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        Impact of patch size on woody tree species richness and abundance in a tropical montane evergreen forest patches of south India

        2018-11-06 06:44:32DharmalingamMohandassMasonCampbellPriyaDavidar
        Journal of Forestry Research 2018年6期

        Dharmalingam Mohandass?Mason J.Campbell?Priya Davidar

        Abstract We examined the effects of forest patch size on woody tree species richness and abundance in tropical montane evergreen forest patches of the Nilgiri region,south India.We sampled woody trees(≥1 cm dbh)from 21 forest patches in the upper Nilgiri hills(>2000 m elevation)and recorded a total of 35,146 individuals of 61 species,45 genera and 30 families.Species richness and abundance of sapling/shrubs (≥ 1 to <10 cm dbh)increased significantly with increasing patch size,but the species richness and abundance of small,medium and larger trees(≥ 10 to<30,≥ 30 to<60 and ≥ 60 cm dbh,respectively)did not.Overall,forest interior species richness and abundance increased significantly with increasing patch size but edge species richness did not.Species richness and abundance of shade-tolerant and shade-demanding tree species also increased with increasing patch size.The abundance of zoochory dispersed tree species was significantly related to increasing patch size,but those dispersed by autochory did not display any clear relationship between patch size and species richness or abundance.Our findings suggest that with increasing forest patch area,tree compositional patterns may be driven by species specific shade-tolerance adaptations and dispersal patterns.Differential responses in these traits by the plant community within the individual habitat zones of forest edge and interiors likely plays a major role in determining the inherent plant community and thus the subsequent ecological processes of forest patches,including their responses to increasing patch area.

        Keywords Dispersal mode·Forest fragments·Shadetolerance ·Species–area relationships ·Western Ghats

        Introduction

        Natural forest patches or fragments are analogues of the theory of island biogeography(MacArthur and Wilson 1963,1967)and have become a central theme for examination in the field of conservation biology.Many studies have focused on species–area relationships(SARs)to estimate the effect of patch size on species richness and abundance and to examine the response of the species within remaining habitats such as forest edges and interiors(Ewers et al.2007;Gonzalez et al.2010;Fahrig 2013;Mohandass et al.2014).Responses by species vary depending on local disturbance and climate in relation to species-specific habitat requirements and remaining habitat area(Pausas and Austin 2001;Martin et al.2007;Gonzalez et al.2010).Different eco-physiological groups of species can also show different responses to fragmentation depending on their life-history traits(Ewers and Didham 2006;Gonzalez et al.2010).Shade tolerance,for instance,ensures tree survival and regeneration in spite of variability in available light.Hence shade tolerance of tree species plays a vital role in forest patch development processes since light is one of the most important factors for tree growth(Zon and Graves 1911;Baker 1949).Patch size is known to positively influence the species richness of shadetolerant plants(Godefroid and Koedam 2003;Kolb and Diekmann 2005).The diversity of woody plants also increases with increasing patch size due to the dispersal mode of tree species in many tropical forests(Kolb and Diekmann 2005;Gonzalez et al.2010).For example,different responses of dispersal guilds to forest edge and forest interior habitat conditions in conjunction with increasing patch size can influence local woody species diversity(Grashof-Bokdam 1997;Hill and Curran 2003).Human disturbance can influence tree species composition in forest edge and interior zones,leading to differing light conditions that may influence forest regeneration success(Ries et al.2004;Harper et al.2005).Forest patch shape can also influence the remnant species diversity of patches,particularly at their edges(Levenson 1981;Collinge 1996),as highly dissected patches can be influenced by the confounding impacts of edge and area effects(Fletcher 2005;Laurance 2008).

        Previous studies of woody-tree species richness in the Western Ghats of India reported a significant increase with increasing patch area(Puyravaud et al.2003;Mohandass and Davidar 2010).However,the relative contributions of forest edge and interior habitats to the observed increase in species richness and abundance of woody trees in response to increasing patch size has received less attention.In addition,the role and relationship between eco-physiological guilds such as shade tolerance and seed dispersal mode,and their interactions with patch size are still unclear for montane forest patches in this region.Thus in this study we examined how these vegetation parameters respond to increasing patch size and how they interact with one another.

        We tested the following hypotheses:(1)species richness and abundance of woody plants within different stem size classes(sapling/shrubs≥1 to<10 cm dbh;small trees≥10 to<30 cm dbh;medium sized trees≥30 to<60 cm dbh;and large trees≥60 cm dbh)increase with increasing patch size;(2)species richness and abundance of woody plants in both the edge and forest interior zones increase with increasing patch size and;(3)species richness and abundance of woody plants of different shadetolerance guilds and dispersal modes in both the forest edge and interior zones respond positively to increasing patch size.

        Materials and methods

        Study area

        The study was carried out at three sites,namely Amaggal,Korakundah and Upper Bhavani,in the tropical montane evergreen forests of the upper Nilgiri Mountains,Western Ghats,south India.The region is encompassed by the Nilgiri Biosphere Reserve(NBR)(Puyravaud and Davidar 2013)and the elevation of our study sites ranged from 2000 to 2350 m above sea level(Fig.1).The study area receives the majority of its rain from two monsoon seasons,viz.southwest(SW)monsoon from June to August and northeast(NE)monsoon from October to Mid-December.Mean annual rainfall recorded from 1996 to 2004 at Korakundah Tea Estate 1 km from our study sites was 1888 mm.Mean temperature recorded from 2002 to 2006 was 20.5°C.Wind speeds can reach up to 120 m/s during the SW monsoon.The basic vegetation types of the study region are montane forest patches(locally named sholas),grasslands,and swamps(wetlands).Montane forests of the Nilgiris usually exist as small discrete patches of vegetation,although they do occasionally occur as medium and large patches.Generally,montane forest patches in the region are largely stunted(10–20 m height),evergreen,and with dense crowned canopies(Mohandass et al.2014).During our study period,most montane forest sites were surrounded by wattle plantations introduced by the forest department in 1950 for dye production;a practice which has now ceased.Adjacent to the montane patches and plantations are grasslands.Swamps occur throughout the landscape in hollows or depressions in the montane grasslands and montane forest areas.

        In the upper Nilgiris,many montane forest patches have been disturbed to varying extents by tea cultivation and exotic tree plantations(e.g.,Acacia dealbata Link.Eucalyptus globulus Labill.And Pinus patula Schiede ex Schltdl.&Cham.).The plantations have fragmented formerly contiguous forest patches and have degraded other nearby forest regions(Kumar 1993;Mohandass et al.2014,2015).In the montane forest patches,most native tree species are shade-tolerant(Meher-Homji 1984,1987),and the degree of shade tolerance appears to have a major role in determining the spatial distribution of plant species especially in relation to their distance from the forest edge and location within edge and interior habitats(Mohandass et al.2014,2016).

        Fig.1 Map of the study area in the Upper Bhavani(UB),Korakundah(KR)and Amaggal(AG),regions of the Nilgiri Mountains.Marked points show different patch sizes S small patches,M medium patches,L large patches(N=21 patches)

        Sampling methods

        Woody trees(≥1 cm dbh)in the tropical montane evergreen forest were investigated at 21 forest patches.Sampling was carried out in the Korakundah and Upper Bhavani sites of upper Nilgiris between November 2002 and September 2004,and in the Amaggal site,during December 2007–March 2008(Table 1).At each study site patches were selected randomly for sampling.To ensure sample independence,a minimum distance of 100 m was maintained between sampled patches.Patches were classified into three size categories,viz.small,medium and large.These categories were used for patch size analysis but also served as a disturbance proxy(Laurance et al.2017)as we did not examine patch disturbance per se.Small patches were approximately>0.1 to<1 ha,medium patches approximately>1 to<5 ha,and large patches were>5 ha.To standardize sample area for data analysis in the small patches,after inventory the sampled area was corrected to 30 m2(Mohandass et al.2014).In the medium and large patches,plants were sampled in plots of 30×30 m,and each plot was further subdivided into quadrats of 10×10 m(Mohandass et al.2014).Sampling of woody trees was stopped in medium and large sized patches when species area curves reached their asymptotes(Mohandass and Davidar2010;Mohandass et al.2014,2015,2016).At the Amaggal sites,due to the patch sizes,a 10×100 m transect was placed randomly and the area subdivided into quadrats of 10×10 m(Mohandass et al.2015).The total area sampled from all 21 patches was approximately 63.17 ha and there were 13 small,four medium,and four large patches(Table 1).After correction,the total sampled area was 13.21 ha,of which 4.56 ha was from the 13 small patches,4.32 ha from the four medium patches,and 4.33 ha from the four large patches(Fig.1;Table 1).Prior to data analysis,the corrected sampled area was logarithmically transformed for each forest patch from all size categories.All woody trees≥1 cm diameter at breast height(dbh)were tagged using sequentially numbered aluminum tags,identified to species level,geo-referenced to the nearest metre,and dbh of each tree was recorded at 1.3 m above ground level(Mohandass and Davidar 2010).Tree samples were identified to species level using various local and regional flora(Fyson 1932;Gamble and Fischer 1915–1935;Matthew 1999)and identification of each sample was con firmed by the Botanical Survey of India,Coimbatore(Mohandass and Davidar 2009).All voucher specimens were deposited in the herbarium of the Department of Ecology and Environmental Sciences,Pondicherry University,India.

        Woody tree species richness and abundance were recorded for edge and forest interior habitat zones.The edge zone was considered as the forested area from the periphery of the patch up to 20 m into the interior of the forest while the rest of the patch was considered as the forest interior zone.Within these zones we recorded tree community composition and selected eco-physiological traits including species shade tolerance and dispersal modes(see below for detailed description).

        Shade tolerance and dispersal mode

        Shade tolerance of woody tree species was divided into three classes.Light-Demanding(LiD)species were considered to be those abundant at the forest edge,but were also recorded at the fringes of forest patches and forest interiors due to colonization of disturbed areas.LiD species were also defined as those that germinate in open conditions or under low-stature tufted grass species.Shade-Demanding(SD)species were deemed as those found only in the forest interior zones and in shaded conditions with no canopy gaps and low light availability.Shade-Tolerant(ST)species were classified as those found in both forest edge and interior zones,but which required canopy openings to become established and needed a light signal for germination(Swaine and White more 1988;Gonzalez et al.2010;Mohandass et al.2014).

        For data analysis,we used the counts of each woody tree species with a minimum of eight occurrences from each forest patch.We then compared tree abundance between forest edge and interior zones for each species.The fruit dispersal method of each woody tree species was classified as one of two types,depending on the dominant dispersal agent.Zoochorous species were classified as those whose seeds dispersed by mammals,including Nilgiri langur(Trachypithecusjohnii),dhole(Cuonalpinus),and Sloth bear(Melursusursinus),or birds.Explosive dispersing species were categorized as those that used an explosive dehiscent method and were recorded within the dispersal category of Autochory.There were no wind or water dispersed species within the studied forest patches(Mohandass et al.2014,2016).

        Data analyses

        We grouped plants based on stem size classes:≥1 to<10 cm dbh were considered sapling/shrubs;≥10 to<30 cm dbh were considered small trees;≥30 to<60 cm dbh were considered medium sized trees;and≥60 cm dbh were considered large trees.Species richness was de fined as the number of woody tree species(Hosseinzadeh et al.2016)recorded in each patch.Abundance was classified as the number of woody trees recorded in each patch.Woody tree species richness and abundance of all dbh size classes(log-transformed)and their relationships to various vegetation parameters and patch sizes were analyzed using principal component analysis.Species diversity of edge and forest interior zones was assumed to increase linearly(when log transformed)with increasing area as argued under the theory of island biogeography(MacArthur and Wilson 1967).We also used principal component analysis to compare species richness and abundance from both the forest patch edge and interior zones with dbh class data again being log-transformed prior to analysis(Gonzalez et al.2010).We included in our analyses,only those species for which at least eight individuals(≥1 cm dbh)were recorded in sampling plots.Exotic tree species were excluded from analyses.

        Mean tree species richness and abundance between habitat zones was tested using the Mann–Whitney test.The relationship between the number of species and abundance of different eco-physiological guilds(i.e.,shade tolerance and fruit dispersal mode)among patch size and between edge and interior zones was analyzed using principal component analysis.We tested differences in mean abundance of tree species by shade-tolerance and dispersal modes between forest edge and interior zones using analysis of variance(ANOVA).We also used the Mann–Whitney test to compare tree species richness and abundance by dispersal mode(Gonzalez et al.2010;Mohandass et al.2014).We tested the correlation of species richness and abundance to patch size using principal component analysis.All statistical analyses were performed using the software SPSS version 16 and Past version.3.01.

        Results

        General patterns

        We recorded a total of 35,146 woody trees(≥1 cm dbh)representing 61 tree species,45 genera and 30 families(Table S1).Over all patch size classes,species richness per patch ranged from 20 to 45(33.9±1.5 species per patch)and tree abundance ranged from 221 to 4233(1674.2±262.8 trees per patch)(Table 1).

        Effects of patch size on tree species richness and abundance

        Total species richness and abundance of sapling/shrubs significantly and positively correlated with increasing patch size(Table 2).Species richness of small trees did not vary significantly with patch size but their abundance did.Species richness and abundance of both medium and large size trees did not significantly correlate with patch size(Table 2).

        Patch size effects by forest edge and forest interior habitat zones

        Tree species richness at forest edges did not vary significantly with patch size but tree abundance did.Both forest interior species richness and abundance significantly and positively correlated with increasing patch size(Table 2).Tree species richness in the edge zone ranged from 5 to 13 species(9.6±0.6 per patch)and tree abundance ranged from 14 to 729 trees(220.6±45.6 per patch).In the forest interior zone,richness ranged from 15 to 38 species(24.6±1.3 per patch)and abundance ranged from 120 to 3806 trees(1453±245 per patch).Of the 61 species analyzed,17 species were more frequent in the edge zone than in the interior zone and three were found only in edge zones (Rhododendron arboretum Sm.Rhodomyrtus tomentosa(Aiton)Hassk.,and Viburnum hebanthum Wight&Arn.,Table S1).Forty-six species were more common in the forest interior than at the forest edge and 59 species were recorded exclusively in the forest interior.Abundance of 13 species was significantly higher at the forest edge than in the interior.While abundance of 48 species was significantly higher in the interior than at the edge(Table S1).

        Patch size effects and tree species shade tolerance

        Shade-demanding(SD)and shade-tolerant(ST)tree species richness and abundance showed a significantly positively relationship with increasing patch size.In contrast,light-demanding(LiD)tree species richness and abundance did not vary significantly among the different sized patchclasses(Table 2).In the edge zone,LiD,SD and ST species richness and abundance were not significantly related to patch size.In the forest-interior,shade tolerant tree species richness was significantly positively related to patch size,whereas LiD and SD richness was not.Treeabundance of SD and ST species was also significantly positively related to patch size(Table 2).

        Table 2 The relationship between patch size,species richness and abundance of different stem size classes,habitat specificity,shade tolerance and dispersal mode of woody trees in the tropical montane evergreen forest patches of the Nilgiris,south India

        Tree species richness ranged between 2–8 per patch for LiD and 1–3 for SD species and 4–33 per patch for ST species.Tree abundance ranged from 3 to 234 per patch for LiD,2 to 318 per patch for SD and 31 to 877 per patch for ST abundance.More shade tolerant species were found at forest edges(84%)and interiors(87%)than both of the other listed light guilds.Shade tolerant species were significantly greater in both richness and abundance than were LiD and SD species in both edge and interior zones(Fig.2).Tree species richness of the three shade tolerance guilds(LiD,SD and ST)did not vary significantly between forest edge and interior zones(Fig.2a).The abundance of LiD species did not vary significantly in either the edge or interior zones,but the abundance of SD and ST species was significantly greater in forest interiors than in edge zones(Fig.2b).

        Patch size effects and their relationship with tree species dispersal mode

        Fig.2 Species richness and abundance of tree species by shadetolerance guild in the forest edge-zones and the forest-interior zones,a tree species richness,b tree abundance in the montane forest patches of the Nilgiri Mountains,south India.Significance levels:**p<0.001;***p<0.0001;ns not significant

        Fig.3 Species richness and abundance of tree species categorized by fruit dispersal modes in the forest edge and the forest-interior habitat zones in the montane forest patches of the Nilgiri Mountains,south India:a tree species richness,b tree abundance.Significance levels are*p<0.05;**p<0.001;ns not significant

        Overall,zoochorous tree abundance was significantly positively related to patch size but species richness was not.Neither species richness nor abundance of autochorous species was related to patch size(Table 1).Richness of autochorous and zoochorous species,and abundance of autochorous species(Fig.3b)were similar in forest edge and interior zones(Fig.3a).The abundance of zoochorous species was significantly greater in forest interior than forest edge zones(Fig.3b).Species richness and abundance of autochorous species were significantly less than for zoochorous species in both the edge and forest interior zones(Table 3).This finding indicated that zoochorous species contributed significantly more to increasing patch size than autochorous tree species.

        Higher loading values in the PC analyses were used to identify the vegetation variables that were correlated with patch size(Table 4).The first three axes explained 64.9%of the variance with the first axis explaining 49.3%,the second axis 16.7%and the third axis 9.0%.The total variance accounted for by the first six PCA axes was 78.9%.The first principal component analysis(PC1)showed the vegetation parameters which were positively correlated(in bold)with patch size axis 1 and those that were weakly and negatively related to axis 2.Some additional parameters were positively related to patch size,and correlated to axis 3(Table 4).However,within these axes,species richness and abundance of all tree dbh classes,specific habitat zones,and both shade tolerance and dispersal mode guilds were the most important variables explaining the strong positive relationship of patch size for montane forest patches.

        Table 3 Frequency of fruit dispersal modes(mean±SE)based on species richness and abundance as recorded from forest edge and forest interior zones in the Nilgiri Mountains,south India

        Discussion

        The significant positive relationship identified between sapling tree species richness and abundance and increasing forest patch area conforms to the Theory of Island Biogeography(MacArthur and Wilson 1967)and the results of several studies of species–area relationships in fragmented forests(Gonzalez et al.2010;Mohandass and Davidar 2010;Hanski et al.2013;Berhanu et al.2017).However,we did find that the richness and abundance of larger stem size classes of small trees,medium trees and large trees were not significantly related to patch size.This finding was probably due to the harvesting of larger trees by local people who use these stems for fire wood and household construction works.As a result,medium and large tree species richness and abundance might be declining in the region due to human disturbances despite an overall trend in the region of increasing average patch size(Cochrane and Schulze 1999;Chazdon 2003).

        According to our results,edge tree species richness did not increase with patch size.This again could be linked with locals harvesting trees from patches but might also be due to the prevalence of exotic trees and tea plantations which surrounded many of the montane forest patches.These exotic plantations can affect the recruitment of native tree species and compete with them for resources as they invade areas of disturbance at the forest edge(Saunders et al.1991).While,certain native pioneer species can recolonize the edges of patches planted to exotic tree and tea plantations(Selwyn and Ganesan 2006;Mohandass et al.2016)the number of species with this capacity is minimal.

        The expansion of the edge and forest interior zones of forest patches appears to be the primary driver of forest patch expansion in the region.Thus edge and forest interior zones might be important habitat to recruit species colonization,growth and survival,as well as ecological process(Laurance et al.2000).Both edge and forest interior zones might in fluence to grow larger patch size by the aid of zoochorous dispersal and shade tolerance pattern.As such,in the present study,pollination and seed dispersal of trees were often greater in edge zones than in interiors(Magrach et al.2014).In particular zoochorous dispersal influence shade tolerant and light demanding species have been shown to be more active on forest edges that could be significant factors for large patch size increases.Moreover forest edge expansion might be occurring because of the greater light availability at forest edges than in the forest interior might facilitate recruitment of pioneer and exotic species that are light demanding or shade tolerant(Baez and Balslev 2007;Dalling and Hubbell 2002).As the area of the patch increases,the perimeter to area ratio of the edge zone also increases leading to an increase in the abundance of edge habitat for woody trees(Laurance 2008;Gonzalez et al.2010).Thus,an increase in light availability and habitat area of expanding forest edges probably allows for the recruitment of more species and individuals,particularly through zoochorous dispersal from adjacent forested habitats,furthering an increase in forest patch size(De Angelis and Waterhouse 1987).

        The forest interior zone can also play an important role in increasing within-patch plant diversity(Forman and Godron 1986;Temple 1986;Li et al.1993).We found that species richness and abundance in the forest interior zones was positively related to patch size.Most of the forest interior species recorded in our study are shade-tolerant and are able to regenerate in the habitat conditions prevailing at the forest edge and in the interior.Thus,they played a major role in the enrichment of tree diversity in the forest interior.However,we did identify some interior zone specialist tree species which were intolerant of the conditions at forest edges such as Pyschotria nilgiriensis Deb&M.G.Gangop.,Lasianthus venulosus(Wight&Arn.)Wight and Saprosma fragrans(Bedd.)Bedd.Such species clearly contributed to the increase in forest interior tree diversity and thus overall patch diversity.These species were likely recruited to the forest interiors through zoochory(Majumdar et al.2016)and the tendency for birdsto perch(and defecate seeds)on the edge of canopy gaps or treefall gaps in forest interiors,thereby influencing the regeneration of shade-tolerant tree species(Echeverria et al.2007).

        Table 4 Effect of patch size on the predictive vegetation parameters of the tropical montane evergreen forest patches of the Nilgiri Mountains,south India

        Shade tolerant tree species appear to contribute the most toward increasing forest area because they can survive at the forest edge and in the interior.As mentioned above,the forest interior specialist species we identified would not be able to survive on the edges of the forests.Analogously,some of the early colonizing,light demanding species we identified might not recruit under shady conditions in the forest interior,i.e.,species such as Rhododendron arboretum Sm.and Rhodomyrtus tomentosa(Aiton)Harsk.Such species respond negatively to canopy closure due to the resulting decrease in light availability and would thus decline in forest interiors(Bin et al.2012;Mohandass and Davidar 2010).As a consequence the more utilitarian shade tolerant tree species might contribute most toward increasing forest area because they can survive under the varying habitat conditions found in both the forest edge and interior zones.If these shade tolerant species disperse between patches,as occurs through zoochory(Echeverria et al.2007)the major dispersal type identified in our study,they would likely contribute even more to tree species richness and abundance.

        Light demanding species were relatively rare on our study sites and might play a lesser role in patch expansion.We postulate that the relative rareness of LiD species may be because they are ephemeral,either dying quickly if shaded,or growing rapidly into larger dbh size classes if light levels remain high(Bin et al.2012).Rapid growth to large size classes by LiD species might explain why they did not show a significant relationship with patch size:large trees face high extraction pressure by locals for use as fire wood,particularly if accessible such as when they are found on forest edge zones or open areas.Thus,LiD species abundance at the forest edges was probably limited by timber harvest in most of the studied patches.As a consequence,this extraction of LiD species may aid in the disproportionate role shade tolerant species play in patch expansion in both the edge and forest interior zones,and thus the identified significant relationship between forest patch size and shade tolerant tree species diversity.

        While species richness by dispersal mode was unrelated to patch size,tree abundance was significantly and positively related to patch size for zoochorous species.Thus it is likely that tree species dispersal through zoochory played a role in the maintenance of diversity of woody species recorded in our forest patches,many of which are habitat generalists(Mohandass 2007).This influence occurs as zoochorous species are often dispersed over larger distances(i.e.between patches and into the interior of forest patches)in our study region and many zoochorous species are shade tolerant(Echeverria et al.2007)so their recruitment is favored in forest interiors.In order of succession,this means that the colonization of light demanding pioneer tree species dispersed via zoochory may facilitate the colonization of patches by shade tolerant species followed by shade-demanding species,particularly those dispersed by zoochory.In addition,zoochory might facilitate the transfer of propagules and genetic diversity between forests patches,and thus play a vital role in sustaining the link between edges and forest interior zones as well as vegetative diversity at the patch and landscape level(Cayuela et al.2006).

        Conclusions

        Sapling/shrubs species richness and abundance increased with increasing patch size,whereas small,medium and large tree species richness and abundance was not related to increasing patch size in our forest patches.This recruitment of small trees suggest that patches are expanding through natural successional processes such as recruitment.Forest interior zones favored the recruitment of zoochorous species due to their provision of bird perches and ease of access by other zoochorous dispersers.Edge and forest interior species richness and abundance positively related to patch size.Therefore,patch size may influence the integrity of habitat zones such as edge and interiors over time and space.Although,shade-tolerant species may predominantly dispersed by zoochorous that creates forest patches extend in size by edge zones.Besides,interior zones may provide suitable conditions for establishment of new recruitment by zoochorous dispersal for increasing the large patch size.Thus indicates shadetolerant and zoochorous dispersal pattern might significantly influence the patch size by the aid of habitat zones(edges and interiors)in the montane evergreen forests of the Nilgiris.Light demanding and shade tolerant species are probably pioneer successional species in the forest patch edge zone,whereas shade demanding species are generally late successional species which strengthen forest cover as patch size increases,especially in the forest interior.However,further study is required to understand the impact of disturbance and climate in relation with patch size effect on woody species richness and abundance,which could reveal the future insights to montane forest patches.

        AcknowledgementsWe are grateful to Dr.Jean-Philippe Puyravaud,Sigur Nature Trust who provided useful suggestions for experimental design and support for this study.We convey our deep gratitude to Prof.Dr.Qing-Jun Li and Dr.T.Muthukumar who provided logistic support and facilities for writing and revise this paper.We thank three anonymous referees especially Language Editor who greatly helped to improve the quality of manuscript and language.We thank Dr.Rama Chandra Prasad for his valuable help with map making and comments given on an earlier version of this manuscript.We thank Dr.Christos Mammides and Dr.V.S.Ramachandran for their valuable comments and language editing in earlier versions.We thank Mr.Hegde and staff of UNITEA Pvt.Ltd,the Tamil Nadu Forest Department and Electricity Board for logistic help and support.This publication was supported by the National Natural Science Foundation of China(NSFC)through Young Scientist Grant No.31200173,P.R.China.This study was partially supported by a small grant from the Center for Tropical Forest Science,Smithsonian Tropical Research Institute and assistance from Dr.Egbert G.Leigh Jr.

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