11
Research Article Trends in Tree Diversity and Stand Structure during Restoration: A Case Study in Fragmented Moist Deciduous Forest Ecosystems of Northeast India Koushik Majumdar, 1 Uma Shankar, 2 and Badal Kumar Datta 1 1 Plant Taxonomy and Biodiversity Laboratory, Department of Botany, Tripura University, Suryamaninagar, Tripura 799022, India 2 Ecosystem Research Laboratory, Centre for Advanced Studies in Botany, School of Life Science, North-Eastern Hill University, Shillong, Meghalaya 793022, India Correspondence should be addressed to Koushik Majumdar; [email protected] Received 19 December 2013; Accepted 29 May 2014; Published 14 July 2014 Academic Editor: Francisco Arreguin-S´ anchez Copyright © 2014 Koushik Majumdar et al. is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Ecological implications for the conservation of fragmented forests to managed local plant diversity have recently drawn much attention of most conservationists. Present study investigates the importance of fragmented forests using different quantitative measures of species diversity and stand structure as indicators of natural restoration from past disturbances. Eleven independent 500 m × 10 m belt transects (5.5 ha) were established within fragmented low land moist deciduous forests of Tripura. All woody plants 10 cm girth at breast height (GBH) were measured. A total of 7,134 individuals with mean density of 648.55 stems ha −1 and 16.36 m 2 ha −1 of basal area were recorded, which represented 134 species, 93 genera, and 43 families of woody plants. Diversity- dominance curve showed that maximum number of species were ranked >10 due to low abundance value. Maximum distributions of stems (>50%) were recorded at lowest girth class (10–30 cm) as an indication of advanced regeneration and significantly declined towards upper girth ( 2 adj = 0.93; < 0.0001) and height ( 2 adj = 0.95; < 0.0001) classes, which also indicated that some plants were adopted with ongoing disturbances through both seeded and nonseeded regeneration. Species richness ( 2 adj = 0.49; = 0.05), dominance ( 2 adj = 0.68; = 0.009), and density ( 2 adj = 0.62; = 0.02) were significantly increased at different disturbance intensities. Disturbances strongly influenced typical community association and structure by increasing diversity and population at certain magnitude and thereby showed declining trend towards maturation. Fragmented moist deciduous forests seek immediate attention as they represent spatial habitat for many economical or ecological important species, thus sustaining local biodiversity for livelihoods. 1. Introduction Small forest fragments or areas lying outside fully protected zones maintain a significant number of species and their genetic diversity [1, 2]. Even forest patches of less than 1 ha in size could play major roles in maintenance of regional bio- diversity [3]. During the past several years, large complexes of natural habitat have been converted into agricultural, industrial, or urbanized landscapes, leading to severe loss of the original habitat and an increasing fragmentation of the remnant patches [4]. High fragmentation rate stimulate habitat modifications and the loss of valuable portions of the forest ecosystem, the number of rare and shade – tolerant species increases due to fragmentation especially during recovery at the early successional stage [5]. Tree species with small population will be the first to be lost in the process of forest fragmentation [6]. Tree species diversity is an important and fundamental aspect to total forest ecosystem biodiversity [7, 8]; they provide resource and habitat for almost all other species [9]. Tree diversity in tropical forests varies greatly from place to place, mainly due to variation in biogeography, climatic responses, habitat suitability, and Hindawi Publishing Corporation Journal of Ecosystems Volume 2014, Article ID 845142, 10 pages http://dx.doi.org/10.1155/2014/845142

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Page 1: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

Research ArticleTrends in Tree Diversity and Stand Structure duringRestoration A Case Study in Fragmented Moist DeciduousForest Ecosystems of Northeast India

Koushik Majumdar1 Uma Shankar2 and Badal Kumar Datta1

1 Plant Taxonomy and Biodiversity Laboratory Department of Botany Tripura University SuryamaninagarTripura 799022 India

2 Ecosystem Research Laboratory Centre for Advanced Studies in Botany School of Life ScienceNorth-Eastern Hill University Shillong Meghalaya 793022 India

Correspondence should be addressed to Koushik Majumdar majumdark80gmailcom

Received 19 December 2013 Accepted 29 May 2014 Published 14 July 2014

Academic Editor Francisco Arreguin-Sanchez

Copyright copy 2014 Koushik Majumdar et al This is an open access article distributed under the Creative Commons AttributionLicense which permits unrestricted use distribution and reproduction in any medium provided the original work is properlycited

Ecological implications for the conservation of fragmented forests to managed local plant diversity have recently drawn muchattention of most conservationists Present study investigates the importance of fragmented forests using different quantitativemeasures of species diversity and stand structure as indicators of natural restoration from past disturbances Eleven independent500m times 10m belt transects (55 ha) were established within fragmented low land moist deciduous forests of Tripura All woodyplants ge10 cm girth at breast height (GBH) were measured A total of 7134 individuals with mean density of 64855 stems haminus1 and1636m2 haminus1 of basal area were recorded which represented 134 species 93 genera and 43 families of woody plants Diversity-dominance curve showed that maximum number of species were ranked gt10 due to low abundance value Maximum distributionsof stems (gt50) were recorded at lowest girth class (10ndash30 cm) as an indication of advanced regeneration and significantly declinedtowards upper girth (1199032adj = 093 119875 lt 00001) and height (1199032adj = 095 119875 lt 00001) classes which also indicated that some plantswere adopted with ongoing disturbances through both seeded and nonseeded regeneration Species richness (1199032adj = 049 119875 = 005)dominance (1199032adj = 068 119875 = 0009) and density (1199032adj = 062 119875 = 002) were significantly increased at different disturbanceintensities Disturbances strongly influenced typical community association and structure by increasing diversity and populationat certainmagnitude and thereby showed declining trend towardsmaturation Fragmentedmoist deciduous forests seek immediateattention as they represent spatial habitat for many economical or ecological important species thus sustaining local biodiversityfor livelihoods

1 Introduction

Small forest fragments or areas lying outside fully protectedzones maintain a significant number of species and theirgenetic diversity [1 2] Even forest patches of less than 1 hain size could play major roles in maintenance of regional bio-diversity [3] During the past several years large complexesof natural habitat have been converted into agriculturalindustrial or urbanized landscapes leading to severe lossof the original habitat and an increasing fragmentation ofthe remnant patches [4] High fragmentation rate stimulate

habitat modifications and the loss of valuable portions of theforest ecosystem the number of rare and shade ndash tolerantspecies increases due to fragmentation especially duringrecovery at the early successional stage [5] Tree specieswith small population will be the first to be lost in theprocess of forest fragmentation [6] Tree species diversity is animportant and fundamental aspect to total forest ecosystembiodiversity [7 8] they provide resource and habitat foralmost all other species [9] Tree diversity in tropical forestsvaries greatly from place to place mainly due to variationin biogeography climatic responses habitat suitability and

Hindawi Publishing CorporationJournal of EcosystemsVolume 2014 Article ID 845142 10 pageshttpdxdoiorg1011552014845142

2 Journal of Ecosystems

anthropogenic disturbances [10 11] Disturbances stronglyinfluence forest dynamics community attributes structureand diversity at both local and regional scales [12 13] evidentin changing size class distributions of the most sought-afterspecies [14]

Most of the quantitative vegetation inventories wereconducted in species-rich forests and data on species-poorforests or forest fragments are inadequate [15] In addi-tion species richness and abundance data are crucial forassessing the impacts of disturbance or fragmentation onlocal biodiversity conservation priority setting and long-term vegetation monitoring [16] Moist deciduous forest isone of the crucial habitat types of India and most of theearlier studies were reported from Western Ghats EasternGhats and Andaman Island [17ndash20] Data on communitycomposition and population structure in lowland moistdeciduous forests are limited Since remaining fragmentedforests are quickly shrinking or modifying under severalmanmade influences immediate attention was felt necessaryto gather quantitative inventories for tree diversity and standstructure in fragmented lowland forests to better understandits nature and dynamics Hence present objectives weredesigned to investigate (1) the status of woody plant diversityand structure in fragmented forests (2) to draw the structuralchanges under present level of disturbance intensity and (3)to test the hypothesis that species diversity stand populationand dominance increased during restoration following inter-mediate disturbance hypothesis of Connell (1978) [21]

2 Materials and Methods

21 Study Site Present study was conducted in Tripurawhich is a small hilly state belonging to the northeastern partof IndiaThe state is surrounded by Bangladesh on three sidesand only northeast part of the state is connected throughAssam and Mizoram with the country accounting for barely0342 of the total area of the country In terms of areait is the third smallest state in the country after Goa andSikkim The geographical area of the state is only 10492 km2and has 4 districts The state is located in biogeographic zoneof 9B-North-East hills endowed with rich and diverse forestresources About 5998 of the geographical area of the stateis under the forest The forests are mainly semievergreenmoist deciduous mixed sal and dry deciduous forests Thepresent study sites located between 23∘5710158405510158401015840 and 24∘0510158403410158401015840north latitude and 91∘1710158401810158401015840 and 91∘2910158402210158401015840 east longitudeThearea is bordered by Bangladesh on northwest side and locatedin extreme northwest part of state The study area falls underlow physiographic zone slope is lt 40∘ and the altitude islt100m (ranging from 1859 to 9022m amsl) The climate ofthe area is characterized by highly humid atmosphere andtemperatures range from 10∘C to 35∘C The monsoon seasonis mainly from June to September with an annual averagerainfall around 2400mmmostly due to southwest monsoonForest characteristic of the study area is similar to moistdeciduous forest (II

3

CC3

b) Total forest area recorded forWest Tripura is 1155409 km2 out of 6292681 km2 Actualstudy locations belong to Mohanpur and Hezamara block

Figure 1 Map of the study area showing eleven different sites infragmented forests of West Tripura India

under Sadar forest division of West Tripura (Figure 1) Northpart of this area particularly the Hezamara block inhabitedby Tripura and Reang tribe many of them still practicingtraditional shifting cultivation Hence both the marginal andinterior forest areas are under threat of sever degradationLarge forest areas have been converted from time to timeinto tea plantation especially inMohanpur block and only fewforest patches still exist [22]

22 Field Sampling Eleven 05 ha belt transects (500m times10m) were established in the inventory of woody vegetationof fragmentedmoist deciduous forest ofWest Tripura districtduring April 2011 and thus the total sampling area was 55 ha(Table 1(A)) Plots were laid randomly within remainingfragmented forests surrounding village boundary plantationand roads In each plot all woody plantsge10 cmgirth at breastheight (GBH) were measured in centimeter and countedindividually at 13m height from the ground level using atape height for all individuals was measured to the nearestmeter [23] We restricted our analysis and interpretation ofdiversity and community composition to species that hadsuccessfully established a minimum girth class We alsoincluded those woody shrubs bamboos and lianas thatattained our girth category The GPS locations of the plotsand physical condition of every individual were noted underalive cut and dead categories sites disturbance scores werealso recorded to obtain quantitative disturbance intensity[24 25] Specimens were identified with the help of theFlora of Tripura State and the Flora of Assam [26 27] Thereference herbarium was deposited in herbarium of BotanyDepartment Tripura University

23 Data Analysis The field data was analyzed for numberof species and quantitative analysis of frequency densityper ha and basal area per ha and their relative values werecalculated and summed to get Importance Value Index (IVI)using Mueller-Dombois and Ellenberg [28] Relative density= (Density of the speciesTotal density of all species) times 100

Journal of Ecosystems 3

Table1Sitecharacteris

ticsa

nddistu

rbance

intensity

with

inthefragm

entedforests

Fragmentedforestsites

AL

ocations

BSitedistu

rbanceslowast

Latitud

e(ddm

mss)

Long

itude

(ddm

mss)

Altitude

(m)

Fire

Grazing

Fuelwoo

dcollection

Thatch

collection

Soil

removal

NTF

Pcollection

Num

bero

fcuttrees

Num

bero

fdead

trees

Total

score

Disturbance

intensity

()

Simna

(Sim

)24∘

051015840

349810158401015840

91∘

261015840

101410158401015840

6857

22

31

12

202

3392

4Diggyatila

(Dig)

24∘

051015840

153610158401015840

91∘

261015840

396610158401015840

6736

13

21

11

368

531485

Taranagar(Tar)

24∘

051015840

141110158401015840

91∘

241015840

301010158401015840

2895

13

33

11

122

2672

8Ka

tlamara(

Kat)

24∘

041015840

132010158401015840

91∘

231015840

197010158401015840

2956

12

22

22

100

21588

Subalsh

ing(Sub

)24∘

001015840

160010158401015840

91∘

271015840

375010158401015840

7559

21

31

22

366

531485

Mon

tala(M

on)

24∘

011015840

104010158401015840

91∘

261015840

121010158401015840

5577

23

13

11

40

15420

Bijoyn

agar

(Bij)

24∘

001015840

238010158401015840

91∘

231015840

390010158401015840

3078

12

22

11

160

2570

0Gajapara(

Gaj)

23∘

551015840

450010158401015840

91∘

291015840

228010158401015840

9022

22

22

21

264

411148

Borkathal(Bo

r)23∘

571015840

550010158401015840

91∘

241015840

040010158401015840

3596

13

33

22

222

381064

Lonk

amura(

Lon)

23∘

521015840

325010158401015840

91∘

141015840

543010158401015840

1859

13

23

21

320

441232

Salbagan

(Sal)

23∘

531015840

018010158401015840

91∘

171015840

182010158401015840

3535

12

11

21

00

8224

Total

1526

2422

1715

214

24357

100

Mean

133

233

211

222

167

133

1756

156

3011

843

Standard

error

017

024

026

028

017

017

412

073

492

138

Standard

deviation

050

071

078

083

050

050

1236

219

1475

413

lowast

Scoringrare1occasional2andrecurrent3and1score

fore

very

cutand

dead

stem

4 Journal of Ecosystems

Relative frequency = (Frequency of the speciesTotal fre-quency of all species) times 100 Relative dominance = (Basal areaof the speciesTotal basal area for all species)times 100 ImportantValue Index (IVI) = Relative Frequency + Relative Density+ Relative Basal Area General diversity was calculated usingShannon andWiener index [29] and the index of dominanceof the community was calculated by Simpsonrsquos index [30]Shannon and Wienerrsquos index = minussum(119899119894119873)log

2

(119899119894119873) ni =IVI of the species i N = IVI of all species in the communitySimpson dominance index = sum(119899119894119873)2 ni = IVI of speciesi N = IVI of all species in the community Species richnessindex was computed using Menhinick [31] and Evennessindex of the forest community was calculated followingPielou [32] Menhinick richness index = SradicN S = totalnumber of species N = total number of individuals Pielouevenness index = Shannon and Wienerrsquos indexlog(S) S =total number of species

The number of species was plotted on 119910-axis against thenumber of individuals on the 119909-axis for comparing speciesrichness among the fragmented forests [33] and speciesabundance data was plotted against species rank to developdiversity versus dominance curves [34] using BioDiversityPro version 2 [35] Woody species was grouped among tengirth classes (ge10ndashlt30 30ndashlt60 60ndashlt90 90ndashlt120 120ndashlt150150ndashlt180 180ndashlt210 210ndashlt240 240ndashlt270 and gt270 cm) andheight classes (lt3 3ndashlt6 6ndashlt9 9ndashlt12 12ndashlt15 15ndashlt18 18ndashlt2121ndashlt23 24ndashlt26 and gt26m) to observe the distribution ofpopulation at different age and height classes To examine thepattern of changes in community types and stand structureacross the studied sites regression curve fitting was used withmidvalue of overall stem density haminus1 girth and height classeson 119909-axis and log transformed density haminus1 on 119910-axis [36]In addition to test intermediate disturbance hypothesis weexamined the relationship by curve fitting between densitydiversity and dominance on 119910-axis and disturbance intensityon119909-axis for those fragmented forests usingOrigin and PASTsoftware [37 38]

3 Results

31 Status of Species Richness and Habitat Diversity AttributesA total of 134 woody species (ge10 cmGBH) representing93 genera and 43 families were recorded within 55 hafragmented moist deciduous forest of West Tripura Habit-wise out of 134woody species 118 species (8806)were trees7 (522) species were woody liana 4 (299) species werewoody shrub and 5 (373) species were bamboo Speciesrichness ranged from 31 to 61 species in eleven sites BothDig and Gaj had 61 species which was two times greater thanSal (31) The observed family richness was the greatest in Gaj(37)The rarefaction curves confirmed that sites Gaj SimDigand Kat were steeper than the others suggesting high speciesrichness (Figure 2)Whereas Simdid not asymptote at similarnumbers of individual which revealed that sufficient numberof species might not covered by our sampling effort and therewas possibility to get additional species in Sim compared toother sites Gaj scored high value of Shannon diversity index(392) while Sal recorded low value (220) compared to the

SimDigTarKatSubMon

BijGajBorLonSal

Expe

cted

num

ber o

f sp

ecie

s (S)

Number of individuals (N)

08

162432404856647280

0 50 100 150 200 250 300 350 400 450 500

Figure 2 Rarefaction curves for comparing the species richness ineleven fragmented forest stands

rest of the sites Mean Simpson dominance index was 006and was recorded high in Sub (007) which was recordedlow in Gaj (002) Species were more evenly distributed inSim Tar Gaj and Mon with a mean value of Evenness indexof 194 (range 147ndash216) while Menhinikrsquos species richnessvaried from 131 to 264 along the fragment sites with meanvalue of 185 (Table 2)

The diversity-dominance curve displayed a log seriesdistribution where species ranked (1ndash100) from most to leastabundant and majority of the species had lower abundanceand population while few species showed higher valuesTectona grandis appeared to be the most abundant in SimSub andMon Schimawallichii inDig Kat andGaj Syzygiumcerasoides in Bij and Bor and Microcos paniculata Gmelinaarborea and Shorea robusta in Tar Lon and Sal respectivelyHolarrhena antidysenterica Microcos paniculata Terminaliabellirica Glochidion multiloculare Chaetocarpus castanicar-pus Syzygium cerasoides and Ziziphus rugosa were observedas codominant association although Aporosa obovata Can-thium glabrum Desmos dumosus Elaeocarpus prunifoliaElaeocarpus robustus Bhesa robusta Leea guineensis Litseacubeba and Litsea laeta were recorded with very low abun-dance and population (Figure 3)

32 Patterns of Stand Density Basal Area and Impor-tant Value Index (IVI) A total of 7134 individuals wereencountered with mean density of 64855 trees haminus1(rangedfrom 428 to 884 haminus1) in eleven 05 ha of fragmentedforest sites The highest woody density was recorded forBor (884 haminus1) and lowest for Sal (428 haminus1) Top tendominant trees namely Microcos paniculata (4382 haminus1)Tectona grandis (4309 haminus1) Holarrhena antidysenterica(3036 haminus1) Syzygium cerasoides (2764 haminus1) Shorea robusta(2491 haminus1) Schima wallichii (2345 haminus1) Syzygium frutico-sum (2145 haminus1) Melocanna baccifera (1782 haminus1) Gmelinaarborea (1745 haminus1) and Terminalia bellirica (1655 haminus1)

Journal of Ecosystems 5

Table 2 Consolidated details of woody plant diversity inventory (ge10 cm GBH) in eleven fragmented moist deciduous forests of Tripura

Parameters Fragmented forest sites Mean Standard error Standard deviationSim Dig Tar Kat Sub Mon Bij Gaj Bor Lon Sal

Number of woody species 60 61 41 45 45 40 37 61 39 47 31 4609 311 1032Number of genera 50 54 31 42 39 37 29 54 33 41 31 4009 276 916Number of families 29 32 22 26 32 28 23 37 25 28 25 2791 134 444Shannon index 373 357 333 349 332 347 275 392 258 338 22 325 016 052Simpson index 003 004 005 004 007 004 002 002 002 006 03 006 002 008Evenness index 21 2 206 21 201 216 164 22 162 201 147 194 007 025Richness index 264 208 164 22 163 17 142 247 131 171 15 185 013 044Density (haminus1) 518 854 628 436 762 552 680 608 884 784 428 64855 4787 15878Basal area (m2haminus1) 528 1408 68 145 1255 9 1644 236 1404 166 4708 1636 343 1137

Spec

ies a

bund

ance

Species rank

0102030405060708090

100110120130140150

1 10010

SimDigTarKatSubMon

BijGajBorLonSal

Figure 3 Diversity-dominance curve and species ranking based onspecies abundance

contributed in 4110 of overall density Mean basal areawas 1636m2 haminus1 for the whole study of which top tendominant trees grasped 944m2 haminus1 (5768) of total basalarea and it recorded the minimum for Sim (528m2 haminus1)and was the greatest in case of Sal (4708m2 haminus1) Basedon the IVI top ten dominant and codominant trees con-tributed in 11011 of total IVI (300) Shorea robusta graspsthe maximum IVI (3053) which was predominant only inSal followed by Tectona grandis (1170) Microcos paniculata(1122) Schima wallichii (1020) Syzygium cerasoides (980)Holarrhena antidysenterica (815) Artocarpus heterophyllus(754) and Terminalia bellirica (712) While 18 species sowedIVI value lt030 Desmos dumosus Leea aequata Elaeocarpusrobustus and Actinodaphne obovata were recorded withvery low IVI contribution to the fragmented forest Overall27 species were selected (top five most dominant speciesfrom each site) based on their IVI contribution to thosefragmented forests which suggested that Microcos panicu-lata Holarrhena antidysentericaChaetocarpus castanicarpusGlochidion multilocure Syzygium cerasoides and Ziziphusrugosawere increasing their IVI over previously predominant

trees These small tress species showed very little differencesin terms of IVI contribution with key dominant speciesnamely Shorea robusta SchimawallichiiTectona grandis andGmelina arborea (Figure 4)

33 Changes in Girth or Age-Class and Height-Class Distribu-tions Stem density across girth classes in all sites decreasesalmost from smallest to largest trees While occurrencesof mean stems haminus1 at lower girth class (10ndashlt30 cm) were5088 (330 plusmn 954) followed by 2910 (18873 plusmn 9694)in 30ndashlt60 cm girth class and 1096 (7109 plusmn 3798) in 60ndashlt90 cm girth class only 006 stems were shown in thehighest (240ndashlt300 cm) girth classes Maximum number ofindividuals at lower girth classwas recorded for Bor (468) andminimum for both Kat and Gaj (212) While few individualswere recorded at higher girth class (gt270 cm) some highergirth classes were absent in Sim Tar Sub Mon Bor andLon Stems were more frequently distributed in all the elevensites within the 10ndashlt120 cm girth classes The overall stemdensity haminus1 at different age classes was significant (1199032adj =093 119875 lt 00001) among the studied sites and showedinverse quadratic relationship (Figure 5) The distributionof stems by height class intervals showed that maximumtrees (8247) were widely distributed within 3ndash12m range3146 of individuals weregt3m followed by 2363 between3 and lt6m and 1435 within 6ndashlt9m whereas 720 ofindividuals distributed between 12 and lt15m only 042individuals were attained gt26m Density of trees was linearlydeclined across the height classes (1199032adj = 095 119875 lt 00001)among the fragmented sites (Figure 6)

34 Effects of Disturbances and Recovery Processes Besidesserious past logging experience for timber several otheranthropogenic disturbances were still ongoing in those frag-mented forests sites Grazing and fuelwood collection weremore intense due to closeness of village A total of 214trees haminus1 (1945 plusmn 1236) were recorded as cut or loggedand 24 trees haminus1 (218 plusmn 275) as injured out of overall7134 individuals maximum number of cut and injured treeswere recorded both from Dig and Sub Both cut and injuredtrees were 238 (2164 plusmn 1450) for overall study but wereabsent in case of Sal Site disturbance intensity was obtained

6 Journal of Ecosystems

minus16

minus8

0

8

16

24

32

40

48M

ean

and

stan

dard

err

or o

f |V

|

Albizia

p

Albizia

s

Artocarpu

Bauhinia

Careyaar

Chaetocar

Ficusrel

Garcinia

Glochidio

Gmelinaa

Holarrhen

Lannea

co

Litseamo

Mallotus

Mang

ifera

Microcos

Schim

awa

Shorea

ro

Streblus

Syzygium

Syzygium

Tamarindu

Tectonag

Terminali

Tremaori

Zanthoxyl

Ziziphus

Figure 4 Dominance and codominance competition of 27 tree species (mean and standard error of IVI) Five most dominant species wereselected from each fragmented forest site based on highest IVI contribution to the community

0 20 40 60 80 100120140160180200220240260280300minus02

00020406081012141618202224262830

Midvalue value of girth class (cm)

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 093 P lt 00001

Log

of d

ensit

y (h

a)

minus1

Figure 5 Relationship between population density (log trans-formed) and midvalue of girth class intervals across the fragmentedforest sites

qualitatively by summing the disturbance scores along andthe number of cut and injured individuals (Table 1(B)) Itcould be assumed that the proportion of scores that showedmaximum value revealed comparatively high level of anthro-pogenic disturbance or fragmentation than the lower valueThe highest intensity was recorded for Sub and Dig (1485)followed by Lon (1232) Gaj (1148) and Bor (1046)and the lowest in case of Sal (224) While the number

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30minus02

00020406081012141618202224262830

Midvalue value of height class (m)

Log

of d

ensit

y (h

a)

minus1

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 095 P lt 00001

Figure 6 Relationship between population density (log trans-formed) and midvalue of height class intervals across the frag-mented forest sites

of species (that occurred on each site) was significantlyincreased with disturbance intensity (1205942 = 006 F = 392119875 = 005) at 15 level and declining trend was observedwhen intensity level was increased among the sites (Figure 7)initially declining trend of the dominance of key communityspecies was observed and at midlevel of disturbance (10)few fragmentation tolerant species significantly (1205942 = 036 F= 872 119875 = 0009) increased in dominance with disturbance

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 2: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

2 Journal of Ecosystems

anthropogenic disturbances [10 11] Disturbances stronglyinfluence forest dynamics community attributes structureand diversity at both local and regional scales [12 13] evidentin changing size class distributions of the most sought-afterspecies [14]

Most of the quantitative vegetation inventories wereconducted in species-rich forests and data on species-poorforests or forest fragments are inadequate [15] In addi-tion species richness and abundance data are crucial forassessing the impacts of disturbance or fragmentation onlocal biodiversity conservation priority setting and long-term vegetation monitoring [16] Moist deciduous forest isone of the crucial habitat types of India and most of theearlier studies were reported from Western Ghats EasternGhats and Andaman Island [17ndash20] Data on communitycomposition and population structure in lowland moistdeciduous forests are limited Since remaining fragmentedforests are quickly shrinking or modifying under severalmanmade influences immediate attention was felt necessaryto gather quantitative inventories for tree diversity and standstructure in fragmented lowland forests to better understandits nature and dynamics Hence present objectives weredesigned to investigate (1) the status of woody plant diversityand structure in fragmented forests (2) to draw the structuralchanges under present level of disturbance intensity and (3)to test the hypothesis that species diversity stand populationand dominance increased during restoration following inter-mediate disturbance hypothesis of Connell (1978) [21]

2 Materials and Methods

21 Study Site Present study was conducted in Tripurawhich is a small hilly state belonging to the northeastern partof IndiaThe state is surrounded by Bangladesh on three sidesand only northeast part of the state is connected throughAssam and Mizoram with the country accounting for barely0342 of the total area of the country In terms of areait is the third smallest state in the country after Goa andSikkim The geographical area of the state is only 10492 km2and has 4 districts The state is located in biogeographic zoneof 9B-North-East hills endowed with rich and diverse forestresources About 5998 of the geographical area of the stateis under the forest The forests are mainly semievergreenmoist deciduous mixed sal and dry deciduous forests Thepresent study sites located between 23∘5710158405510158401015840 and 24∘0510158403410158401015840north latitude and 91∘1710158401810158401015840 and 91∘2910158402210158401015840 east longitudeThearea is bordered by Bangladesh on northwest side and locatedin extreme northwest part of state The study area falls underlow physiographic zone slope is lt 40∘ and the altitude islt100m (ranging from 1859 to 9022m amsl) The climate ofthe area is characterized by highly humid atmosphere andtemperatures range from 10∘C to 35∘C The monsoon seasonis mainly from June to September with an annual averagerainfall around 2400mmmostly due to southwest monsoonForest characteristic of the study area is similar to moistdeciduous forest (II

3

CC3

b) Total forest area recorded forWest Tripura is 1155409 km2 out of 6292681 km2 Actualstudy locations belong to Mohanpur and Hezamara block

Figure 1 Map of the study area showing eleven different sites infragmented forests of West Tripura India

under Sadar forest division of West Tripura (Figure 1) Northpart of this area particularly the Hezamara block inhabitedby Tripura and Reang tribe many of them still practicingtraditional shifting cultivation Hence both the marginal andinterior forest areas are under threat of sever degradationLarge forest areas have been converted from time to timeinto tea plantation especially inMohanpur block and only fewforest patches still exist [22]

22 Field Sampling Eleven 05 ha belt transects (500m times10m) were established in the inventory of woody vegetationof fragmentedmoist deciduous forest ofWest Tripura districtduring April 2011 and thus the total sampling area was 55 ha(Table 1(A)) Plots were laid randomly within remainingfragmented forests surrounding village boundary plantationand roads In each plot all woody plantsge10 cmgirth at breastheight (GBH) were measured in centimeter and countedindividually at 13m height from the ground level using atape height for all individuals was measured to the nearestmeter [23] We restricted our analysis and interpretation ofdiversity and community composition to species that hadsuccessfully established a minimum girth class We alsoincluded those woody shrubs bamboos and lianas thatattained our girth category The GPS locations of the plotsand physical condition of every individual were noted underalive cut and dead categories sites disturbance scores werealso recorded to obtain quantitative disturbance intensity[24 25] Specimens were identified with the help of theFlora of Tripura State and the Flora of Assam [26 27] Thereference herbarium was deposited in herbarium of BotanyDepartment Tripura University

23 Data Analysis The field data was analyzed for numberof species and quantitative analysis of frequency densityper ha and basal area per ha and their relative values werecalculated and summed to get Importance Value Index (IVI)using Mueller-Dombois and Ellenberg [28] Relative density= (Density of the speciesTotal density of all species) times 100

Journal of Ecosystems 3

Table1Sitecharacteris

ticsa

nddistu

rbance

intensity

with

inthefragm

entedforests

Fragmentedforestsites

AL

ocations

BSitedistu

rbanceslowast

Latitud

e(ddm

mss)

Long

itude

(ddm

mss)

Altitude

(m)

Fire

Grazing

Fuelwoo

dcollection

Thatch

collection

Soil

removal

NTF

Pcollection

Num

bero

fcuttrees

Num

bero

fdead

trees

Total

score

Disturbance

intensity

()

Simna

(Sim

)24∘

051015840

349810158401015840

91∘

261015840

101410158401015840

6857

22

31

12

202

3392

4Diggyatila

(Dig)

24∘

051015840

153610158401015840

91∘

261015840

396610158401015840

6736

13

21

11

368

531485

Taranagar(Tar)

24∘

051015840

141110158401015840

91∘

241015840

301010158401015840

2895

13

33

11

122

2672

8Ka

tlamara(

Kat)

24∘

041015840

132010158401015840

91∘

231015840

197010158401015840

2956

12

22

22

100

21588

Subalsh

ing(Sub

)24∘

001015840

160010158401015840

91∘

271015840

375010158401015840

7559

21

31

22

366

531485

Mon

tala(M

on)

24∘

011015840

104010158401015840

91∘

261015840

121010158401015840

5577

23

13

11

40

15420

Bijoyn

agar

(Bij)

24∘

001015840

238010158401015840

91∘

231015840

390010158401015840

3078

12

22

11

160

2570

0Gajapara(

Gaj)

23∘

551015840

450010158401015840

91∘

291015840

228010158401015840

9022

22

22

21

264

411148

Borkathal(Bo

r)23∘

571015840

550010158401015840

91∘

241015840

040010158401015840

3596

13

33

22

222

381064

Lonk

amura(

Lon)

23∘

521015840

325010158401015840

91∘

141015840

543010158401015840

1859

13

23

21

320

441232

Salbagan

(Sal)

23∘

531015840

018010158401015840

91∘

171015840

182010158401015840

3535

12

11

21

00

8224

Total

1526

2422

1715

214

24357

100

Mean

133

233

211

222

167

133

1756

156

3011

843

Standard

error

017

024

026

028

017

017

412

073

492

138

Standard

deviation

050

071

078

083

050

050

1236

219

1475

413

lowast

Scoringrare1occasional2andrecurrent3and1score

fore

very

cutand

dead

stem

4 Journal of Ecosystems

Relative frequency = (Frequency of the speciesTotal fre-quency of all species) times 100 Relative dominance = (Basal areaof the speciesTotal basal area for all species)times 100 ImportantValue Index (IVI) = Relative Frequency + Relative Density+ Relative Basal Area General diversity was calculated usingShannon andWiener index [29] and the index of dominanceof the community was calculated by Simpsonrsquos index [30]Shannon and Wienerrsquos index = minussum(119899119894119873)log

2

(119899119894119873) ni =IVI of the species i N = IVI of all species in the communitySimpson dominance index = sum(119899119894119873)2 ni = IVI of speciesi N = IVI of all species in the community Species richnessindex was computed using Menhinick [31] and Evennessindex of the forest community was calculated followingPielou [32] Menhinick richness index = SradicN S = totalnumber of species N = total number of individuals Pielouevenness index = Shannon and Wienerrsquos indexlog(S) S =total number of species

The number of species was plotted on 119910-axis against thenumber of individuals on the 119909-axis for comparing speciesrichness among the fragmented forests [33] and speciesabundance data was plotted against species rank to developdiversity versus dominance curves [34] using BioDiversityPro version 2 [35] Woody species was grouped among tengirth classes (ge10ndashlt30 30ndashlt60 60ndashlt90 90ndashlt120 120ndashlt150150ndashlt180 180ndashlt210 210ndashlt240 240ndashlt270 and gt270 cm) andheight classes (lt3 3ndashlt6 6ndashlt9 9ndashlt12 12ndashlt15 15ndashlt18 18ndashlt2121ndashlt23 24ndashlt26 and gt26m) to observe the distribution ofpopulation at different age and height classes To examine thepattern of changes in community types and stand structureacross the studied sites regression curve fitting was used withmidvalue of overall stem density haminus1 girth and height classeson 119909-axis and log transformed density haminus1 on 119910-axis [36]In addition to test intermediate disturbance hypothesis weexamined the relationship by curve fitting between densitydiversity and dominance on 119910-axis and disturbance intensityon119909-axis for those fragmented forests usingOrigin and PASTsoftware [37 38]

3 Results

31 Status of Species Richness and Habitat Diversity AttributesA total of 134 woody species (ge10 cmGBH) representing93 genera and 43 families were recorded within 55 hafragmented moist deciduous forest of West Tripura Habit-wise out of 134woody species 118 species (8806)were trees7 (522) species were woody liana 4 (299) species werewoody shrub and 5 (373) species were bamboo Speciesrichness ranged from 31 to 61 species in eleven sites BothDig and Gaj had 61 species which was two times greater thanSal (31) The observed family richness was the greatest in Gaj(37)The rarefaction curves confirmed that sites Gaj SimDigand Kat were steeper than the others suggesting high speciesrichness (Figure 2)Whereas Simdid not asymptote at similarnumbers of individual which revealed that sufficient numberof species might not covered by our sampling effort and therewas possibility to get additional species in Sim compared toother sites Gaj scored high value of Shannon diversity index(392) while Sal recorded low value (220) compared to the

SimDigTarKatSubMon

BijGajBorLonSal

Expe

cted

num

ber o

f sp

ecie

s (S)

Number of individuals (N)

08

162432404856647280

0 50 100 150 200 250 300 350 400 450 500

Figure 2 Rarefaction curves for comparing the species richness ineleven fragmented forest stands

rest of the sites Mean Simpson dominance index was 006and was recorded high in Sub (007) which was recordedlow in Gaj (002) Species were more evenly distributed inSim Tar Gaj and Mon with a mean value of Evenness indexof 194 (range 147ndash216) while Menhinikrsquos species richnessvaried from 131 to 264 along the fragment sites with meanvalue of 185 (Table 2)

The diversity-dominance curve displayed a log seriesdistribution where species ranked (1ndash100) from most to leastabundant and majority of the species had lower abundanceand population while few species showed higher valuesTectona grandis appeared to be the most abundant in SimSub andMon Schimawallichii inDig Kat andGaj Syzygiumcerasoides in Bij and Bor and Microcos paniculata Gmelinaarborea and Shorea robusta in Tar Lon and Sal respectivelyHolarrhena antidysenterica Microcos paniculata Terminaliabellirica Glochidion multiloculare Chaetocarpus castanicar-pus Syzygium cerasoides and Ziziphus rugosa were observedas codominant association although Aporosa obovata Can-thium glabrum Desmos dumosus Elaeocarpus prunifoliaElaeocarpus robustus Bhesa robusta Leea guineensis Litseacubeba and Litsea laeta were recorded with very low abun-dance and population (Figure 3)

32 Patterns of Stand Density Basal Area and Impor-tant Value Index (IVI) A total of 7134 individuals wereencountered with mean density of 64855 trees haminus1(rangedfrom 428 to 884 haminus1) in eleven 05 ha of fragmentedforest sites The highest woody density was recorded forBor (884 haminus1) and lowest for Sal (428 haminus1) Top tendominant trees namely Microcos paniculata (4382 haminus1)Tectona grandis (4309 haminus1) Holarrhena antidysenterica(3036 haminus1) Syzygium cerasoides (2764 haminus1) Shorea robusta(2491 haminus1) Schima wallichii (2345 haminus1) Syzygium frutico-sum (2145 haminus1) Melocanna baccifera (1782 haminus1) Gmelinaarborea (1745 haminus1) and Terminalia bellirica (1655 haminus1)

Journal of Ecosystems 5

Table 2 Consolidated details of woody plant diversity inventory (ge10 cm GBH) in eleven fragmented moist deciduous forests of Tripura

Parameters Fragmented forest sites Mean Standard error Standard deviationSim Dig Tar Kat Sub Mon Bij Gaj Bor Lon Sal

Number of woody species 60 61 41 45 45 40 37 61 39 47 31 4609 311 1032Number of genera 50 54 31 42 39 37 29 54 33 41 31 4009 276 916Number of families 29 32 22 26 32 28 23 37 25 28 25 2791 134 444Shannon index 373 357 333 349 332 347 275 392 258 338 22 325 016 052Simpson index 003 004 005 004 007 004 002 002 002 006 03 006 002 008Evenness index 21 2 206 21 201 216 164 22 162 201 147 194 007 025Richness index 264 208 164 22 163 17 142 247 131 171 15 185 013 044Density (haminus1) 518 854 628 436 762 552 680 608 884 784 428 64855 4787 15878Basal area (m2haminus1) 528 1408 68 145 1255 9 1644 236 1404 166 4708 1636 343 1137

Spec

ies a

bund

ance

Species rank

0102030405060708090

100110120130140150

1 10010

SimDigTarKatSubMon

BijGajBorLonSal

Figure 3 Diversity-dominance curve and species ranking based onspecies abundance

contributed in 4110 of overall density Mean basal areawas 1636m2 haminus1 for the whole study of which top tendominant trees grasped 944m2 haminus1 (5768) of total basalarea and it recorded the minimum for Sim (528m2 haminus1)and was the greatest in case of Sal (4708m2 haminus1) Basedon the IVI top ten dominant and codominant trees con-tributed in 11011 of total IVI (300) Shorea robusta graspsthe maximum IVI (3053) which was predominant only inSal followed by Tectona grandis (1170) Microcos paniculata(1122) Schima wallichii (1020) Syzygium cerasoides (980)Holarrhena antidysenterica (815) Artocarpus heterophyllus(754) and Terminalia bellirica (712) While 18 species sowedIVI value lt030 Desmos dumosus Leea aequata Elaeocarpusrobustus and Actinodaphne obovata were recorded withvery low IVI contribution to the fragmented forest Overall27 species were selected (top five most dominant speciesfrom each site) based on their IVI contribution to thosefragmented forests which suggested that Microcos panicu-lata Holarrhena antidysentericaChaetocarpus castanicarpusGlochidion multilocure Syzygium cerasoides and Ziziphusrugosawere increasing their IVI over previously predominant

trees These small tress species showed very little differencesin terms of IVI contribution with key dominant speciesnamely Shorea robusta SchimawallichiiTectona grandis andGmelina arborea (Figure 4)

33 Changes in Girth or Age-Class and Height-Class Distribu-tions Stem density across girth classes in all sites decreasesalmost from smallest to largest trees While occurrencesof mean stems haminus1 at lower girth class (10ndashlt30 cm) were5088 (330 plusmn 954) followed by 2910 (18873 plusmn 9694)in 30ndashlt60 cm girth class and 1096 (7109 plusmn 3798) in 60ndashlt90 cm girth class only 006 stems were shown in thehighest (240ndashlt300 cm) girth classes Maximum number ofindividuals at lower girth classwas recorded for Bor (468) andminimum for both Kat and Gaj (212) While few individualswere recorded at higher girth class (gt270 cm) some highergirth classes were absent in Sim Tar Sub Mon Bor andLon Stems were more frequently distributed in all the elevensites within the 10ndashlt120 cm girth classes The overall stemdensity haminus1 at different age classes was significant (1199032adj =093 119875 lt 00001) among the studied sites and showedinverse quadratic relationship (Figure 5) The distributionof stems by height class intervals showed that maximumtrees (8247) were widely distributed within 3ndash12m range3146 of individuals weregt3m followed by 2363 between3 and lt6m and 1435 within 6ndashlt9m whereas 720 ofindividuals distributed between 12 and lt15m only 042individuals were attained gt26m Density of trees was linearlydeclined across the height classes (1199032adj = 095 119875 lt 00001)among the fragmented sites (Figure 6)

34 Effects of Disturbances and Recovery Processes Besidesserious past logging experience for timber several otheranthropogenic disturbances were still ongoing in those frag-mented forests sites Grazing and fuelwood collection weremore intense due to closeness of village A total of 214trees haminus1 (1945 plusmn 1236) were recorded as cut or loggedand 24 trees haminus1 (218 plusmn 275) as injured out of overall7134 individuals maximum number of cut and injured treeswere recorded both from Dig and Sub Both cut and injuredtrees were 238 (2164 plusmn 1450) for overall study but wereabsent in case of Sal Site disturbance intensity was obtained

6 Journal of Ecosystems

minus16

minus8

0

8

16

24

32

40

48M

ean

and

stan

dard

err

or o

f |V

|

Albizia

p

Albizia

s

Artocarpu

Bauhinia

Careyaar

Chaetocar

Ficusrel

Garcinia

Glochidio

Gmelinaa

Holarrhen

Lannea

co

Litseamo

Mallotus

Mang

ifera

Microcos

Schim

awa

Shorea

ro

Streblus

Syzygium

Syzygium

Tamarindu

Tectonag

Terminali

Tremaori

Zanthoxyl

Ziziphus

Figure 4 Dominance and codominance competition of 27 tree species (mean and standard error of IVI) Five most dominant species wereselected from each fragmented forest site based on highest IVI contribution to the community

0 20 40 60 80 100120140160180200220240260280300minus02

00020406081012141618202224262830

Midvalue value of girth class (cm)

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 093 P lt 00001

Log

of d

ensit

y (h

a)

minus1

Figure 5 Relationship between population density (log trans-formed) and midvalue of girth class intervals across the fragmentedforest sites

qualitatively by summing the disturbance scores along andthe number of cut and injured individuals (Table 1(B)) Itcould be assumed that the proportion of scores that showedmaximum value revealed comparatively high level of anthro-pogenic disturbance or fragmentation than the lower valueThe highest intensity was recorded for Sub and Dig (1485)followed by Lon (1232) Gaj (1148) and Bor (1046)and the lowest in case of Sal (224) While the number

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30minus02

00020406081012141618202224262830

Midvalue value of height class (m)

Log

of d

ensit

y (h

a)

minus1

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 095 P lt 00001

Figure 6 Relationship between population density (log trans-formed) and midvalue of height class intervals across the frag-mented forest sites

of species (that occurred on each site) was significantlyincreased with disturbance intensity (1205942 = 006 F = 392119875 = 005) at 15 level and declining trend was observedwhen intensity level was increased among the sites (Figure 7)initially declining trend of the dominance of key communityspecies was observed and at midlevel of disturbance (10)few fragmentation tolerant species significantly (1205942 = 036 F= 872 119875 = 0009) increased in dominance with disturbance

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 3: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

Journal of Ecosystems 3

Table1Sitecharacteris

ticsa

nddistu

rbance

intensity

with

inthefragm

entedforests

Fragmentedforestsites

AL

ocations

BSitedistu

rbanceslowast

Latitud

e(ddm

mss)

Long

itude

(ddm

mss)

Altitude

(m)

Fire

Grazing

Fuelwoo

dcollection

Thatch

collection

Soil

removal

NTF

Pcollection

Num

bero

fcuttrees

Num

bero

fdead

trees

Total

score

Disturbance

intensity

()

Simna

(Sim

)24∘

051015840

349810158401015840

91∘

261015840

101410158401015840

6857

22

31

12

202

3392

4Diggyatila

(Dig)

24∘

051015840

153610158401015840

91∘

261015840

396610158401015840

6736

13

21

11

368

531485

Taranagar(Tar)

24∘

051015840

141110158401015840

91∘

241015840

301010158401015840

2895

13

33

11

122

2672

8Ka

tlamara(

Kat)

24∘

041015840

132010158401015840

91∘

231015840

197010158401015840

2956

12

22

22

100

21588

Subalsh

ing(Sub

)24∘

001015840

160010158401015840

91∘

271015840

375010158401015840

7559

21

31

22

366

531485

Mon

tala(M

on)

24∘

011015840

104010158401015840

91∘

261015840

121010158401015840

5577

23

13

11

40

15420

Bijoyn

agar

(Bij)

24∘

001015840

238010158401015840

91∘

231015840

390010158401015840

3078

12

22

11

160

2570

0Gajapara(

Gaj)

23∘

551015840

450010158401015840

91∘

291015840

228010158401015840

9022

22

22

21

264

411148

Borkathal(Bo

r)23∘

571015840

550010158401015840

91∘

241015840

040010158401015840

3596

13

33

22

222

381064

Lonk

amura(

Lon)

23∘

521015840

325010158401015840

91∘

141015840

543010158401015840

1859

13

23

21

320

441232

Salbagan

(Sal)

23∘

531015840

018010158401015840

91∘

171015840

182010158401015840

3535

12

11

21

00

8224

Total

1526

2422

1715

214

24357

100

Mean

133

233

211

222

167

133

1756

156

3011

843

Standard

error

017

024

026

028

017

017

412

073

492

138

Standard

deviation

050

071

078

083

050

050

1236

219

1475

413

lowast

Scoringrare1occasional2andrecurrent3and1score

fore

very

cutand

dead

stem

4 Journal of Ecosystems

Relative frequency = (Frequency of the speciesTotal fre-quency of all species) times 100 Relative dominance = (Basal areaof the speciesTotal basal area for all species)times 100 ImportantValue Index (IVI) = Relative Frequency + Relative Density+ Relative Basal Area General diversity was calculated usingShannon andWiener index [29] and the index of dominanceof the community was calculated by Simpsonrsquos index [30]Shannon and Wienerrsquos index = minussum(119899119894119873)log

2

(119899119894119873) ni =IVI of the species i N = IVI of all species in the communitySimpson dominance index = sum(119899119894119873)2 ni = IVI of speciesi N = IVI of all species in the community Species richnessindex was computed using Menhinick [31] and Evennessindex of the forest community was calculated followingPielou [32] Menhinick richness index = SradicN S = totalnumber of species N = total number of individuals Pielouevenness index = Shannon and Wienerrsquos indexlog(S) S =total number of species

The number of species was plotted on 119910-axis against thenumber of individuals on the 119909-axis for comparing speciesrichness among the fragmented forests [33] and speciesabundance data was plotted against species rank to developdiversity versus dominance curves [34] using BioDiversityPro version 2 [35] Woody species was grouped among tengirth classes (ge10ndashlt30 30ndashlt60 60ndashlt90 90ndashlt120 120ndashlt150150ndashlt180 180ndashlt210 210ndashlt240 240ndashlt270 and gt270 cm) andheight classes (lt3 3ndashlt6 6ndashlt9 9ndashlt12 12ndashlt15 15ndashlt18 18ndashlt2121ndashlt23 24ndashlt26 and gt26m) to observe the distribution ofpopulation at different age and height classes To examine thepattern of changes in community types and stand structureacross the studied sites regression curve fitting was used withmidvalue of overall stem density haminus1 girth and height classeson 119909-axis and log transformed density haminus1 on 119910-axis [36]In addition to test intermediate disturbance hypothesis weexamined the relationship by curve fitting between densitydiversity and dominance on 119910-axis and disturbance intensityon119909-axis for those fragmented forests usingOrigin and PASTsoftware [37 38]

3 Results

31 Status of Species Richness and Habitat Diversity AttributesA total of 134 woody species (ge10 cmGBH) representing93 genera and 43 families were recorded within 55 hafragmented moist deciduous forest of West Tripura Habit-wise out of 134woody species 118 species (8806)were trees7 (522) species were woody liana 4 (299) species werewoody shrub and 5 (373) species were bamboo Speciesrichness ranged from 31 to 61 species in eleven sites BothDig and Gaj had 61 species which was two times greater thanSal (31) The observed family richness was the greatest in Gaj(37)The rarefaction curves confirmed that sites Gaj SimDigand Kat were steeper than the others suggesting high speciesrichness (Figure 2)Whereas Simdid not asymptote at similarnumbers of individual which revealed that sufficient numberof species might not covered by our sampling effort and therewas possibility to get additional species in Sim compared toother sites Gaj scored high value of Shannon diversity index(392) while Sal recorded low value (220) compared to the

SimDigTarKatSubMon

BijGajBorLonSal

Expe

cted

num

ber o

f sp

ecie

s (S)

Number of individuals (N)

08

162432404856647280

0 50 100 150 200 250 300 350 400 450 500

Figure 2 Rarefaction curves for comparing the species richness ineleven fragmented forest stands

rest of the sites Mean Simpson dominance index was 006and was recorded high in Sub (007) which was recordedlow in Gaj (002) Species were more evenly distributed inSim Tar Gaj and Mon with a mean value of Evenness indexof 194 (range 147ndash216) while Menhinikrsquos species richnessvaried from 131 to 264 along the fragment sites with meanvalue of 185 (Table 2)

The diversity-dominance curve displayed a log seriesdistribution where species ranked (1ndash100) from most to leastabundant and majority of the species had lower abundanceand population while few species showed higher valuesTectona grandis appeared to be the most abundant in SimSub andMon Schimawallichii inDig Kat andGaj Syzygiumcerasoides in Bij and Bor and Microcos paniculata Gmelinaarborea and Shorea robusta in Tar Lon and Sal respectivelyHolarrhena antidysenterica Microcos paniculata Terminaliabellirica Glochidion multiloculare Chaetocarpus castanicar-pus Syzygium cerasoides and Ziziphus rugosa were observedas codominant association although Aporosa obovata Can-thium glabrum Desmos dumosus Elaeocarpus prunifoliaElaeocarpus robustus Bhesa robusta Leea guineensis Litseacubeba and Litsea laeta were recorded with very low abun-dance and population (Figure 3)

32 Patterns of Stand Density Basal Area and Impor-tant Value Index (IVI) A total of 7134 individuals wereencountered with mean density of 64855 trees haminus1(rangedfrom 428 to 884 haminus1) in eleven 05 ha of fragmentedforest sites The highest woody density was recorded forBor (884 haminus1) and lowest for Sal (428 haminus1) Top tendominant trees namely Microcos paniculata (4382 haminus1)Tectona grandis (4309 haminus1) Holarrhena antidysenterica(3036 haminus1) Syzygium cerasoides (2764 haminus1) Shorea robusta(2491 haminus1) Schima wallichii (2345 haminus1) Syzygium frutico-sum (2145 haminus1) Melocanna baccifera (1782 haminus1) Gmelinaarborea (1745 haminus1) and Terminalia bellirica (1655 haminus1)

Journal of Ecosystems 5

Table 2 Consolidated details of woody plant diversity inventory (ge10 cm GBH) in eleven fragmented moist deciduous forests of Tripura

Parameters Fragmented forest sites Mean Standard error Standard deviationSim Dig Tar Kat Sub Mon Bij Gaj Bor Lon Sal

Number of woody species 60 61 41 45 45 40 37 61 39 47 31 4609 311 1032Number of genera 50 54 31 42 39 37 29 54 33 41 31 4009 276 916Number of families 29 32 22 26 32 28 23 37 25 28 25 2791 134 444Shannon index 373 357 333 349 332 347 275 392 258 338 22 325 016 052Simpson index 003 004 005 004 007 004 002 002 002 006 03 006 002 008Evenness index 21 2 206 21 201 216 164 22 162 201 147 194 007 025Richness index 264 208 164 22 163 17 142 247 131 171 15 185 013 044Density (haminus1) 518 854 628 436 762 552 680 608 884 784 428 64855 4787 15878Basal area (m2haminus1) 528 1408 68 145 1255 9 1644 236 1404 166 4708 1636 343 1137

Spec

ies a

bund

ance

Species rank

0102030405060708090

100110120130140150

1 10010

SimDigTarKatSubMon

BijGajBorLonSal

Figure 3 Diversity-dominance curve and species ranking based onspecies abundance

contributed in 4110 of overall density Mean basal areawas 1636m2 haminus1 for the whole study of which top tendominant trees grasped 944m2 haminus1 (5768) of total basalarea and it recorded the minimum for Sim (528m2 haminus1)and was the greatest in case of Sal (4708m2 haminus1) Basedon the IVI top ten dominant and codominant trees con-tributed in 11011 of total IVI (300) Shorea robusta graspsthe maximum IVI (3053) which was predominant only inSal followed by Tectona grandis (1170) Microcos paniculata(1122) Schima wallichii (1020) Syzygium cerasoides (980)Holarrhena antidysenterica (815) Artocarpus heterophyllus(754) and Terminalia bellirica (712) While 18 species sowedIVI value lt030 Desmos dumosus Leea aequata Elaeocarpusrobustus and Actinodaphne obovata were recorded withvery low IVI contribution to the fragmented forest Overall27 species were selected (top five most dominant speciesfrom each site) based on their IVI contribution to thosefragmented forests which suggested that Microcos panicu-lata Holarrhena antidysentericaChaetocarpus castanicarpusGlochidion multilocure Syzygium cerasoides and Ziziphusrugosawere increasing their IVI over previously predominant

trees These small tress species showed very little differencesin terms of IVI contribution with key dominant speciesnamely Shorea robusta SchimawallichiiTectona grandis andGmelina arborea (Figure 4)

33 Changes in Girth or Age-Class and Height-Class Distribu-tions Stem density across girth classes in all sites decreasesalmost from smallest to largest trees While occurrencesof mean stems haminus1 at lower girth class (10ndashlt30 cm) were5088 (330 plusmn 954) followed by 2910 (18873 plusmn 9694)in 30ndashlt60 cm girth class and 1096 (7109 plusmn 3798) in 60ndashlt90 cm girth class only 006 stems were shown in thehighest (240ndashlt300 cm) girth classes Maximum number ofindividuals at lower girth classwas recorded for Bor (468) andminimum for both Kat and Gaj (212) While few individualswere recorded at higher girth class (gt270 cm) some highergirth classes were absent in Sim Tar Sub Mon Bor andLon Stems were more frequently distributed in all the elevensites within the 10ndashlt120 cm girth classes The overall stemdensity haminus1 at different age classes was significant (1199032adj =093 119875 lt 00001) among the studied sites and showedinverse quadratic relationship (Figure 5) The distributionof stems by height class intervals showed that maximumtrees (8247) were widely distributed within 3ndash12m range3146 of individuals weregt3m followed by 2363 between3 and lt6m and 1435 within 6ndashlt9m whereas 720 ofindividuals distributed between 12 and lt15m only 042individuals were attained gt26m Density of trees was linearlydeclined across the height classes (1199032adj = 095 119875 lt 00001)among the fragmented sites (Figure 6)

34 Effects of Disturbances and Recovery Processes Besidesserious past logging experience for timber several otheranthropogenic disturbances were still ongoing in those frag-mented forests sites Grazing and fuelwood collection weremore intense due to closeness of village A total of 214trees haminus1 (1945 plusmn 1236) were recorded as cut or loggedand 24 trees haminus1 (218 plusmn 275) as injured out of overall7134 individuals maximum number of cut and injured treeswere recorded both from Dig and Sub Both cut and injuredtrees were 238 (2164 plusmn 1450) for overall study but wereabsent in case of Sal Site disturbance intensity was obtained

6 Journal of Ecosystems

minus16

minus8

0

8

16

24

32

40

48M

ean

and

stan

dard

err

or o

f |V

|

Albizia

p

Albizia

s

Artocarpu

Bauhinia

Careyaar

Chaetocar

Ficusrel

Garcinia

Glochidio

Gmelinaa

Holarrhen

Lannea

co

Litseamo

Mallotus

Mang

ifera

Microcos

Schim

awa

Shorea

ro

Streblus

Syzygium

Syzygium

Tamarindu

Tectonag

Terminali

Tremaori

Zanthoxyl

Ziziphus

Figure 4 Dominance and codominance competition of 27 tree species (mean and standard error of IVI) Five most dominant species wereselected from each fragmented forest site based on highest IVI contribution to the community

0 20 40 60 80 100120140160180200220240260280300minus02

00020406081012141618202224262830

Midvalue value of girth class (cm)

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 093 P lt 00001

Log

of d

ensit

y (h

a)

minus1

Figure 5 Relationship between population density (log trans-formed) and midvalue of girth class intervals across the fragmentedforest sites

qualitatively by summing the disturbance scores along andthe number of cut and injured individuals (Table 1(B)) Itcould be assumed that the proportion of scores that showedmaximum value revealed comparatively high level of anthro-pogenic disturbance or fragmentation than the lower valueThe highest intensity was recorded for Sub and Dig (1485)followed by Lon (1232) Gaj (1148) and Bor (1046)and the lowest in case of Sal (224) While the number

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30minus02

00020406081012141618202224262830

Midvalue value of height class (m)

Log

of d

ensit

y (h

a)

minus1

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 095 P lt 00001

Figure 6 Relationship between population density (log trans-formed) and midvalue of height class intervals across the frag-mented forest sites

of species (that occurred on each site) was significantlyincreased with disturbance intensity (1205942 = 006 F = 392119875 = 005) at 15 level and declining trend was observedwhen intensity level was increased among the sites (Figure 7)initially declining trend of the dominance of key communityspecies was observed and at midlevel of disturbance (10)few fragmentation tolerant species significantly (1205942 = 036 F= 872 119875 = 0009) increased in dominance with disturbance

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

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Environmental and Public Health

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Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

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MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

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Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

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Environmental Chemistry

Atmospheric SciencesInternational Journal of

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Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

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ClimatologyJournal of

Page 4: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

4 Journal of Ecosystems

Relative frequency = (Frequency of the speciesTotal fre-quency of all species) times 100 Relative dominance = (Basal areaof the speciesTotal basal area for all species)times 100 ImportantValue Index (IVI) = Relative Frequency + Relative Density+ Relative Basal Area General diversity was calculated usingShannon andWiener index [29] and the index of dominanceof the community was calculated by Simpsonrsquos index [30]Shannon and Wienerrsquos index = minussum(119899119894119873)log

2

(119899119894119873) ni =IVI of the species i N = IVI of all species in the communitySimpson dominance index = sum(119899119894119873)2 ni = IVI of speciesi N = IVI of all species in the community Species richnessindex was computed using Menhinick [31] and Evennessindex of the forest community was calculated followingPielou [32] Menhinick richness index = SradicN S = totalnumber of species N = total number of individuals Pielouevenness index = Shannon and Wienerrsquos indexlog(S) S =total number of species

The number of species was plotted on 119910-axis against thenumber of individuals on the 119909-axis for comparing speciesrichness among the fragmented forests [33] and speciesabundance data was plotted against species rank to developdiversity versus dominance curves [34] using BioDiversityPro version 2 [35] Woody species was grouped among tengirth classes (ge10ndashlt30 30ndashlt60 60ndashlt90 90ndashlt120 120ndashlt150150ndashlt180 180ndashlt210 210ndashlt240 240ndashlt270 and gt270 cm) andheight classes (lt3 3ndashlt6 6ndashlt9 9ndashlt12 12ndashlt15 15ndashlt18 18ndashlt2121ndashlt23 24ndashlt26 and gt26m) to observe the distribution ofpopulation at different age and height classes To examine thepattern of changes in community types and stand structureacross the studied sites regression curve fitting was used withmidvalue of overall stem density haminus1 girth and height classeson 119909-axis and log transformed density haminus1 on 119910-axis [36]In addition to test intermediate disturbance hypothesis weexamined the relationship by curve fitting between densitydiversity and dominance on 119910-axis and disturbance intensityon119909-axis for those fragmented forests usingOrigin and PASTsoftware [37 38]

3 Results

31 Status of Species Richness and Habitat Diversity AttributesA total of 134 woody species (ge10 cmGBH) representing93 genera and 43 families were recorded within 55 hafragmented moist deciduous forest of West Tripura Habit-wise out of 134woody species 118 species (8806)were trees7 (522) species were woody liana 4 (299) species werewoody shrub and 5 (373) species were bamboo Speciesrichness ranged from 31 to 61 species in eleven sites BothDig and Gaj had 61 species which was two times greater thanSal (31) The observed family richness was the greatest in Gaj(37)The rarefaction curves confirmed that sites Gaj SimDigand Kat were steeper than the others suggesting high speciesrichness (Figure 2)Whereas Simdid not asymptote at similarnumbers of individual which revealed that sufficient numberof species might not covered by our sampling effort and therewas possibility to get additional species in Sim compared toother sites Gaj scored high value of Shannon diversity index(392) while Sal recorded low value (220) compared to the

SimDigTarKatSubMon

BijGajBorLonSal

Expe

cted

num

ber o

f sp

ecie

s (S)

Number of individuals (N)

08

162432404856647280

0 50 100 150 200 250 300 350 400 450 500

Figure 2 Rarefaction curves for comparing the species richness ineleven fragmented forest stands

rest of the sites Mean Simpson dominance index was 006and was recorded high in Sub (007) which was recordedlow in Gaj (002) Species were more evenly distributed inSim Tar Gaj and Mon with a mean value of Evenness indexof 194 (range 147ndash216) while Menhinikrsquos species richnessvaried from 131 to 264 along the fragment sites with meanvalue of 185 (Table 2)

The diversity-dominance curve displayed a log seriesdistribution where species ranked (1ndash100) from most to leastabundant and majority of the species had lower abundanceand population while few species showed higher valuesTectona grandis appeared to be the most abundant in SimSub andMon Schimawallichii inDig Kat andGaj Syzygiumcerasoides in Bij and Bor and Microcos paniculata Gmelinaarborea and Shorea robusta in Tar Lon and Sal respectivelyHolarrhena antidysenterica Microcos paniculata Terminaliabellirica Glochidion multiloculare Chaetocarpus castanicar-pus Syzygium cerasoides and Ziziphus rugosa were observedas codominant association although Aporosa obovata Can-thium glabrum Desmos dumosus Elaeocarpus prunifoliaElaeocarpus robustus Bhesa robusta Leea guineensis Litseacubeba and Litsea laeta were recorded with very low abun-dance and population (Figure 3)

32 Patterns of Stand Density Basal Area and Impor-tant Value Index (IVI) A total of 7134 individuals wereencountered with mean density of 64855 trees haminus1(rangedfrom 428 to 884 haminus1) in eleven 05 ha of fragmentedforest sites The highest woody density was recorded forBor (884 haminus1) and lowest for Sal (428 haminus1) Top tendominant trees namely Microcos paniculata (4382 haminus1)Tectona grandis (4309 haminus1) Holarrhena antidysenterica(3036 haminus1) Syzygium cerasoides (2764 haminus1) Shorea robusta(2491 haminus1) Schima wallichii (2345 haminus1) Syzygium frutico-sum (2145 haminus1) Melocanna baccifera (1782 haminus1) Gmelinaarborea (1745 haminus1) and Terminalia bellirica (1655 haminus1)

Journal of Ecosystems 5

Table 2 Consolidated details of woody plant diversity inventory (ge10 cm GBH) in eleven fragmented moist deciduous forests of Tripura

Parameters Fragmented forest sites Mean Standard error Standard deviationSim Dig Tar Kat Sub Mon Bij Gaj Bor Lon Sal

Number of woody species 60 61 41 45 45 40 37 61 39 47 31 4609 311 1032Number of genera 50 54 31 42 39 37 29 54 33 41 31 4009 276 916Number of families 29 32 22 26 32 28 23 37 25 28 25 2791 134 444Shannon index 373 357 333 349 332 347 275 392 258 338 22 325 016 052Simpson index 003 004 005 004 007 004 002 002 002 006 03 006 002 008Evenness index 21 2 206 21 201 216 164 22 162 201 147 194 007 025Richness index 264 208 164 22 163 17 142 247 131 171 15 185 013 044Density (haminus1) 518 854 628 436 762 552 680 608 884 784 428 64855 4787 15878Basal area (m2haminus1) 528 1408 68 145 1255 9 1644 236 1404 166 4708 1636 343 1137

Spec

ies a

bund

ance

Species rank

0102030405060708090

100110120130140150

1 10010

SimDigTarKatSubMon

BijGajBorLonSal

Figure 3 Diversity-dominance curve and species ranking based onspecies abundance

contributed in 4110 of overall density Mean basal areawas 1636m2 haminus1 for the whole study of which top tendominant trees grasped 944m2 haminus1 (5768) of total basalarea and it recorded the minimum for Sim (528m2 haminus1)and was the greatest in case of Sal (4708m2 haminus1) Basedon the IVI top ten dominant and codominant trees con-tributed in 11011 of total IVI (300) Shorea robusta graspsthe maximum IVI (3053) which was predominant only inSal followed by Tectona grandis (1170) Microcos paniculata(1122) Schima wallichii (1020) Syzygium cerasoides (980)Holarrhena antidysenterica (815) Artocarpus heterophyllus(754) and Terminalia bellirica (712) While 18 species sowedIVI value lt030 Desmos dumosus Leea aequata Elaeocarpusrobustus and Actinodaphne obovata were recorded withvery low IVI contribution to the fragmented forest Overall27 species were selected (top five most dominant speciesfrom each site) based on their IVI contribution to thosefragmented forests which suggested that Microcos panicu-lata Holarrhena antidysentericaChaetocarpus castanicarpusGlochidion multilocure Syzygium cerasoides and Ziziphusrugosawere increasing their IVI over previously predominant

trees These small tress species showed very little differencesin terms of IVI contribution with key dominant speciesnamely Shorea robusta SchimawallichiiTectona grandis andGmelina arborea (Figure 4)

33 Changes in Girth or Age-Class and Height-Class Distribu-tions Stem density across girth classes in all sites decreasesalmost from smallest to largest trees While occurrencesof mean stems haminus1 at lower girth class (10ndashlt30 cm) were5088 (330 plusmn 954) followed by 2910 (18873 plusmn 9694)in 30ndashlt60 cm girth class and 1096 (7109 plusmn 3798) in 60ndashlt90 cm girth class only 006 stems were shown in thehighest (240ndashlt300 cm) girth classes Maximum number ofindividuals at lower girth classwas recorded for Bor (468) andminimum for both Kat and Gaj (212) While few individualswere recorded at higher girth class (gt270 cm) some highergirth classes were absent in Sim Tar Sub Mon Bor andLon Stems were more frequently distributed in all the elevensites within the 10ndashlt120 cm girth classes The overall stemdensity haminus1 at different age classes was significant (1199032adj =093 119875 lt 00001) among the studied sites and showedinverse quadratic relationship (Figure 5) The distributionof stems by height class intervals showed that maximumtrees (8247) were widely distributed within 3ndash12m range3146 of individuals weregt3m followed by 2363 between3 and lt6m and 1435 within 6ndashlt9m whereas 720 ofindividuals distributed between 12 and lt15m only 042individuals were attained gt26m Density of trees was linearlydeclined across the height classes (1199032adj = 095 119875 lt 00001)among the fragmented sites (Figure 6)

34 Effects of Disturbances and Recovery Processes Besidesserious past logging experience for timber several otheranthropogenic disturbances were still ongoing in those frag-mented forests sites Grazing and fuelwood collection weremore intense due to closeness of village A total of 214trees haminus1 (1945 plusmn 1236) were recorded as cut or loggedand 24 trees haminus1 (218 plusmn 275) as injured out of overall7134 individuals maximum number of cut and injured treeswere recorded both from Dig and Sub Both cut and injuredtrees were 238 (2164 plusmn 1450) for overall study but wereabsent in case of Sal Site disturbance intensity was obtained

6 Journal of Ecosystems

minus16

minus8

0

8

16

24

32

40

48M

ean

and

stan

dard

err

or o

f |V

|

Albizia

p

Albizia

s

Artocarpu

Bauhinia

Careyaar

Chaetocar

Ficusrel

Garcinia

Glochidio

Gmelinaa

Holarrhen

Lannea

co

Litseamo

Mallotus

Mang

ifera

Microcos

Schim

awa

Shorea

ro

Streblus

Syzygium

Syzygium

Tamarindu

Tectonag

Terminali

Tremaori

Zanthoxyl

Ziziphus

Figure 4 Dominance and codominance competition of 27 tree species (mean and standard error of IVI) Five most dominant species wereselected from each fragmented forest site based on highest IVI contribution to the community

0 20 40 60 80 100120140160180200220240260280300minus02

00020406081012141618202224262830

Midvalue value of girth class (cm)

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 093 P lt 00001

Log

of d

ensit

y (h

a)

minus1

Figure 5 Relationship between population density (log trans-formed) and midvalue of girth class intervals across the fragmentedforest sites

qualitatively by summing the disturbance scores along andthe number of cut and injured individuals (Table 1(B)) Itcould be assumed that the proportion of scores that showedmaximum value revealed comparatively high level of anthro-pogenic disturbance or fragmentation than the lower valueThe highest intensity was recorded for Sub and Dig (1485)followed by Lon (1232) Gaj (1148) and Bor (1046)and the lowest in case of Sal (224) While the number

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30minus02

00020406081012141618202224262830

Midvalue value of height class (m)

Log

of d

ensit

y (h

a)

minus1

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 095 P lt 00001

Figure 6 Relationship between population density (log trans-formed) and midvalue of height class intervals across the frag-mented forest sites

of species (that occurred on each site) was significantlyincreased with disturbance intensity (1205942 = 006 F = 392119875 = 005) at 15 level and declining trend was observedwhen intensity level was increased among the sites (Figure 7)initially declining trend of the dominance of key communityspecies was observed and at midlevel of disturbance (10)few fragmentation tolerant species significantly (1205942 = 036 F= 872 119875 = 0009) increased in dominance with disturbance

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 5: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

Journal of Ecosystems 5

Table 2 Consolidated details of woody plant diversity inventory (ge10 cm GBH) in eleven fragmented moist deciduous forests of Tripura

Parameters Fragmented forest sites Mean Standard error Standard deviationSim Dig Tar Kat Sub Mon Bij Gaj Bor Lon Sal

Number of woody species 60 61 41 45 45 40 37 61 39 47 31 4609 311 1032Number of genera 50 54 31 42 39 37 29 54 33 41 31 4009 276 916Number of families 29 32 22 26 32 28 23 37 25 28 25 2791 134 444Shannon index 373 357 333 349 332 347 275 392 258 338 22 325 016 052Simpson index 003 004 005 004 007 004 002 002 002 006 03 006 002 008Evenness index 21 2 206 21 201 216 164 22 162 201 147 194 007 025Richness index 264 208 164 22 163 17 142 247 131 171 15 185 013 044Density (haminus1) 518 854 628 436 762 552 680 608 884 784 428 64855 4787 15878Basal area (m2haminus1) 528 1408 68 145 1255 9 1644 236 1404 166 4708 1636 343 1137

Spec

ies a

bund

ance

Species rank

0102030405060708090

100110120130140150

1 10010

SimDigTarKatSubMon

BijGajBorLonSal

Figure 3 Diversity-dominance curve and species ranking based onspecies abundance

contributed in 4110 of overall density Mean basal areawas 1636m2 haminus1 for the whole study of which top tendominant trees grasped 944m2 haminus1 (5768) of total basalarea and it recorded the minimum for Sim (528m2 haminus1)and was the greatest in case of Sal (4708m2 haminus1) Basedon the IVI top ten dominant and codominant trees con-tributed in 11011 of total IVI (300) Shorea robusta graspsthe maximum IVI (3053) which was predominant only inSal followed by Tectona grandis (1170) Microcos paniculata(1122) Schima wallichii (1020) Syzygium cerasoides (980)Holarrhena antidysenterica (815) Artocarpus heterophyllus(754) and Terminalia bellirica (712) While 18 species sowedIVI value lt030 Desmos dumosus Leea aequata Elaeocarpusrobustus and Actinodaphne obovata were recorded withvery low IVI contribution to the fragmented forest Overall27 species were selected (top five most dominant speciesfrom each site) based on their IVI contribution to thosefragmented forests which suggested that Microcos panicu-lata Holarrhena antidysentericaChaetocarpus castanicarpusGlochidion multilocure Syzygium cerasoides and Ziziphusrugosawere increasing their IVI over previously predominant

trees These small tress species showed very little differencesin terms of IVI contribution with key dominant speciesnamely Shorea robusta SchimawallichiiTectona grandis andGmelina arborea (Figure 4)

33 Changes in Girth or Age-Class and Height-Class Distribu-tions Stem density across girth classes in all sites decreasesalmost from smallest to largest trees While occurrencesof mean stems haminus1 at lower girth class (10ndashlt30 cm) were5088 (330 plusmn 954) followed by 2910 (18873 plusmn 9694)in 30ndashlt60 cm girth class and 1096 (7109 plusmn 3798) in 60ndashlt90 cm girth class only 006 stems were shown in thehighest (240ndashlt300 cm) girth classes Maximum number ofindividuals at lower girth classwas recorded for Bor (468) andminimum for both Kat and Gaj (212) While few individualswere recorded at higher girth class (gt270 cm) some highergirth classes were absent in Sim Tar Sub Mon Bor andLon Stems were more frequently distributed in all the elevensites within the 10ndashlt120 cm girth classes The overall stemdensity haminus1 at different age classes was significant (1199032adj =093 119875 lt 00001) among the studied sites and showedinverse quadratic relationship (Figure 5) The distributionof stems by height class intervals showed that maximumtrees (8247) were widely distributed within 3ndash12m range3146 of individuals weregt3m followed by 2363 between3 and lt6m and 1435 within 6ndashlt9m whereas 720 ofindividuals distributed between 12 and lt15m only 042individuals were attained gt26m Density of trees was linearlydeclined across the height classes (1199032adj = 095 119875 lt 00001)among the fragmented sites (Figure 6)

34 Effects of Disturbances and Recovery Processes Besidesserious past logging experience for timber several otheranthropogenic disturbances were still ongoing in those frag-mented forests sites Grazing and fuelwood collection weremore intense due to closeness of village A total of 214trees haminus1 (1945 plusmn 1236) were recorded as cut or loggedand 24 trees haminus1 (218 plusmn 275) as injured out of overall7134 individuals maximum number of cut and injured treeswere recorded both from Dig and Sub Both cut and injuredtrees were 238 (2164 plusmn 1450) for overall study but wereabsent in case of Sal Site disturbance intensity was obtained

6 Journal of Ecosystems

minus16

minus8

0

8

16

24

32

40

48M

ean

and

stan

dard

err

or o

f |V

|

Albizia

p

Albizia

s

Artocarpu

Bauhinia

Careyaar

Chaetocar

Ficusrel

Garcinia

Glochidio

Gmelinaa

Holarrhen

Lannea

co

Litseamo

Mallotus

Mang

ifera

Microcos

Schim

awa

Shorea

ro

Streblus

Syzygium

Syzygium

Tamarindu

Tectonag

Terminali

Tremaori

Zanthoxyl

Ziziphus

Figure 4 Dominance and codominance competition of 27 tree species (mean and standard error of IVI) Five most dominant species wereselected from each fragmented forest site based on highest IVI contribution to the community

0 20 40 60 80 100120140160180200220240260280300minus02

00020406081012141618202224262830

Midvalue value of girth class (cm)

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 093 P lt 00001

Log

of d

ensit

y (h

a)

minus1

Figure 5 Relationship between population density (log trans-formed) and midvalue of girth class intervals across the fragmentedforest sites

qualitatively by summing the disturbance scores along andthe number of cut and injured individuals (Table 1(B)) Itcould be assumed that the proportion of scores that showedmaximum value revealed comparatively high level of anthro-pogenic disturbance or fragmentation than the lower valueThe highest intensity was recorded for Sub and Dig (1485)followed by Lon (1232) Gaj (1148) and Bor (1046)and the lowest in case of Sal (224) While the number

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30minus02

00020406081012141618202224262830

Midvalue value of height class (m)

Log

of d

ensit

y (h

a)

minus1

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 095 P lt 00001

Figure 6 Relationship between population density (log trans-formed) and midvalue of height class intervals across the frag-mented forest sites

of species (that occurred on each site) was significantlyincreased with disturbance intensity (1205942 = 006 F = 392119875 = 005) at 15 level and declining trend was observedwhen intensity level was increased among the sites (Figure 7)initially declining trend of the dominance of key communityspecies was observed and at midlevel of disturbance (10)few fragmentation tolerant species significantly (1205942 = 036 F= 872 119875 = 0009) increased in dominance with disturbance

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 6: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

6 Journal of Ecosystems

minus16

minus8

0

8

16

24

32

40

48M

ean

and

stan

dard

err

or o

f |V

|

Albizia

p

Albizia

s

Artocarpu

Bauhinia

Careyaar

Chaetocar

Ficusrel

Garcinia

Glochidio

Gmelinaa

Holarrhen

Lannea

co

Litseamo

Mallotus

Mang

ifera

Microcos

Schim

awa

Shorea

ro

Streblus

Syzygium

Syzygium

Tamarindu

Tectonag

Terminali

Tremaori

Zanthoxyl

Ziziphus

Figure 4 Dominance and codominance competition of 27 tree species (mean and standard error of IVI) Five most dominant species wereselected from each fragmented forest site based on highest IVI contribution to the community

0 20 40 60 80 100120140160180200220240260280300minus02

00020406081012141618202224262830

Midvalue value of girth class (cm)

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 093 P lt 00001

Log

of d

ensit

y (h

a)

minus1

Figure 5 Relationship between population density (log trans-formed) and midvalue of girth class intervals across the fragmentedforest sites

qualitatively by summing the disturbance scores along andthe number of cut and injured individuals (Table 1(B)) Itcould be assumed that the proportion of scores that showedmaximum value revealed comparatively high level of anthro-pogenic disturbance or fragmentation than the lower valueThe highest intensity was recorded for Sub and Dig (1485)followed by Lon (1232) Gaj (1148) and Bor (1046)and the lowest in case of Sal (224) While the number

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30minus02

00020406081012141618202224262830

Midvalue value of height class (m)

Log

of d

ensit

y (h

a)

minus1

SimDigTarKatSubMon

BijGajBorLonSal

r2adj = 095 P lt 00001

Figure 6 Relationship between population density (log trans-formed) and midvalue of height class intervals across the frag-mented forest sites

of species (that occurred on each site) was significantlyincreased with disturbance intensity (1205942 = 006 F = 392119875 = 005) at 15 level and declining trend was observedwhen intensity level was increased among the sites (Figure 7)initially declining trend of the dominance of key communityspecies was observed and at midlevel of disturbance (10)few fragmentation tolerant species significantly (1205942 = 036 F= 872 119875 = 0009) increased in dominance with disturbance

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 7: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

Journal of Ecosystems 7

Sim Dig

TarKat Sub

MonBij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

14815215616

16416817217618

minus0001375x 2 + 003994x + 1426

Log

num

ber o

f sp

ecie

s

Figure 7 Relationship between species richness and disturbanceintensity

intensity (Figure 8) In addition relationship between stemdensity and disturbance intensity also suggested that presentnumber of stems per ha were increased significantly withfragmentation intensity (1205942 = 005 F = 665119875 = 002) duringrestoration (Figure 9)

4 Discussion

Rarefaction curves suggested that the maximum number ofspecies was counted for Gaj Sim and Dig while Sim didnot show asymptote and species richness was much lower inthe relatively managed Sal forest Higher species richness inthe fragmented sites might attribute to intermediate distur-bance hypothesis [21] Species richness might be enhancedin some of the sites through quick accumulation of somelocally adaptable early successional species The range ofspecies richness (31ndash61) recorded within the present studywas more than Puerto Rico 30ndash35 species [39] and moistdeciduous forest of Western Ghats was 24ndash26 species [40]However overall 134 species encountered in this inventorywere higher than the number of species reported fromsome well-protected Indian forests [41ndash43] The value ofShannon-Winner diversity index ranged from 220 to 392whereas this value generally ranges between 083 and 41 forIndian forests [41] hence present studiesrsquo value was fallenwithin the reported range Species diversity was increasedin case of Gaj Sim Dig Lon and Sub (Figure 7) along thedisturbance intensity (1199032adj = 049 119875 = 005) because diversityin communities increases under intermediate disturbanceregime [21] Instead of protection and management in Salby Forest Department it showed minimum species richnesswhichmight be due to presence of high density of voluminousS robusta which generally has strong competitive effects onseedling growth and regeneration of other species [36 42]However disturbances allow species to grow well in newenvironment recruited and established by increasing thenumber of species that are not found in managed areas [43]hence species richness fairly increased in those fragmentedforests

Stem density haminus1 at ge10 cm girth class in tropical forestgenerally varies from 245 to 859 trees haminus1 [44 45] present

SimDigTarKat

Sub

MonBij Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18 20Disturbance intensity ()

minus2minus18minus16minus14minus12minus1

minus08minus06minus04minus02

0 001614x 2 minus 03159x minus 008125

Log

of d

omin

ance

inde

x

Figure 8 Relationship between species dominance and disturbanceintensity

mean stand density of 64855 trees haminus1 (range 428ndash884 haminus1)was observed higher than protected Western Ghats moistdeciduous forest [46 47] Such relatively higher density ofthe present fragmented forests might be due to their patchycondition which resulted from past disturbances since thetrend of smaller treesrsquo density was reported to increaseafter historic disturbances [48] Significant inverse relationof overall stem density-age classes suggested that 5088of stems were at lower girth classes that is 10ndashlt30 cm(Figure 5) which was the indication of advance regenerationstages and possibly those small fragmented forests wererecuperating from past disturbances 214 trees haminus1 wererecorded as logged or cut in the whole study which alsosuggested illegal logging and heavy timber requirementsLower abundance or absence of upper canopy forming orold mature trees was also evident from overall stem density-height classes (Figure 6) and so forth which might result inenough gaps or forest openings where fragmentation tolerantsmall tree species were facilitated to regenerate quickly andincrease in their stem density [36 42] After removal ofmature or old aged key dominant tree species especially fortimber many opportunistic species might gather throughseeded or nonseeded multiplication in absence of previouslykey dominant species Hence stem density was significantlyincreased with disturbance intensity (1199032adj = 062 119875 = 002)in those sites as an early successional stage and the trendpossibly declined very slowly from this level towards forestmaturation (Figure 9)

Mean basal area value of 1636m2 haminus1 obtained inthe present study (range 528ndash4708m2 haminus1) was close tomoist deciduous forests of Western Ghats 147m2 haminus1 and1283m2 haminus1 [46 47] andwas less than 49m2 haminus1 in tropicalmoist deciduous forest of Eastern Ghats [49] Dominanceand codominance association in those fragmented forestsuggested that S robusta T grandis S wallichii and so forthstill have the control over upper canopy structure and basalarea But simultaneously locally adapted small oligarchicspecies were increasing their population density to keepthe codominant rank by maximum IVI contribution whichmight lead to the variation in community association overexisting forest structure by shifting of codominant species

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 8: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

8 Journal of Ecosystems

Sim

Dig

Tar

Kat

Sub

Mon

Bij

Gaj

Bor

Lon

Sal

0 2 4 6 8 10 12 14 16 18Disturbance intensity ()

26264268272276

28284288292296

minus00005312x 2 + 003021x + 2577

Log

of d

ensit

y (h

a)

minus1

Figure 9 Relationship between stem density and disturbanceintensity

into the most dominant species rank at such intensity of dis-turbances (Figure 4) Species dominance generally increasesas a function of stress or past disturbance [50] and thesespecies have vital role in the microenvironment within thecommunity [51] Higher abundance of several local oligarchicsmall tree species as codominant associations in those frag-mented forests was increased dominance especially in caseof Sub Lon and Dig (Figure 8) due to strong adaptabilityof those species with fragmentation intensity (1199032adj = 068119875 = 0009) The intensity of disturbances greatly influencedspecies dominance in those fragmented forests which mayaffect future forest structure and composition successionphases and species restoration Fragmentation may establishseveral transitional community associationswhich aremostlydominated by few local adaptable species through direct orindirect competitions through increasing IVI over previouslykey dominated species

5 Conclusions

Bamboo shared 9125 of overall IVI of the present frag-mented forest which contributed enormously in rural liveli-hood by fulfilling the NTFPs and fuelwood demand for thepeople living proximate of those fragmented forests Manybamboo patches were dried due to flowering especially inMelocanna baccifera Bambusa balcooa and B tulda whichin turn drastically changed the trend of demands for forestresources in those fragmented forests Demands for fuel-woods thatches and structural and fencing materials weremore or less likely to shift into the small and medium sizedtrees in absence of bamboo in those fragmented forests (afterflowering) which may further influence the natural processof restoration in those fragmented forests may restrictthe possibilities of species richness and locally enhancethe process of either species monodominance or rarenessFragmented forest plays a crucial role for conservation andsustaining local plant genetic diversity whichmay be the onlyremaining store for many natural products and threatenedspecies beyond protected areas Monitoring of fragmentedtree species and plantation of locally prioritized species in the

openedpatcheswill contribute towards effectivemanagementbetween forest dependent people and fragmented forests

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors are thankful to Department of Biotechnology(DBT) Government of India for funding received throughDBT Network Project (BTPR7928NDB5292006) Theauthors also thank Professor S Sinha HOD Departmentof Botany Tripura University for his valuable suggestionsand facilities Thanks also are extended to Sri Samir KumarDebnath and Sri Monthosh Roy for their consistent fieldassistance and several anonymous reviewers

References

[1] I M Turner H T W Tan Y C Wee A B Ibrahim P TChew and R T Corlett ldquoA study of plant species extinction inSingapore lessons for the conservation of tropical biodiversityrdquoConservation Biology vol 8 no 3 pp 705ndash712 1994

[2] M B Mulder and P Coppolillo Conservation Linking EcologyEconomics and Culture Princeton University Press PrincetonNJ USA 2005

[3] R Pither and M Kellman ldquoTree species diversity in smalltropical riparian forest fragments in Belize Central AmericardquoBiodiversity and Conservation vol 11 no 9 pp 1623ndash1636 2002

[4] H Jacquemyn J Butaye and M Hermy ldquoInfluence of envi-ronmental and spatial variables on regional distribution offorest plant species in a fragmented and changing landscaperdquoEcography vol 26 no 6 pp 768ndash776 2003

[5] J L Hill and P J Curran ldquoSpecies composition in fragmentedforests conservation implications of changing forest areardquoApplied Geography vol 21 no 2 pp 157ndash174 2001

[6] H Zhu Z F Xu H Wang and B G Li ldquoTropical rain forestfragmentation and its ecological and species diversity changesin southern Yunnanrdquo Biodiversity and Conservation vol 13 no7 pp 1355ndash1372 2004

[7] K Rennolls and Y Laumonier ldquoSpecies diversity structureanalysis at two sites in the tropical rain forest of SumatrardquoJournal of Tropical Ecology vol 16 no 2 pp 253ndash270 2000

[8] W Huang V Pohjonen S Johansson M Nashanda M I LKatigula and O Luukkanen ldquoSpecies diversity forest structureand species composition in Tanzanian tropical forestsrdquo ForestEcology and Management vol 173 no 1ndash3 pp 11ndash24 2003

[9] C H Cannon D R Peart and M Leighton ldquoTree speciesdiversity in commercially logged Bornean rainforestrdquo Sciencevol 281 no 5381 pp 1366ndash1368 1998

[10] T C Whitmore An Introduction to Tropical Rain ForestsOxford University Press Oxford UK 1993

[11] M A Hare D O Lantagne P G Murphy and H ChecoldquoStructure and tree species composition in a subtropical dryforest in the Dominican Republic comparison with a dry forestin Puerto Ricordquo Tropical Ecology vol 38 no 1 pp 1ndash17 1997

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 9: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

Journal of Ecosystems 9

[12] S P Hubbell R B Foster S T OrsquoBrien et al ldquoLight-gapdisturbances recruitment limitation and tree diversity in aneotropical forestrdquo Science vol 283 no 5401 pp 554ndash557 1999

[13] O I Sumina ldquoPlant communities on anthropogenically dis-turbed sites on the Chukotka Peninsula Russiardquo Journal ofVegetation Science vol 5 no 6 pp 885ndash896 1994

[14] E J Luoga E T F Witkowski and K Balkwill ldquoRegenerationby coppicing (resprouting) of miombo (African savanna) treesin relation to land userdquo Forest Ecology andManagement vol 189no 1ndash3 pp 23ndash35 2004

[15] M Johnston and M Gillman ldquoTree population studies in low-diversity forests Guyana I Floristic composition and standstructurerdquo Biodiversity and Conservation vol 4 no 4 pp 339ndash362 1995

[16] A S van Jaarsveld S Freitag S L Chown et al ldquoBiodiversityassessment and conservation strategiesrdquo Science vol 279 no5359 pp 2106ndash2108 1998

[17] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat low-elevation evergreen and moist deciduous forests atKodayar (W Ghats India)rdquo International Journal of Ecology andEnvironmental Sciences vol 23 no 4 pp 363ndash379 1997

[18] U Ghate N V Joshi and M Gadgil ldquoOn the patterns of treediversity in the Western Ghats of Indiardquo Current Science vol75 no 6 pp 594ndash603 1998

[19] H Padalia N Chauhan M C Porwal and P S Roy ldquoPhytoso-ciological observations on tree species diversity of AndamanIslands IndiardquoCurrent Science vol 87 no 6 pp 799ndash806 2004

[20] A Sen T Johri and N S Bisht ldquoAnalysis of the effects ofanthropogenic interferences on tree species composition in theforests of Dadra and Nagar Haveli Indiardquo Current Science vol95 no 1 pp 50ndash58 2008

[21] J H Connell ldquoDiversity in tropical rain forests and coral reefsrdquoScience vol 199 no 4335 pp 1302ndash1310 1978

[22] AnonymousThe State of Forest Report Forest Survey of IndiaDehradun India 2009

[23] U Shankar ldquoA case of high tree diversity in a sal (Shorearobusta)-dominated lowland forest of Eastern Himalaya floris-tic composition regeneration and conservationrdquo Current Sci-ence vol 81 no 7 pp 776ndash786 2001

[24] P Rao S K Barik H N Pandey and R S Tripathi ldquoCommu-nity composition and tree population structure in a sub-tropicalbroad-leaved forest along a disturbance gradientrdquoVegetatio vol88 no 2 pp 151ndash162 1990

[25] S Mani and N Parthasarathy ldquoTree diversity and stand struc-ture in inland and coastal tropical dry evergreen forests ofpeninsular Indiardquo Current Science vol 90 no 9 pp 1238ndash12462006

[26] D B Deb The Flora of Tripura State vol 1-2 Today andTomorrowrsquos Printers and Publishers New Delhi India 1981

[27] UNKanjilal P C Kanjilal ADas RN de andN L BorFloraof Assam vol 1-5 Government Press Shillong India 1940

[28] D Muller-Dombois and H Ellenberg Aims and Methods ofVegetation Ecology John Wiley and Sons 1974

[29] C E Shannon and W Wiener The Mathematical Theory ofCommunities University of Illinois Press Urbana Ill USA1963

[30] E H Simpson ldquoMeasurement of diversityrdquo Nature vol 163article 688 1949

[31] E F Menhinick ldquoA comparison of some speciesindividualsdiversity indices applied to samples of field insectsrdquoEcology vol45 pp 859ndash861 1964

[32] T Ganesh R Ganesan M Soubadra Devy P Davidar and K SBawa ldquoAssessment of plant biodiversity at amid-elevation ever-green forest of Kalakad-Mundanthurai Tiger Reserve WesternGhats Indiardquo Current Science vol 71 no 5 pp 379ndash392 1996

[33] D Simberloff ldquoProperties of the rarefaction diversity measure-mentrdquoThe American Naturalist vol 106 pp 414ndash418 1972

[34] K M Shaw P J D Lambshead and H M Platt ldquoDetection ofpollution induced disturbance in marine benthic assemblageswith special reference to nematodesrdquo Marine Ecology ProgressSeries vol 11 pp 195ndash202 1983

[35] N McAleece J D G Gage P J D Lambshead and G L JPaterson BioDiversity Professional Statistics Analysis SoftwareScottishAssociation forMarine Science and theNaturalHistoryMuseum London UK 1997

[36] I P Sapkota M Tigabu and P C Oden ldquoSpatial distributionadvanced regeneration and stand structure of Nepalese Sal(Shorea robusta) forests subject to disturbances of differentintensitiesrdquo Forest Ecology and Management vol 257 no 9 pp1966ndash1975 2009

[37] Origin 2013 Free version OriginLab Corporationhttpwwworiginlabcom

[38] O Hammer D A T Harper and P D Ryan ldquoPAST pale-ontological statistics software package for education and dataanalysis version 203rdquo Paleontologia Electronica vol 4 no 1 p9 2001

[39] P G Murphy and A E Lugo ldquoEcology of tropical dry forestrdquoAnnual Review of Ecology and Systematics vol 17 pp 67ndash881986

[40] N I Valappil and K Swarupanandan ldquoRegeneration dynamicsand sylvigenesis in the moist deciduous forests of southwestIndiardquo New Forests vol 11 no 3 pp 185ndash205 1996

[41] N Parthasarathy V Kinhal and L Praveenkumar ldquo Plantspecies diversity and human impacts in the tropical wet ever-green forests of southern Western Ghatsrdquo in Proceedings ofthe IndondashFrench Workshop on Tropical Forest Ecosystems Nat-ural Functioning and Anthropogenic Impact French InstitutePondicherry India November 1992

[42] S K Pandey and R P Shukla ldquoPlant diversity in managed sal(Shorea robusta Gaertn) forests of Gorakhpur India speciescomposition regeneration and conservationrdquo Biodiversity andConservation vol 12 no 11 pp 2295ndash2319 2003

[43] T Banda W Schwartz and T M Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of Western Tanzaniardquo Forest Ecology andManagement vol 230 no 1ndash3 pp 179ndash185 2006

[44] P S Ashton ldquoA quantitative phytosociological techniqueapplied to tropical mixed rain forest vegetationrdquo MalayanForester vol 27 pp 304ndash317 1964

[45] D G Campbell J L Stone and A Rosas Jr ldquoA comparison ofthe phytosociology and dynamics of three floodplain (Varzea)forests of known ages Rio Jurua Western Brazilian AmazonrdquoBotanical Journal of the Linnean Society vol 108 no 3 pp 213ndash237 1992

[46] S K Seth and O N Kaul ldquoTropical forest ecosystems of Indiathe teak forestrdquo in Unesco UNEP and FAO Tropical ForestEcosystems pp 628ndash640 Unesco-UNEP France 1978

[47] S M Sundarapandian and P S Swamy ldquoPlant biodiversityat Lowelevation Evergreen and Moist Deciduous Forests atKodayar in theWestern Ghatsrdquo International Journal of Ecologyand Environmental Science vol 23 no 4 pp 363ndash379 1997

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 10: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

10 Journal of Ecosystems

[48] P Bhuyan M L Khan and R S Tripathi ldquoTree diversityand population structure in undisturbed and human-impactedstands of tropical wet evergreen forest in Arunachal PradeshEastern Himalayas Indiardquo Biodiversity and Conservation vol12 no 8 pp 1753ndash1773 2003

[49] C S Reddy S Babar G Amarnath andC Pattanaik ldquoStructureand floristic composition of tree stand in tropical forest in theEastern Ghats of northern Andhra Pradesh Indiardquo Journal ofForestry Research vol 22 no 4 pp 491ndash500 2011

[50] R J Burnham ldquoDominance diversity and distribution of lianasin Yasunı Ecuador who is on toprdquo Journal of Tropical Ecologyvol 18 no 6 pp 845ndash864 2002

[51] M Cao and J Zhang ldquoTree species diversity of tropical forestvegetation in Xishuangbanna SW Chinardquo Biodiversity andConservation vol 6 no 7 pp 995ndash1006 1997

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 11: Research Article Trends in Tree Diversity and Stand Structure ...downloads.hindawi.com/archive/2014/845142.pdfT : Consolidated details of woody plant diversity inventory ( cm GBH)

Submit your manuscripts athttpwwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of