Vegetation dynamics in the western himalayas, diversity indices and climate change

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Vegetation provides the first tropic trophic level in mountain ecosystems and hence requires proper documentation and quantification in relation to abiotic environmental variables both at individual and aggregate levels. The complex and dynamic Himalayas with their varying climate and topography exhibit diverse vegetation that provides a range of ecosystem services. The biodiversity of these mountains is also under the influence of diverse human cultures and land uses. The present paper is not only first of its kind but also quite unique because of the use of modern statistical techniques for the quantification of Diversity Indices of plant species and communities. The vegetation was sampled in three categories, i.e., trees, shrubs and herbs, as follows: a height of ≥ 5m were classified in the tree layer, shrubs were all woody species of height 1m and 5m and, finally, the herb layer comprised all herbaceous species less than 1m in height. The presence/absence of all vascular plants was recorded on pre-prepared data sheets (1, 0 data). For the tree layer, the diameter of trees at breast height was measured using diameter tape. Coverage of herbaceous vegetation was visually estimated according to Daubenmire and Braun Blanquet methods. It gives overall abundance of vascular plants on one hand and composition of these species on the other. Data was analysed in Canonical Community Coordination Package (CANOCO) to measure diversity indices of plant communities and habitat types. Results for five plant communities/habitat types indicated that plant biodiversity decreased along the altitude. Shannon Diversity Index values range between 3.3 and 4. N2 index and Index of Sample Variance were also designed. All of these Diversity Indices showed the highest values for the communities/habitats of north facing slopes at middle altitudes. Higher plant diversity at these slopes and altitudes can be associated to the period of snow cover which is longer and a relatively denser tree cover as compared to the southern slopes and hence the soil has high moisture which supports high biodiversity in return. Global warming causes desertification in number of fragile mountain ecosystem around the globe. These findings suggest that species diversity decreases along the measured ecological gradient under the influence of deforestation coupled with global climatic change.

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Vegetation dynamics in the western himalayas, diversity indices and climate change

  1. 1. Sci., Tech. and Dev., 31 (3): 232-243, 2012*Author for correspondence E-mail: shuja60@gmail.comVEGETATION DYNAMICS IN THE WESTERNHIMALAYAS, DIVERSITY INDICES AND CLIMATECHANGESHUJAUL M. KHAN1*, SUE PAGE2, HABIB AHMAD3, HAMAYUN SHAHEEN4AND DAVID HARPER51Department of Botany, Hazara University, Mansehra, Pakistan.2Department of Geography, University of Leicester, UK3Department of Genetics, Hazara University, Mansehra, Pakistan.4Department of Botany, Azad Kashmir University, Muzaffarabad, Pakistan.5Department of Biology, University of Leicester, UKAbstractVegetation provides the first tropic trophic level in mountain ecosystems and hencerequires proper documentation and quantification in relation to abiotic environmental variablesboth at individual and aggregate levels. The complex and dynamic Himalayas with theirvarying climate and topography exhibit diverse vegetation that provides a range of ecosystemservices. The biodiversity of these mountains is also under the influence of diverse humancultures and land uses. The present paper is not only first of its kind but also quite uniquebecause of the use of modern statistical techniques for the quantification of Diversity Indices ofplant species and communities. The vegetation was sampled in three categories, i.e., trees,shrubs and herbs, as follows: a height of ≥ 5m were classified in the tree layer, shrubs were allwoody species of height 1m and 5m and, finally, the herb layer comprised all herbaceousspecies less than 1m in height. The presence/absence of all vascular plants was recorded on pre-prepared data sheets (1, 0 data). For the tree layer, the diameter of trees at breast height wasmeasured using diameter tape. Coverage of herbaceous vegetation was visually estimatedaccording to Daubenmire and Braun Blanquet methods. It gives overall abundance of vascularplants on one hand and composition of these species on the other. Data was analysed inCanonical Community Coordination Package (CANOCO) to measure diversity indices of plantcommunities and habitat types. Results for five plant communities/habitat types indicated thatplant biodiversity decreased along the altitude. Shannon Diversity Index values range between3.3 and 4. N2 index and Index of Sample Variance were also designed. All of these DiversityIndices showed the highest values for the communities/habitats of north facing slopes at middlealtitudes. Higher plant diversity at these slopes and altitudes can be associated to the period ofsnow cover which is longer and a relatively denser tree cover as compared to the southernslopes and hence the soil has high moisture which supports high biodiversity in return. Globalwarming causes desertification in number of fragile mountain ecosystem around the globe.These findings suggest that species diversity decreases along the measured ecological gradientunder the influence of deforestation coupled with global climatic change.Keywords: Biodiversity, Diversity Index, Climate, Mountain Ecosystem, Western Himalayas,Vegetation.IntroductionMountains are the most remarkable landforms on earth surface with prominent vegetationzones based mainly on altitudinal and climaticvariations. Variations in aspects also enhancehabitat heterogeneity and bring micro-environmental variation in vegetation pattern(Clapham, 1973; Khan et al., 2011b). In north-western Pakistan, three of the world’s highestmountain ranges, i.e., the Himalayan, Hindu Kush
  2. 2. VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE 233and Karakoram, come together, ensuring highfloral diversity and phytogeographic interest.Plant biodiversity survives at the edge of life atthese high mountains where climatic changes aremore visible and species extinction very rapid.Studies on mountainous vegetation around theglobe show that ecological amplitude of alpinespecies shifted to even higher elevations over therecent decades. Species preferred temperatures inhigher altitudinal zones. Therefore, many alpinespecies are under the risk of extinction due totheir requirements for germination andreproduction (Grabherr et al., 1994; Holzinger etal., 2008). Unlike the eastern Himalayas, wheremonsoon-driven vegetation predominates underhigher rainfall and humidity (Chawla et al., 2008;Dutta and Agrawal, 2005; Anthwal et al., 2010;Behera et al., 2005; Roy and Behera, 2005), thevegetation in the western Himalayas in general(Chawla et al., 2008; Kukshal et al., 2009;Shaheen et al., 2011; Ahmad et al., 2009; Dickoréand Nüsser, 2000; Shaheen et al., 2012) and in theNaran Valley (Khan et al., 2011b) in particularhave closer affinities with that of the Hindukushmountains, which have a drier and cooler climate(Ali and Qaiser, 2009; Wazir et al., 2008;Noroozi et al., 2008). A recent study showed thatwith the passage of time, homogeneity takes placein the vegetation of a region due to continuousdominance of certain resistant and vigorousspecies. This phenomenon is further enhanced byselective utilisation of species by humanintervention (Del Moral et al., 2010). Similarly,the tree-line vegetation and indicator species shiftupwards due to climate change.Mountainous vegetation has manifoldfunctions, not only within the system where itexists regionally in the lowland ecosystem byregulating floods and flow in streams andglobally in combating the climate change andgreenhouse effects. Regionally, shrubbyvegetation of high altitude regulates avalanchemovements and protects soil but in majority ofplaces, it is threatened due to human and climaticinfluences (Hester and Brooker, 2007). Plantbiodiversity is necessary for regulation of overallsystem in the mountain, e.g., the Himalayas is thebirth place for 10 largest rivers in the Asia and abig and important carbon sink. Ecologicalchanges in the Himalayas affect global climate bybringing changes in temperature and precipitationpatterns of the world. Himalayan Vegetation isdiverse and range from tropical evergreen speciesin the south east to thorn steppe and alpinespecies in the north western parts (Behera andKushwaha, 2007). Melting of its snow in aregular fashion is related to its vegetation cover.Irregular loss of its ice might have dangerous risein world sea-levels (Xu et al., 2009). Thesemountains are extremely sensitive to globalclimatic change. Such hazardous glimpses havealready been observed in the form of flood inPakistan, India, China and Thailand in the lastthree years.The Naran Valley is located at the far west ofthe Western Himalayas on the border with theHindu Kush range which lie to the west and nearto the Karakorum Range to the north (Figure 1).Due to this transitional location the valley hostrepresentative vegetation types from all threemountain ranges. Monsoon winds are mainsource of precipitation and also a primary force ofcontrolling erosion and climatology over millionsyears of time and thus modify its climate,topography and vegetation of Himalayas but inthe western Himalaya, especially in Naran Valley,high mountains situated at the opening of thevalley act as barriers to the incoming summermonsoon from the south and limit its saturationinto upper northern parts. Thus, summers remaincool and relatively dry and make most of thevalley as a dry temperate-type of habitat. Thereare clear seasonal variations. Total averageannual precipitation is low at only 900-1000mmbut there is heavy snowfall in winter which mayoccur any time from November to April (averageannual snowfall 3m). The range reflects a sharpincrease in depth of snow with increasing altitude.There is a distinct wet season in January-Aprilwhilst the driest months of the year are June-November. As far as temperature of the area isconcerned, most of the year, it remains below10°C. December, January, February and Marchare the coldest months of the year in whichtemperature remains around the freezing point oreven below most of the times. June to August isthe main season for growth, with average daytimetemperatures in the range 15-20°C. Geologically,the valley is located where the Eurasian andIndian tectonic plates meet and where the aridclimate of the western Eurasian mountains givesway to the moister monsoon climate of the Sino-
  3. 3. 234 SHUJAUL M. KHAN ET AL.Japanese region (Qaiser and Abid, 2005;Takhtadzhian and Cronquist, 1986; Kuhle, 2007).The valley thus occupies a unique transitionalposition in the region. In remote mountainousvalleys like the Naran, the complexity ofecosystems, their inaccessibility and the cost andtime factors make it extremely difficult to observeeach and every aspect of vegetation features.Perhaps those were the reasons that in spite of itshigh phytogeographic importance, there havebeen no previous quantitative studies of thevegetation in this valley. The present study is notonly the first of its kind but also quite uniquebecause of the use of modern statisticaltechniques for the quantification of plant speciesand communities along geo-climaticenvironmental gradients that has limitedcomparator studies in this region (Wazir et al.,2008; Saima et al., 2009; Dasti et al., 2007). TheNaran Valley, which is floristically located in theWestern Himalayan province of the Irano-Turanian region, forms a botanical transitionalzone between the moist temperate (from theSouth East) and dry temperate (from the NorthWest) vegetation zones of the Hindu Kush andHimalayan mountain ranges, respectively. Aphyto-climatic gradient of vegetation based onlife forms further emphasizes the nature of thearea and makes apparent the transitional positionof the region, though predominated by alpinespecies (Khan et al., 2011b & c).The quantitative approaches to vegetationdescription and analyses deployed in this studynot only fill methodological deficiencies (Khan,2012) and gaps in the literature, i.e., evaluation ofdiversity indices but also provide a firm basis forextending this approach to the adjacent mountainsystems that are in need of up to date vegetationmapping (Fosaa, 2004; Mucina, 1997). Inaddition, the present study documents andprovides suggestions for the conservation ofmountain plant biodiversity under a scenario ofcontinuous human exploitation and climatechange. It is necessary to maintain ecosystemservices in general and food security in particular,not only within mountain system but also for thepeople and ecosystems of the lowlands thatdepend on those mountains (Rasul, 2010;Manandhar and Rasul, 2009; Sharma et al., 2010;Khan et al., 2011a).Fig. 1. Physiographic map showing the elevation zones and position of the Naran Valley, westernHimalayas (study area) in relation to Karakorum and Hindu Kush mountains
  4. 4. VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE 235MethodologyIn total of 432 quadrats in 144 replicates, thevegetation was sampled in three layers i.e., trees,shrubs and herbs. Quadrats sizes used in thisstudy were 10x5 (for trees layer), 5x2 (for shrubslayer) and 1x0.5 (for herbs layer). Trees had aheight of ≥ 5m, shrubs were all woody species ofheight in the range 1-5 m and the herb layercomprised all herbaceous species less than 1m inheight. The presence of all vascular plants wasrecord on pre-prepared data sheets (1, 0 data). Forthe tree layer, the diameter of trees at breastheight was measured using diameter tape(Figure 2). This enabled an assessment of treecover values in the quadrats. For the shrub andherb layers, abundance/cover values were visuallyestimated according to a regulated Braun-Blanquet scale later on modified by Daubenmire(Table 1) (Braun-Blanquet et al., 1932;Daubenmire, 1968). Absolute and relative valuesof density, cover and frequency of each vascularplant species at each station were calculated usingphytosociological formulae to finally calculateImportant Value Index (McIntosh, 1978).Fig. 2. Measuring diameter of tree species through diameter tape in a 10x5m quadrat in the field atLalazar, Naran.Table 1. Braun Blanquet covers classes modified by Daubenmire.Cover Class Range of Percent Cover Midpoint1 0-5% 2.5%2 5-25% 15.0%3 25-50% 37.5%4 50-75% 62.5%5 75-95% 85%6 95-100% 97.5%
  5. 5. 236 SHUJAUL M. KHAN ET AL.In the past, it was a very common techniqueto classify plant communities based on ImportantValue Index (IVI) of plants species. In this sort ofcharacterization and classification inphytosociology, the species with the highestImportance Values IV are considered as dominantplant species and communities are attributed withthem. Even today, it is a very common method ofclassifying plant communities. The present dayrevolution of computer based technology in everyfield of science has modified the older techniques.The IVs were calculated by adding the values ofrelative density, cover and frequency and thendividing by 3. In our study, we use IV datamainly for ordination purposes.Data analysisEcologists use number of diversity indices formeasuring the plant species diversity in differenthabitat types and plant communities. CANOCOversion 4.5 (Ter Braak 1988, Ter Braak 1989, TerBraak and Smilauer 2002) was used to analyseIVI data to assess diversity indices. The mainobjective of these statistical packages is toformulate the study more precisely. Asphytosociology is concerned with vegetationitself, the sites in which it occurs and theenvironmental variables related with those siteshence need to be examined in a statisticalframework (Kent and Coker, 2002; 1995,Lambert and Dale, 1964). Plant biologists oftenneed to test hypotheses regarding the effects ofinvestigational factors on whole groups of species(Khan et al., 2011b; Shaheen et al., 2011;Anderson et al., 2006). CANODRAW is a utilityof CANOCO and was used to generate dataattribute plots (graphic forms) ofindicator/characteristic species. It also gives theopportunity to utilise index of number of species,Shannon Weiner index, index of species richness,evenness, sample variance, etc. Diversity indiceswere calculated at the community through dataattribute plot under the DCA function ofCANODRAW.ResultsFloristic composition of the Naran ValleyA total of 198 plant species belonging to 150genera and 68 families were recorded at the 432replicates of relives/quadrats. The division intomajor taxonomical groups (Table 2) indicates thatdicotyledonous angiosperms are the mostabundant. Five plant communities were identifiedand discussed in our first paper from this project(Khan et al., 2011b). There is a clear altitudinalzonations, i.e., (i) the lower altitude (2450-3250m) dominated by temperate vegetation and (ii)the higher altitude (3250-4100 m) dominated bysubalpine and alpine vegetation. The mostabundant plant family was Asteraceae(Compositae) with seventeen species and a 25%share of all species. Rosaceae represented byfourteen species (with a 21% share) was thesecond most species rich family in the study area.Lamiaceae, Ranunculaceae, Poaceae andPolygonaceae were represented by 13, 12, 11 and10 plant species respectively. The remainingfamilies all had less than 10 species each.Importance Values of the top 15 abundant speciesare given in Table 3.Table 2. Taxonomic divisions of recorded plant species.Taxonomic distributionNo. ofFamiliesNo. ofSpeciesDicot 53 161Monocot 9 23Subtotal of Angiosperms 62 184Gymnosperms 3 8SUBTOTAL OFSPERMATOPHYTA65 192Pteridophyta (Ferns and allies) 3 6TOTAL NO. OF PLANTSPECIES68 198Table 3. Important Value Index (IVI) of the top 15species reported from the region.S.No. Name of the SpeciesImportanceValue (IV)1 Abies pindrow 5842 Betula utilis 5823 Sambucus weghtiana 4464 Juniperus communis 3385 Pinus wallichiana 3236 Salix flabillaris 1607 Rosa webbiana 1378 Artemesia absinthium 1359 Rhododendron hypenanthum 11610 Fragaria nubicola 11211 Berberis pseudoumbellata 11112 Thymus linearis 10913 Iris hookeriana 10414 Cedrus deodara 10315 Viola canescens 96Based upon plant growth habit, 12 trees, 20shrubs and 166 herbs shared 6, 10 and 84%,respectively, in the plant diversity of thevegetation of the Naran Valley (Figure 3).
  6. 6. VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE 237Fig. 3. Percent share of various habit forms of vegetation of the Naran Valley.Species richness along environmentalgradientsAnalyses show that the main influencingenvironmental variables are altitude, aspect (slopedirection) and soil depth and that both speciesrichness and diversity vary along these gradients.Analysis of the elevation gradient showed thatspecies richness was higher at lower altitudes andlower at higher elevations. Species β-diversitygradually decreased along the altitudinal gradientas soil depth and temperature decreased.Similarly species richness on slopes with anorthern aspect was slightly higher than thosewith a southern aspect (Fig. 4).Fig. 4. Average β-diversity of plant species along the altitudinal gradient.
  7. 7. 238 SHUJAUL M. KHAN ET AL.Diversity indices using DCA analysisData attribute plots using DeterendedCorrespondence Analysis (DCA) were used tocalculate diversity indices. The first sort ofdiversity index based on species richness was thecalculation of species number per community(Figure 5). The highest number of plant specieswas reported in community 2 at middle altitudenorthern aspect habitats followed by loweraltitude valley bottom habitats. The high diversityof these habitat types can be attributed to highsoil depth with high moisture retaining capacityand relatively high temperature at thosesomewhat lower altitudes. The lowest number ofspecies scored in the index was for thecommunity number 5 at peak elevations abovethe tree line habitats.The Shannon Diversity index values rangebetween 3.3 and 4. Being constituted of bothNorthern and Southern aspect stations,community 1 has the highest value of 3.98amongst all the groups whilst community 5 hasthe lowest value due to narrow ecologicalamplitude (Figure 6). High values of index aredue to the inclusion of considerable number ofstations in a single community.Fig. 5. Index of species number (alpha diversity) amongst all community types through DCA dataattribute plots along the gradients.Fig. 6. Shannon Weiner diversity index at community level through DCA data attribute plots show thegradient of the community.
  8. 8. VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE 239The N2 index is the reciprocal of Sampsondiversity index and is available in theCANODRAW utility of CANOCO. This indexshowed the highest value for community 2followed by community 1 (Figure 7).Index of sample variance showed the highestvariance at community 2 amongst the groups. Asthe altitudinal pattern (Figure 4) showed thatspecies richness is optimum at the middle altitudeespecially on north facing slopes. This indexreconfirmed the phenomenon of species richnessphenomenon through sample variance index(Figure 8).Fig. 7. N2 diversity index at community level through DCA data attribute plots along the gradient of thecommunity.Fig. 8. Index of Sample Variance amongst all community types through DCA data attributeplots along the gradient.The pattern of plant communities in thevalley is largely determined by aspect andaltitude. Four plant communities can be separatedon the basis of these two variables, whilst onecommunity is established under the combinedeffect of lower altitude and greater depth of soil(the third most important variable). Relativelyhigher summer temperatures and soil moisture arethe co-variables associated with the low elevationand deeper soil. Predominantly, community 1reflects the latitudinal gradient of vegetation frommoist temperate to dry temperate along the valleyon either side of the River Kunhar at loweraltitudes. Species diversity and richness areoptimal at the middle altitudes (2800-3400m), incontrast to the lower altitudes (2400-2800m)where direct anthropogenic activities have hadtheir greatest impact.
  9. 9. 240 SHUJAUL M. KHAN ET AL.DiscussionDiversity amongst plant communities/habitattypesShort summer, deep snow, low temperature,intense solar radiation and cold speedy winds inthe valley in general and higher altitude andspecial slope orientation in particular result xericcondition for plant growth and hence the 𝛽-diversity is gradually decreasing both along thealtitudinal and latitudinal gradients of the valley.Though drawing a sharp line in any naturalecosystem of mountains is difficult as the rapidmicro climatic and edaphic variations overlapeach other due to number of driving agencies andhistorical perspectives but based on someindicator species, vegetation zonation andassociation can be established. Both at habitat andcommunity level, the plant species richness anddiversity is strongly influenced by elevation,aspect and soil depth which in turn are associatedwith precipitation and soil moisture. Variousdiversity indices including number of species,Shannon Weiner, sample variance and N2 showmaximum values for the middle altitude northernaspect plant community (Com. 2) followed by thesubalpine (Com. 4) and middle altitude southernaspect plant communities (Com. 3). Similarpatterns of diversity across altitudinal gradientshave been observed in other studies in theHimalayan regions (Kharkwal et al., 2005;Tanner et al., 1998; Vázquez and Givnish, 1998).Implications of the present project for futurestudiesObserving biodiversity is a lifelong processand needs inputs from various disciplines fromthe natural as well as the social sciences. Forbetter ecosystem management and protection,plant biodiversity should be studied at species,community and ecosystem levels. Themountainous valleys need more botanicalexploration for the complete description of theirplant taxa, abundance and conservation status.There are also extensive gaps in information onecosystem services in the Himalayan region.Most of the vegetation studies have been carriedout solitarily either based on scientific approachesor public perception for making inventories. Wealso believe and propose that such approach mustbe statistically sound and communicable toconservationists, planners, politicians and policymakers. Furthermore, we suggest that theidentification of indicator species for specifichabitats will assist in monitoring and assessmentof the biological diversity on the one hand and theeffects of climatic change on the other. Inaddition, a broad scale deterioration of the naturalecosystems due to agriculture, expansion ofroads, increases in population and deforestationcause enormous losses to the natural vegetation.In spite of IUCN recommendations, there is verylimited and small scale documentation on RedList Categories for plant species in the Himalayasas well as in Pakistan more generally (Ali, 2008;Pant and Samant, 2006; Chettri et al., 2008).Being a member state of the CBD, Pakistanshould give top priority to this issue as addressedby Khan (2012) and Khan et al., (2011b & c).The three mountain ranges i.e., theHimalayas, Hindu Kush and Karakorum, provideessential ecosystem services to millions of peopleacross 10 countries of the world (Dong et al.,2010; Khan et al., 2007). These services includeprovisioning, regulating, supporting and culturalservices. In the present study, the main focus wason vegetation mapping and provisioning servicesin one of the Himalayan valley. Most of theprovisioning services considered in the presentstudy can further be evaluated at molecular andbiochemical levels in the future. Beyond thedirect role of plant biodiversity in thesocioeconomics of the mountain people, it isindispensable for the people living in the plainsand hence this mountain valley can be studied forthe evaluation of the other kinds of services,including those of regulatory nature, that itprovides on a broader level, e.g., flood control,erosion control, irrigation water and hydropowerdevelopment. These mountains also provideSupporting Services, e.g., soil formation,biogeochemical and nutrient cycling, etc. Thesemountain systems host characteristic biodiversityon the one hand and a long established andunique cultural diversity on the other.Preservation of their indigenous knowledge andits utilization in environmental management canalso be potential topics for future studies (Fig. 9).
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