia modesta Acacia nilotica Acacia torta Acer cappadocicum Acer negundo Aesculus indica Ailanthus altissima Albizia julibrissin Albizia lebbeck Amorpha fruticosa Ampelopsis vitifolia Antidesma acidum Azadirachta indica Bougainvillea glabra Bougainvillea spectabilis Bridelia retusa Bridelia verrucosa Buddleja alternifolia Buddleja asiatica Buddleja davidii Buddleja lindleyana Life form DS DS DT DT DCL DT DT DT DT DT DT DS DS DS DT DCL DCL DT DT ES ES DS DS Flowering 79 24 34 711 910 56 45 46 57 56 46 45 56 67 35 911 89 89 910 34 24 89 78 Altitudes (m) 11001500 12001400 11001500 9001500 11001700 12002300 15002500 12002500 15002200 14001600 8001600 16001700 13001500 12001600 11001800 8001100 9001300 7001300 12001500 16001700 16001700 16001700 16001800

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Table B1 (continued ) Alien species Magnolia liliiora Malus baccata Malus pumila Melia azedarach Mimosa pudica Morus alba Morus macroura Morus nigra Morus serrata Murraya koenigii Myrsine africana Nerium oleander Olea europaea subsp. cuspidata Osmanthus fragrans Paeonia suffruticosa Parthenocissus himalayana Parthenocissus quinquefolia Parthenocissus tricuspidata Philadelphus confusus Philadelphus incanus Philadelphus pubescens Philadelphus tomentosus Pinus canariensis Pinus halepensis Pinus radiata Pistacia integerrima Pistacia vera Platanus occidentalis Platanus orientalis Pogostemon benghalensis Populus alba Populus deltoides Populus euphratica Populus nigra Prinsepia utilis Prunus armeniaca Prunus bokhariensis Prunus cerasifera Prunus cerasus Prunus domestica Prunus dulcis Prunus persica Pterospermum acerifolium Punica granatum Pyrus communis Pyrus pyrifolia Quercus baloot Quercus robur Ricinus communis Robinia pseudoacacia Rosa banksiae Rosa brunonii Rosa chinensis Rosa foetida Rosa moschata Rosa multiora Rubus ellipticus Rubus niveus Rubus ulmifolius Salix alba Salix babylonica Salix fragilis Salix viminalis Sambucus nigra Sapindus mukorossi Sapium sebiferum Sequoiadendron giganteum Sida cordifolia Smilax menispermoidea Solanum anguivi Solanum erianthum Solanum incanum Solanum pseudocapsicum Solanum torvum Sophora japonica Sophora mollis Life form ET DT DT DT DSS DT DT DT DT DS ES ES DT ET DS ECL ECL DS DS DS DS DS ET ET ET DT DT DT DT DS DT DT DT DT DS DT DT DT DT DT DT DT DT DS DT DT DT DT DSS DT ECL DC DCL DS DCL DS ES DS DS DT DT DT DT DS DT DT ET DSS DC DSS DS DS DSS DS DS DS Flowering 45 46 35 35 68 34 56 34 56 45 34 45 34 45 45 67 67 67 45 56 45 35 57 57 56 57 57 45 45 34 45 45 23 45 24 35 34 34 45 45 34 34 35 57 45 45 45 56 57 45 45 67 35 67 57 45 24 68 89 35 34 34 34 67 78 67 57 67 67 34 23 56 56 67 78 68 Altitudes (m) 16001700 18003200 14002700 12001700 12001500 12003200 15001700 13002000 13002300 12001500 12001700 12002100 15001700 16001800 16001800 11001500 15002000 16001700 16001700 16001800 16001700 14002200 16002000 16001800 17001900 7002000 11001700 16001700 13002100 8001400 16003000 17001800 16003200 17003000 12001900 14003200 16001800 16002000 17002100 17001900 17001800 17002300 9001400 11002100 14003100 12002000 16001700 17001900 9001800 11002000 16001800 19002800 16001800 14001800 17001900 16001900 12001800 17002800 15002300 12003000 16003000 16002800 16003200 16001700 11001500 7001600 18001900 8001400 10001800 10001400 12001700 11001400 14003300 12001400 16001900 14001900

Table B1 (continued ) Alien species Sorbus domestica Spartium junceum Spiraea canotoniensis Spiraea hypericifolia Spiraea prunifolia Syringa emodi Syringa vulgaris Tamarix dioca Tamarix parviora Tamarix troupii Taxodium distichum Taxus baccata Thuja orientalis Tilia platyphyllos Toxicodendron succedaneum Triumfetta rhomboidea Tylophora hirsuta Viburnum cotinifolium Viburnum nervosum Vitex negundo Vitis labruscana Vitis riparia Vitis vinifera Weigela orida Yucca aloifolia Native species Abelia triora Abies pindrow Abies spectabilis Abrus precatorius Abutilon indicum Abutilon ramosum Acacia catechu Acer acuminatum Acer caesium Acer oblongum Acer pentapomicum Acer sterculiaceum Acer villosum Adhatoda vasica Aegle marmelos Albizia odoratissima Alnus nitida Ampelocissus latifolia Arachne cordifolia Aralia cashemirica Asparagus adscendens Asparagus gracilis Aspidopterys wallichii Aster albescens Astragalus bakeri Astragalus candolleanus Astragalus chlorostachys Astragalus grahamianus Astragalus oplites Astragalus zanskarensis Atylosia mollis Barleria prionitis Bauhinia vahlii Bauhinia variegata Berberis calliobotrys Berberis chitria Berberis edgeworthiana Berberis glaucocarpa Berberis jaeschkeana Berberis kashmiriana Berberis kunawurensis Berberis lycium Berberis orthobotrys Berberis pachyacantha Berberis royleana Berberis ulicina Betula utilis Boehmeria platyphylla Life form DS DS DS DS DS DS DS DS DS DT ET ES ES DT DT DS ES DS DS DS DCL DCL DCL DS ES Life form DS ET ET DCL DSS DSS DT DT DT DT DT DT DT ES DT ET DT DS DS DS DSS DSS ECL DS DS DSS DSS DSS DS DSS DCL DS ECL DT DS DS DS DS DS DS DS ES DS DS DS DS DT DS Flowering 67 67 57 46 56 67 45 45 45 89 78 68 67 56 56 67 34 45 34 45 67 67 57 46 910 Flowering 67 68 78 89 79 89 46 46 45 45 34 34 57 23 67 56 68 67 45 89 910 810 810 79 58 78 67 47 68 78 68 912 45 34 46 56 67 78 57 67 57 56 46 45 45 67 56 78 Altitudes (m) 18002200 16001800 16002000 14001800 16001700 16002100 17001800 10001700 16001700 9001500 18002000 16001700 16001800 16001700 17002300 9002100 16002200 13003400 18002400 11001500 16001800 16001700 10001800 16001700 10001700 Altitudes (m) 15003400 23002900 26003100 12001400 10001500 9001600 8001400 21003000 17002400 15002500 16002300 13002100 23003000 12001600 7001300 12001900 16001900 11001200 12002100 17002200 12001900 14001700 10001500 11001300 14002000 30003500 18003000 15002200 30003500 28003600 11002100 12001600 11002000 16001900 17001900 20003700 20002900 22003700 30003300 27003100 11003100 17002400 28002900 15002500 16001800 23003700 23003000 8002100

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Table B1 (continued ) Native species Bombax ceiba Buddleja crispa Butea monosperma Buxus papillosa Buxus wallichiana Callicarpa macrophylla Calotropis procera Campylotropis macrostyla Capparis decidua Capparis sepiaria Capparis spinosa Caragana brevifolia Caragana brevispina Caragana versicolor Carissa opaca Carpinus viminea Caryopteris bicolour Casearia elliptica Cassiope fastigiata Cedrela serrata Cedrus deodara Celastrus paniculata Clematis buchananiana Clematis connata Clematis grata Clematis graveolens Clematis montana Clematis orientalis Clematis puberula Clematis tibetiana Cocculus laurifolius Cocculus pendulus Colebrookea oppositifolia Cordia gharaf Cordia vestita Coriaria nepalensis Corylus ferox Corylus jacquemontii Cotinus coggygria Cotoneaster acuminatus Cotoneaster afnis Cotoneaster aitchisonii Cotoneaster frigidus Cotoneaster humilis Cotoneaster integerrima Cotoneaster microphylla Cotoneaster minuta Cotoneaster nummularia Cotoneaster obovatus Cotoneaster rosea Crataegus songarica Crotalaria spectabilis Dalbergia sissoo Daphne mucronata Daphne papyracea Daphne retusa Debregeasia salicifolia Deeringia amaranthoides Desmodium elegans Desmodium motorium Desmodium velutinum Deutzia staminea Dictamnus albus Diospyros lotus Ehretia aspera Ehretia laevis Elaeagnus parvifolia Elaeodendron roxburghii Elsholtzia fruticosa Embelia robusta Ephedra gerardiana Ephedra intermedia Ephedra prezewalskyi Ephedra procera Ephedra regeliana Eranthemum pulchellum Life form DT DS DT ES ES ES ES DS DS ES DS DS DS DS ES DT DS DT DS DT ET DC DCL DCL DCL DCL DCL DCL DCL DCL ES ES DS DS DT DS DT DT DS DS DS DS DS DS DS ES DS DS DS DS DT DSS DT ES DS DS ES DS DS DSS DSS DS DS DT DS DT DS ET DS DS ES ES ES ES ES ES Flowering 23 45 34 37 35 89 27 78 46 56 56 68 56 78 67 68 23 45 67 89 56 46 89 78 78 78 45 89 89 57 45 39 23 46 45 34 45 34 57 56 56 56 67 67 67 56 56 57 57 67 56 69 36 45 34 57 56 45 67 89 79 45 67 56 56 34 45 56 89 34 57 78 56 46 67 35 Altitudes (m) 9001400 18002300 9001700 10001600 13001700 11001400 12001400 16001900 10001500 11001400 21003200 23004500 19002800 29003800 11001900 12002600 11001500 13002500 28004500 20002500 20002600 12001400 15002400 16002200 14002400 15002100 17002100 12003500 13001600 28003600 10001900 9001700 17002700 7001600 8001500 16002500 26003200 16003300 15002500 21004000 19002400 18002200 22003000 28003700 26004000 25003100 17001900 20003200 15002000 18002900 15002200 19002700 8001600 14002300 16002100 30003700 10002300 11001600 12002600 12001900 12001700 13001600 25003000 17001900 10001500 6001200 16003000 10001300 21002500 12001600 25004500 27004000 30004000 25003800 19002900 11001700

Table B1 (continued ) Native species Erythrina suberosa Euonymus mbriatus Euphorbia royleana Ficus bengalensis Ficus palmata Ficus racemosa Ficus religiosa Ficus sarmentosa Ficus semicordata Flacourtia indica Flemingia macrophylla Flueggea virosa Fraxinus hookeri Fraxinus xanthoxyloides Gaultheria trichophylla Glochidion velutinum Grewia optiva Grewia tenax Gymnosporia royleana Helicteres isora Helinus lanceolatus Hippophae rhamnoides Holarrhena pubescens Holmskioidia sanguinea Hypericum dyeri Hypericum hookerianum Hypericum oblongifolium Hypericum uralum Ichnocarpus frutescens Ilex dipyrena Ilex excelsa Indigofera atropurpurea Indigofera cassioides Indigofera hebepetala Indigofera heterantha Isodon rugosus Jasminum arborescens Jasminum auriculatum Jasminum dispermum Jasminum multiorum Jasminum ofcinale Juniperus communis Juniperus semiglobosa Juniperus squamata Kydia calycina Lanea coromandelica Lepidagathis cuspidata Leptodermis lanceolata Leptodermis virgata Lespedeza elegans Lespedeza eriocarpa Lonicera angustifolia Lonicera asperifolia Lonicera discolor Lonicera hispida var. bracteata Lonicera myrtillus Lonicera obovata Lonicera purpurascens Lonicera quinquelocularis Lonicera semenovii Lonicera spinosa Lonicera vaccinioides Lonicera webbiana Lycium ruthenicum Mahonia borealis Mallotus philippensis Measa montana Meliosma simplicifolia Mimosa rubicaulis Mitragyna parvifolia Moringa oleifera Murraya paniculata Myricaria davurica Myricaria germanica Neolitsea pallens Nyctanthes arbour-tristis Life form DT DT DS ET DS DT DT ECL ET DT DS DT DT DT ES DT DT DS ES DT DS DS DT ES DS DS ES DS ES ET ES DS DS DS DS DS DS DS DS ES DCL ES ES ES DT DT DSS DS DS DS DS DS DS DS DS DS DS DS DS DS DS DS DS DS ES DT DS ET DS DT DT DS DS DS ET DT Flowering 34 57 67 23 34 57 57 57 59 34 79 57 57 46 57 45 46 46 78 78 35 34 56 89 67 67 45 49 46 45 56 89 34 57 57 810 45 89 57 46 57 67 78 67 910 34 123 68 78 89 67 56 79 56 56 67 67 68 46 67 67 78 67 67 78 89 13 46 78 78 36 34 67 67 35 911 Altitudes (m) 9001200 16002300 15001700 8001800 7001600 8001400 8001400 7001200 11001700 17002500 10001700 15001700 14003000 16002800 27004500 12001500 11001800 10001500 10001600 10001400 10001600 21003600 11001600 10001300 14002300 18003200 10001600 15001800 11002000 15002300 11002300 9001500 11001500 16002800 12002800 14002100 8001500 12001400 14002300 8001300 15002300 21003100 22002600 27003500 8001200 8001400 11002000 12001800 11002400 16002500 18002600 18002300 27004000 28003300 20003200 24003000 30004400 30003500 11002300 35004800 36004600 30004000 21004300 29003500 20003000 7001500 9001400 7001200 11001500 8001500 11001800 13001700 17002800 22003400 11002300 16001800

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Table B1 (continued ) Native species Oroxylum indicum Paeonia emodi Parrotiopsis jacquemontiana Parthenocissus semicordata Persea duthiei Persea odoratissima Phyllanthus emblica Picea smithiana Pinus gerardiana Pinus roxburghii Pinus wallichiana Pittosporum napaliense Populus ciliata Premna barbata Premna latifolia Prunus cornuta Prunus prostrata Prunus tomentosa Pyrus pashia Quercus oribunda Quercus leucotrichophora Quercus semecarpifolia Randia spinosa Randia tetrasperma Reinwardtia indica Rhamnus pentapomica Rhamnus prostrata Rhamnus purpureus Rhamnus triquetra Rhamnus virgata Rhododenron arboreum Rhododendron campanulatum Rhododendron hypenanthum Rhododendron lepidotum Rhus javanica Rhus punjabensis Ribes alpestre Ribes glaciale Ribes orientale Ribes villosum Rosa corymbifera Rosa hemispherica Rosa macrophylla Rosa webbiana Rubus antennifer Rubus biorus Rubus macilantus Rubus paniculatus Rubus pedunculosus Rubus pungens Sabia campanulata Sageretia liformis Sageretia thea Salix acmophylla Salix daphnoides Salix denticulata Salix abellaris Salix karelinii Salix lindleyana Salix oxycarpa Salix pycnostachya Salix sericocarpa Salix wallichiana Salix wilhelmsiana Sarcococca saligna Sida rhombifolia Skimmia anquetilia Smilax aspera Smilax glaucophylla Smilax vaginata Sophora moorcroftiana Sorbaria tomentosa Sorbus lantata Spermadictyon suaveolens Spiraea arcuata Spiraea bella Life form ET DS DS DCL ET ET DT ET ET ET ET DS DT DT DT DT DS DS DT ET ET DT DT DS DS DS DS DT DS DS ET ES ES ES DS DT DS DS DS DS DS DS DS DS DS DS ES DS DS DS DCL DS ES DT DS DS DS DS DS DT DS DT DS DS ES DS ES DC DC DSS DS DS DT DS DS DS Flowering 67 56 67 67 34 34 45 57 67 45 68 67 34 56 45 45 45 34 45 67 34 56 57 45 34 34 67 46 89 46 45 57 67 78 89 57 56 57 56 67 56 57 45 46 57 45 45 45 56 46 45 911 78 24 34 67 67 67 67 45 57 56 46 57 89 911 67 56 47 67 57 67 45 1012 78 67 Altitudes (m) 11001200 14002000 17002100 10001800 10002000 12002200 7001500 24002800 16002500 11001500 19003000 11001500 14002500 11001500 10001500 18002600 18002300 17002400 12002200 18002200 14002800 8001900 11001600 9001700 11002500 10001800 17003000 15003000 8001500 18002200 15002800 31004000 30003600 30004000 12002400 20002300 29003900 24004200 21004000 32004000 20002700 16002900 19002900 14003100 17002200 14002100 14003000 11001500 17002000 18002900 8001900 12001500 15002700 14002100 16003000 23002900 26003600 22003200 27003000 16002000 18003200 18002200 19002300 28003300 18003000 11001700 22003000 12001500 11002300 13001800 28003500 18002500 20002500 10001500 13001900 20003200

Table B1 (continued ) Native species Spiraea canescens Spiraea vaccinifolia Staphylea emodi Stereospermum suaveolens Strobilanthes urticifolia Symplocos paniculata Syzygium cumini Tamaricaria elegans Tamarix indica Taxus wallichiana Terminalia arjuna Terminalia bellirica Terminalia chebula Tinospora malabarica Toona ciliata Trema politoria Ulmus villosa Ulmus wallichiana Urena lobata Vallaris solanacea Viburnum grandiorum Viburnum mullaha Villebrunea frutescens Vitis jacquemontii Vitis parvifolia Vitis trifolia Wendlandia heynei Wikstroemia canescens Woodfordia fruticosa Wrightia tomentosa Xylosma longifolium Zanthoxylum armatum Ziziphus mauritiana Ziziphus oxyphylla Ziziphus vulgaris Life form DS DS DS DT DSS DT ET DS DS ET ET ET ET DCL DT ET DT DT DSS ES DS DS ES DCL DCL DCL DT DS ES DT DS DS DT DS DS Flowering 57 57 45 56 711 56 56 67 67 67 45 46 56 79 45 56 34 78 710 45 34 910 45 56 67 78 35 56 23 67 1011 45 89 67 68 Altitudes (m) 14002600 12001900 18002900 10001200 12002600 14002400 9001400 25003500 28003400 22002800 7001200 7001700 9001300 8001600 15002300 10001300 11002100 16003200 9001200 11002300 17002500 13003400 16002000 10001400 10001500 11001300 8001500 18002800 12001900 11002300 9001400 10001700 14002200 16002200 15001900

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