Tree Species Diversity and Composition of Miombo Woodlands ...downloads.hindawi.com/journals/ijfr/2017/6202093.pdf · The diversity profiles indicate that mature forests were more

Research ArticleTree Species Diversity and Composition of MiomboWoodlands in South-Central Angola A Chronosequence ofForest Recovery after Shifting Cultivation

Francisco M P Gonccedilalves12 Rasmus Revermann2 Amacircndio L Gomes23

Marcos P M Aidar4 Manfred Finckh2 and Norbert Juergens2

1Herbarium of Lubango ISCED Huıla Sarmento Rodrigues Str CP 230 Lubango Angola2Biocentre Klein Flottbek University of Hamburg Ohnhorststr 18 22609 Hamburg Germany3Department of Biology Faculty of Sciences Agostinho Neto University Av 4 de Fevereiro 71 4 Andar Luanda Angola4Plant Physiology and Biochemistry Institute of Botany CP 3005 01061-970 Sao Paulo SP Brazil

Correspondence should be addressed to Francisco M P Goncalves franciscomaiatogmailcom

Received 15 July 2016 Revised 29 November 2016 Accepted 14 December 2016 Published 1 March 2017

Academic Editor Piermaria Corona

Copyright copy 2017 Francisco M P Goncalves et al This is an open access article distributed under the Creative CommonsAttribution License which permits unrestricted use distribution and reproduction in any medium provided the original work isproperly cited

The study was carried out in the Cusseque area of the Municipality of Chitembo in south-central Angola Our objectives were toassess the floristic diversity the species composition and stand structure of Miombo woodlands during regeneration after shiftingcultivation A total of 40 plots of 1000m2 were surveyed and analyzed corresponding tomature forestswoodlands and three fallowtypes of different ageThe analyseswere based on plot inventories of all treeswithDBHge 5 cmA total of 51woody species 38 generaand 19 families were recordedThe dominant family was Fabaceae with subfamily Caesalpinioideae being very abundant ShannonDiversity and Evenness were highest in mature forests and young fallows while the mature forest stands showed the highest speciesrichness A Principal Coordinates Analysis (PCoA) showed many species shared between the intermediate fallow types but onlyfew species were shared with young fallows Mature forests formed a clearly distinct groupThis study shows potential pathways offorest recovery in terms of faster regeneration after agricultural abandonment and thus the results presented here can be used infuture conservation and management plans in order to reduce the pressure on mature forests

1 Introduction

Forests andwoodlands in Africa play an important role in thelivelihood of many communities and in the economic devel-opment of many countries [1] Forest ecosystems are impor-tant not only for supplying timber and other economicallyimportant products (eg charcoal) but also for generatingindirect benefits such as controlling soil erosion supportingsoil fertility providing shade and buffering hydrologicalcycles [2]

The forest resources in most African countries are threat-ened by various natural and anthropogenic disturbancesespecially overexploitation Thus the problem of unsustain-able use of forest resources is of increasing concern in thesecountries due to the fact that forest losses are causing land

degradation and habitat loss for forest taxa [3] Direct causesof deforestation and land degradation in south-central Africainclude land conversion for example by expansion of shiftingcultivation practiced by small holders and infrastructural oragroindustrial expansion and overexploitation for examplefor timber fuelwood and charcoal [4]

Shifting cultivation is an agricultural system practiced inthe tropics for thousands of years being the main sourceof subsistence for people in many rural areas [5] Thisagricultural practice is characterized by a rotation of fieldsrather than of crops by short periods of cropping (one to fiveyears) alternated with long fallow periods (up to twenty ormore years but often as short as six to eight years) and byclearing the fields using slash-and-burn techniques Fields arenormally prepared using a hoe or digging stick the plow only

HindawiInternational Journal of Forestry ResearchVolume 2017 Article ID 6202093 13 pageshttpsdoiorg10115520176202093

2 International Journal of Forestry Research

rarely being employed [6 7] The use of fire serves to clearthe field and enrich the soil with nutrients from the ash of thewoody biomass [8]

This agricultural system has been labeled as the mostserious land use problem and a major driver of deforestationin tropical areas ofAfrica [9 10] It has been however an idealsolution for soil fertility in the humid tropics at least in thepast when the human population density was low and fallowperiods were long enough to restore soil fertility [11 12] Therapid growth of human population in rural areas of Africaand increasing needs for food production have caused theshortening of fallow periods of abandoned fields and peopleto encroach into new areas that have not been subjected toagriculture before [12]The increasing demands of the rapidlygrowing human population have led to land scarcity andthe remaining forests are being increasingly over utilized farbeyond their capacity for regeneration [13] This process asa whole seriously impacts forest biodiversity patterns forexample species richness and composition [14]

TheAngolanMiombo which refers to dry tropical forestsin south-central Africa dominated by tree species of the gen-era Brachystegia Julbernardia and Isoberlinia [15] are poorlyknown in terms of woody species diversity compositionand resilience after abandonment Data on species diversityand composition are confined to sporadic and outdatedstudies (eg Monteiro [16]) Almost nothing is known onthe dynamics of forest recovery following disturbance whichhampers any systematic management approaches aimed atsustainable use of forest resources To obtain a holistic viewon forest resources of the Angolan Miombo it is crucial tounderstand variations in tree species composition to analyzepatterns of species distributions in the forests and to quantifythe relative contribution of different species to overarchingpatterns [17]

In order tomanage disturbed and undisturbed stands andto understand the provision of nontimber ecosystem servicesit is imperative to describe patterns of species composition[13] Additionally data on woody species composition serveas baseline information for projecting changes in vegetationover time which is fundamental to understand regenerationprocesses such as tree growth tree mortality understorydevelopment and the spread of disturbances [18]

The present study was conducted in the Cusseque areaof the Municipality of Chitembo in southern Bie provincewhich is situated in the center of theAngolanMiombo regionThis area is at the current front of deforestation of Africandry tropical forests [19] This study aims to assess the woodyspecies diversity and composition on abandoned agriculturalcrop fields and mature forest stands in the Cusseque area inorder to

(i) analyze changes on woody species composition dur-ing forest recovery

(ii) assess changes of woody species diversity during aslash-and-burn cycle

(iii) understand the temporal dynamics and timelineof forest regeneration from field abandonment tomature forest

2 Material and Methods

21 Study Area The study area is located on the southernslope of the Angolan Central Plateau in the Bie province(136985∘S 170382∘E) and it is one of the research sites ofthe Future Okavango Project in Angola (Figure 1) The meanelevation is 1560mwith amean annual temperature of 204∘C[20 21]

The study area has a tropical subhumid climate witha rainy season of approximately six months lasting fromOctober to April and a dry season corresponding to theremaining period of the year For the period 1971 to 2000the mean annual precipitation was 987mmyear Howeverthe annual rainfall in Cusseque shows a high interannualvariabilitywith a slightly decreasing trend since the 1970s [21]During the dry season frosts frequently occur especially inthe valleys [22]The soils of the study area correspondmainlyto ferralitic soils with a shallow sandy top horizon

The vegetation of the study site is dominated mainly bytypical Miombo woodlands species [23] Brachystegia sppJulbernardia paniculata Cryptosepalum exfoliatum subsppseudotaxus and Erythrophleum africanum are locally domi-nant tree species [16 24ndash28] The woodlands range from 4 to15m in height with sparse to moderate grass or shrub layersbelow the continuous canopy [27]While undisturbed forestsseem to bemostly single stemmedmany stands show signs ofearlier slash-and-burn agriculture with multistemmed treesabove knee height Fire also plays an important role in thecurrent forest dynamics [29] especially during the grassdominated early fallow stages

The human population of the study area mostly belongsto the ethnic group Cokwe Subsistence agriculture basedon shifting cultivation is their main source of livelihoodwith maize (Zea mays) cassava (Manihot esculenta) beans(Phaseolus vulgaris) and peanut (Arachis hypogaea) beingthe main crops Other sources of income include livestockkeeping wage labour and notably in the nearby townof Chitembo the retail of natural resources like charcoalbushmeat and honey [30]

22 Site Selection and Data Collection Forests and fallowswere sampled to represent a chronosequence of forest recov-ery in order to study the state of origin and the dynamicsof regrowth The site selection was done through informalmeetings with traditional authorities and the communityin general aiming to identify sites of former slash-and-burn agriculture and known age since abandonment Theinformation obtained regarding the fallows ages was con-firmed with remote sensing data obtained from LandTrendra programme that detects disturbance within Landsat time-series [32]

The selected areas were then grouped into four cate-gories according to fallow age young fallows (range 7-8 yrs)medium-aged fallows (range 10ndash14 yrs) old fallows (range 20-21 yrs) andmature forests (undisturbed forests orwoodlandsshowing structural features characteristic of later stages ofstand and succession development)

Field work was carried out during March May and June2013 and in September 2014 using 20m times 50m plots that

International Journal of Forestry Research 3

Projection WGS 1984 background RapidEye high resolution satellite imagery recorded May 1 2013 We acknowledge the DLR for the provision of the data from the RapidEye Science Archive

(km)0 8642

N

16∘56

㰀E 17∘0㰀E 17

∘4㰀E 17

∘8㰀E

13∘40

㰀S

13∘44

㰀S

Figure 1 Location of the Okavango basin in southern Africa and the study site in Cusseque (denoted in red) The Okavango basin as shownhere follows the definition of ldquoThe Future Okavango Projectrdquo httpwwwfuture-okavangoorg [31]

were divided into ten 100m2 subplots [33] A total of fortyplots were surveyed corresponding to ten mature standsand ten in each of the fallow stages On every plot werecorded the occurrence of tree species with DBH ge 5 cm Ascomplementary information covers of herbaceous and shrubspecies in a central 100m2 subplot were also recorded

If not already known to the authors the species wereidentified by their local name in Cokwe (given by local fieldguides) collected and taxonomically identified followingfieldwork Voucher specimens are stored at the Herbariumof Lubango (LUBA) and Kew Herbarium (K) The scientificnames and authors of the species follow the checklist ofAngolan vascular plants [34]

23 Data Analysis Diversity refers to different measuresof the species richness and composition Measures includespecies richness and evenness [35] Species richness refers tothe total number of species recorded in each forest stage Inthis study we also calculated the Shannon Diversity Index[36]

1198671015840 = minussum119901119894ln (119901119894) (1)

In the Shannon Diversity Index (1198671015840) 119901119894is the number of

individuals of species in a given plot divided by the totalnumber of individuals in the plot ln is the natural logarithmand sum is the sum of the calculations The index incorporatesthe species richness and the proportion of each species in allsampled plots (evenness) [35 37] The evenness in species

abundance was assessed by Buzas amp Gibsonrsquos Evenness Index[37]

119864 = 119890119867

119878 (2)

In the BuzasampGibsonrsquos Evenness Index (119864)119867 is the ShannonDiversity Index 119890 is the natural logarithm base and 119878 is thetotal number of species Evenness is the equitability of theabundances of different species and the index varies from 0to 1 with 1 indicating that all species have the same abundance[37 38]

To compare the diversity of the four forest stages wecalculated diversity profiles for every stage Diversity pro-files provide a graphical representation of the shape ofa community have the advantage of combining multiplediversity measures (eg species richness Shannon Diversityand Simpsonrsquos diversity) at once and display one curve percategory in this case fallow stage [39ndash41]

This allows for a more complete picture of the multi-dimensional term biodiversity instead of calculating a fewisolated diversity indices only The scale parameter alpha (120572)gives the order of Renyirsquos diversity 120572 = 0 gives the totalnumber of species (species richness) 120572 = 1 gives the indexproportional to the Shannon Diversity Index and 120572 = 2 givesan index which behaves like Simpsonrsquos index while 120572 = Infrefers to the relative abundance of the most abundant species[42]

Stand structure was calculated from stems density in eachfallow and mature forest site A nonparametric test (Kruskal-Wallis) was used to assess differences on stand density


between fallows and mature forest stands Population struc-ture was used to assess dynamics in population for thematurestands Woody species composition was analyzed using theImportanceValues Index (see (3)) which is the summation ofthe relative values of frequency density and dominance [43]The Importance Values Index (IVI) describes the floristicstructure and composition of the woodlands and has beenfrequently used in the Miombo systems [44ndash46]

IVI = (RF + RD + RDo)3

(3)

where IVI is Importance Value Index RF is relative fre-quency RD is relative density and RDo is relative domi-nance

The frequency is a statistical parameter which reflects thespread of a species in a given area The relative frequency(see (4)) of species was obtained from absolute frequencydividing the number of sampling units (100m2 subplots) inwhich the species occurs by the total number of samplingunits

The density of a species reflects the abundance of a speciesin a given community The relative density (see (5)) wasobtained from absolute density calculated from the totalnumber of individual of a species present in a plot dividedby the total area sampled (01 ha)

The dominance is defined as the area occupied by thebasal area of a species per plot (01 ha) The relative domi-nance (see (6)) was obtained by dividing the total basal areafor a given species by the total basal area of all species per plotTo calculate relative frequency density and dominance of thespecies we used standard formulas [47ndash49]

RF = AFTFtimes 100 (4)

where RF is relative frequency of species AF is absolute fre-quency of the species and TF is sum of absolute frequenciesof all species

RD = ADTDtimes 100 (5)

where RD is relative density of species AD is absolutedensity of species (per ha) and TD is total density of trees(per ha)

RDo = ADoTDotimes 100 (6)

where RDo is relative dominance of species ADo is absolutedominance (or basal area) of species and TDo is totaldominance (or basal area) of all species

To explore information on samples and variables(species) we used a Principal Component Analysis biplot[50] A biplot analysis gives graphical representation ofa multivariate sample and it superimpose on the displaya representation of the variables on which the sample ismeasured [51]

Patterns in the species composition of vegetation plotsacross fallow stages were explored using the Principal Coor-dinates Analysis (PCoA) based on the original abundance

matrix data [50]The Principal Coordinates Analysis (PCoA)is an eigen-vector-based method which reduces the dimen-sionality by projecting multidimensional datasets into asmaller number of dimensions [52] allowing a much widerdefinition of dissimilarity than simple Euclidian distancein the species space focusing on the associations betweenindividual observation in a dataset [52 53] Calculations ofdiversity indices evenness diversity profiles Principal Coor-dinatesAnalysis (PCoA) and biplot of Principal ComponentsAnalysis (PCA) were computed in PAST (PaleontologicalStatistics Software) (version 217)

The overlap of the vegetation composition of the four cat-egories was assessed based on an Analysis of Similarity (One-way ANOSIM) Values of 119877ANOSIM range from 0 to 1 Valuesclose to 0 indicate strong overlap [53] while values close to 1indicate complete separation among the groups [54] To illus-trate how many woody species are shared between the fourcategories a Venn diagram was drawn with the Bioinformat-ics amp Evolutionary Genomic Tool (httpbioinformaticspsbugentbewebtoolsVenn)

Additionally Indicator Species Analysis (ISA) was calcu-lated in the R Statistical Analysis Software [55] to identify thediagnostic species of each fallow stage Associations of speciesto more than one fallow stage were allowed [56 57]

3 Results

31 Woody Species Diversity The species richness measuredby the total number of tree species recorded was higherin mature forests (44) and medium-aged fallows (41) beinglower in old (36) and young fallows (31) The overall speciesdiversity as measured with the Shannon Diversity Index(1198671015840) was highest in mature forests (291) followed by youngfallows (281) andmedium-aged fallows (271)Theold fallowsshowed the lowest species diversity (261) The highest valueof evenness (119864) was found in young fallows (053) followedby mature forests (041) and old fallows (037) Medium-agedfallows showed the lowest evenness value (036)

The diversity profiles indicate that mature forests weremore diverse than medium-aged fallows and old fallowsas the graphs of the respective diversity profiles did notintercept and also they feature higher species richness thanyoung fallows (graph higher for low alpha) Only if evenness(high alpha) is considered young fallows have higher valuesthan mature forests (Figure 2)

32 Stand Structure and Species Composition We recorded atotal of 3157 individuals 445 individuals in young fallows760 individuals in medium-aged fallows 965 individuals inold fallows and 987 individuals in mature forests The meanstand density in early regrowth fallows was 45plusmn16 stems haminus1stems ranging from 17 to 69 stems haminus1 In the medium-aged fallows the mean stand density was 76 plusmn 28 stems haminus1ranging from 35 to 125 stems haminus1 Old fallows showed amean density of 97 plusmn 23 stems haminus1 stems ranged from 65 to140 stems haminus1 While the mature forests mean stand densitywas 99 plusmn 48 stems haminus1 ranging from 45 to 191 stems haminus1Stand density was significantly different between the fallowtypes and mature forests (119867 = 1618 119901 = 0001) being


025 05 1 2 4 8 Inf0Alpha

0

1

2

3

H-a

lpha

Figure 2 Diversity profiles using Renyirsquos diversity of three fallowsstages and mature forest stands (mature forests red old fallowsblue medium-aged fallows yellow and young fallows green)

2 4 6 8 10 12 140DBH classes (cm)

050

100150200250300350400450500

No

of s

tem

shaminus1

Figure 3 Size class distribution of mature forests showing a reverseJ shape the numbers represent the DBH classes in cm 2 (10ndash15) 4(20ndash25) 6 (30ndash35) 8 (40ndash45) 10 (50ndash55) 12 (60ndash65) and 14 (ge65)

more evident between early regrowth stages old fallows andmature stands The mean diameter in mature stands was132plusmn88 cm showing reverse J-shaped size class distributionwith the majority of trees found in the smaller diameterclasses accounting for 47 of stems (Figure 3)

These individuals belong to 51 species from 38 generaand 19 families Two species could not be identified andanother two were only identified to genus level (Table 1)The most diverse family was Fabaceae subfamily Caesalpin-ioideae representing (30) of all woody species followedby Strychnaceae (Loganiaceae) and Combretaceae (9)Myrtaceae and Proteaceae (7) Euphorbiaceae (6) andApocynaceae Dipterocarpaceae Euphorbiaceae and Melas-tomataceae (4) Rubiaceae and other smaller families rep-resent less than 2 of species

The structural analysis of woody species represented bythe relative values of frequency density dominance andImportance Values Index (IVI) is summarized in Table 1in alphabetical order of the species The dominant species

AntAgmAbmBpqBmd

BbkBsp

BafCbl

Ccl

Csp

CzhCba

Cef

DngDbt

Dcn

Ebn

Eaf

FrcGclHac

Jpn

MflMaf

Mkt

Ocn

OdtPcrPagPggPptPspPmfPngPanRegSlpStrSccSpgSspSpySzbSzcSzg

TbrUntUkrUsp1Usp2VmdWsp

Com

pone

nt 2

8

16

24

32

40

48

minus16

minus8minus30 minus20 minus10 10 20 30 40 50minus40

Component 1

Figure 4 Biplot of the Principal Components Analysis for thevariables (species) quantified across the plots (young fallows greenmedium fallows yellow old fallows blue and mature forests red)

ranked by their IVI in all categories were Brachystegiaspiciformis Burkea africana Combretum collinum Combre-tum zeyheri Cryptosepalum exfoliatum subsp pseudotaxusDiplorhynchus condylocarpon Erythrophleum africanum Jul-bernardia paniculata Monotes africanus Pericopsis angolen-sis and Terminalia brachystemma

The biplot analysis of PCA explained 60 (39 and 21for the first and second axis resp) of the species variationobserved across the fallows andmature forest plots (Figure 4)The early regrowth stages are characterized by the dominanceof Albizia spp Anisophyllea boehmii Pericopsis angolensisand Combretum species Cryptosepalum exfoliatum subsppseudotaxus and Brachystegia spiciformis dominated themature stands while Erythrophleum africanum Julbernardiapaniculata and Diplorhynchus condylocarpon characterizedthe medium-aged stands and old fallows Strychnos speciesshowed high variation in terms of frequency density anddominance in all stages Two exploitable woody speciesGuibourtia coleosperma and Pterocarpus angolensis showedfew individuals in all fallows stages

33 Principal Coordinates Analysis (PCoA) and Analysis ofSimilarity (ANOSIM) ThePrincipal Coordinates Analysis ofspecies density and composition distinguished four differentgroupsThefirst axis explained 184of variation (Eigenvalue= 19) and the second axis explained 148 of variation(Eigenvalue = 15) The first group was formed by theyoung fallows the second and third groups were formed bymedium-aged and old fallows while the fourth group wasformed by the mature forests (Figure 5)

Theold andmedium fallows showedmany shared speciesas indicated byAnalysis of Similarity (119877ANOSIM = 0052) Onlyfew species are shared between young and medium fallows(119877ANOSIM = 0487) The group formed by mature stands


Table1Standstructureof

woo

dyspeciesandIm

portance

Values

Indexof

four

forestrsquosstagesY

F=youn

gfallo

wsMF=medium

fallo

wsOF=oldfallo

ws

andMW

=mature

forests

woo

dlands

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Albiziaantunesia

namdash

147

mdash043

mdash064

mdash011

mdash018

mdash002

mdash217

mdash055

Albiziagummifera

258

mdashmdash

043

097

mdashmdash

023

043

mdashmdash

003

369

mdashmdash

067

Aniso

phyllea

boehmii

065

245

mdash043

024

090

mdash011

009

168

mdash005

092

391

mdash056

Baphiabequaertii

258

392

471

388

121

167

239

331

057

186

132

298

398

621

754

818

Bobgun

niamadagascarie

nsis

452

294

314

216

362

090

087

091

186

035

053

081

876

396

419

334

Brachyste

giabakeria

na19

414

7419

431

121

039

087

046

040

017

053

052

328

191

523

494

Brachyste

giaspicifo

rmis

581

490

524

431

362

437

380

1906

325

541

692

2740

1051

1108

1134

3251

Burkea

africana

516

490

471

388

628

463

423

354

652

205

309

212

1361

1021

997

813

Chrysophyllum

bangwe

olense

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdash19

0mdash

mdashCo

mbretum

collinu

m645

343

524

216

1304

244

651

126

745

144

694

062

2198

635

1406

362

Combretum

sp(139009)

mdashmdash

mdash388

mdashmdash

mdash17

1mdash

mdashmdash

090

mdashmdash

mdash589

Combretum

zeyh

eri

645

490

471

129

1039

617

651

046

799

1006

1493

019

1950

1443

1619

181

Copaifera

baum

iana

mdashmdash

mdash086

mdashmdash

mdash034

mdashmdash

mdash003

mdashmdash

mdash12

2Cr

yptosepa

lum

exfolia

tum

subsppseudo

taxu

s12

919

6471

431

121

090

141

1530

072

048

064

1076

274

302

634

2319

Dialiu

mengle

rianu

m323

441

314

172

242

219

076

080

316

242

136

070

670

740

435

276

Diospyros

batocana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash014

mdashmdash

mdash114

Diplorhyn

chus

cond

ylocarpo

n645

392

524

388

628

308

1020

605

214

336

845

565

1345

813

1825

1181

Ekebergiabenguelen

sismdash

196

mdashmdash

mdash051

mdashmdash

mdash030

mdashmdash

mdash257

mdashmdash

Erythrophleum

africanu

m645

490

524

431

1329

1195

1182

1347

1863

886

1413

1488

2595

1981

2177

2274

Faurea

rochetiana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash009

mdashmdash

mdash112

Guibourtiacoleo

sperma

258

049

052

259

097

013

011

126

023

004

004

325

362

063

064

493

Hym

enocardiaacida

452

441

471

388

314

398

271

251

675

363

177

097

991

961

801

671

Julberna

rdia

paniculata

581

490

524

431

483

3368

3221

285

372

2583

2700

660

1188

4719

4645

936

Mem

ecylon

flavovirens

mdashmdash

mdash216

mdashmdash

mdash12

6mdash

mdashmdash

117

mdashmdash

mdash380

Mon

otesafricanu

s19

4294

209

431

169

129

076

582

124

144

045

823

404

471

300

1287

Monoteskatangensis

258

245

157

345

242

077

076

126

161

027

033

191

553

331

244

534

Ochna

schw

einfurthiana

mdashmdash

419

345

mdashmdash

119

308

mdashmdash

029

073

mdashmdash

548

677

Oldfieldiadactylophylla

mdash049

209

043

mdash013

054

011

mdash004

030

003

mdash063

274

056

Parin

aricuratellifolia

mdash14

7mdash

259

mdash039

mdash10

3mdash

038

mdash036

mdash19

8mdash

373

Pericopsisan

golensis

129

441

524

mdash048

270

488

mdash334

809

355

mdash289

981

1130

mdashProtea

gaguedi

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

Protea

petio

laris

129

mdash10

5043

048

mdash022

011

034

mdash007

009

189

mdash12

9057

Protea

sp(139072)

mdash19

6mdash

mdashmdash

064

mdashmdash

mdash038

mdashmdash

mdash273

mdashmdash

Pseudolachnosty

lismaproun

eifoliavardekind

tii12

9245

419

302

048

090

152

148

029

220

286

134

187

408

666

495


Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227

Schreberatrichoclada

387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070

Strychnoscocculoides

194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439


Table 2 Indicator species 119901 values and significance stars [lowast] of single and combined groups of fallows stages and mature stands as observedin the Cusseque area

Fallow age Species Ind value p values Sig starsYoung fallows Albizia gummifera 0516 0036 lowastOld fallows Pericopsis angolensis 0808 0001 lowast lowast lowastMedium amp old fallows Julbernardia paniculata 0971 0001 lowast lowast lowastMedium fallows amp mature forests Parinari curatellifolia 0632 003 lowastMedium old fallows amp mature forests Baphia bequaertii 0827 0028 lowast

Medium old amp young fallows Combretum collinum 0919 0002 lowastlowastCombretum zeyheri 0919 0001 lowast lowast lowast

Old fallows amp mature forests Ochna schweinfurthiana 0765 0002 lowastlowast

Mature forests

Cryptosepalum exfoliatum subsp pseudotaxus 0857 0001 lowast lowast lowastBrachystegia bakeriana 0796 0002 lowastlowast

Strychnos cf mitis 0707 0002 lowastlowastWarneckea sapinii 0674 0009 lowastlowastCombretum sp 0632 0005 lowastlowast

Memecylon flavovirens 0548 0048 lowast

Coordinate 1

Coo

rdin

ate 2

016 024 032 040008minus032 minus024 minus016 minus008minus040

minus040

minus032

minus024

minus016

minus008

008

016

024

032

040

Figure 5 Principal Coordinates Analysis (PCoA) showing therelationship between the tree species composition and fallows stagesin the Cusseque study area (young fallows green medium-agedfallows yellow old fallows blue and mature forests red)

is clearly separated from all fallow types The more ampledispersion of the medium fallows in the ordination diagramindicates that they aremore heterogeneous in terms of speciescomposition than the other stages The Analysis of Similarityalso showed slight variations in species composition amongfallows and mature stands

The analysis of the Venn diagram found a total of twentyspecies shared between all categories Mature stands showedten unique species and medium fallows showed seven whiletwo species were found to be unique to old fallows (Figure 6)

34 Indicator Species Analysis (ISA) The analysis of indicatorspecies (Table 2) identified 6 indicator species for the matureforest plots For the fallow stages only a limited number ofspecies were identified as diagnostic Combretum collinum

Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4

Figure 6 Nonsymmetric Venn diagram showing the number ofspecies shared between the forest stages (young fallows greenmedium fallows yellow old fallows blue and mature forests red)

and C zeyheri were indicative of all fallow types Albiziagummiferawas characteristic for young fallows and Pericopsisangolensis for old fallows The Indicator Species Analysisshows that typical Miombo species rarely were indicative offallow types with the exception of Julbernardia paniculata inmedium-aged and old fallows

4 Discussion

41 Woody Species Diversity Species diversity is the mostcommonly used representation of ecological diversity and canbe measured from the number of species (species richness)and relative abundance of individuals within each species(species abundance) [58] We assessed woody species diver-sity in multiple ways including species richness which was


measured from the total number of species occurring indifferent fallows stages and mature forests We observed anincrease in species richness with fallow age However standagemaynot necessarily be themost important determinant offloristic composition and even stand structure Other factorssuch as initial floristic composition and land use history mayneed to be considered further [59 60] The observed speciesrichness values in the early stage of regrowth are comparableto those of mature stands while the species composition didnot completely recover in the observed time frame and maytake several decades to be the same as mature forests [61]

Further diversity measures such as Shannon Diversityand Evenness (Buzas amp Gibsonrsquos) were used to assess thediversity of woody species in the study area The ShannonDiversity Index normally varies from 15 to 35 rarely exceed-ing 45 [48] The overall species diversity values found inour study area are in the range of the values encounteredin other studies in Miombo woodlands of south-centralAfrica indicating that young regrowth and mature forestshad high species diversity while the intermediate stages hadlow species diversity [62]

The species diversity values encountered were consider-ably higher compared to other studies conducted in Miombowoodlands inTanzaniawhere the values foundwere1198671015840 = 1311198671015840 = 132 and1198671015840 = 142 [18] and close to those encountered inZambianMiombo (1198671015840 = 25 in 20 yrs of fallow and1198671015840 = 26 in30 yrs of fallow) [44]The evenness values found were similarin the intermediate stages implying that the individuals ofdifferent species recorded had relatively similar abundancesat both stages Differences in species richness and diversity ofthe forests are reported to be dependent upon the amount ofrainfall that the area receives annually [46]

The high diversity value found in young fallows is a bitunexpected but can be explained by the high environmentalheterogeneity of fallows in early successional stages Youngfallows still host some sprouts from stumps of mature forestspecies while strong disturbances (like slash-and-burn agri-culture) allow light demanding pioneer species to recruit andestablish thus increasing the number of species [63] Withongoing succession some of the original forest remnantsare disappearing in part due to environmental stress andto losing out in light competition with early successionspecies which causes a decline in species diversity in themid succession stages At the same time the light demandingpioneer species are already disappearing in mid succession

Other studies reported similarly that species diversitytended to reach levels close to mature forests relativelyquickly after disturbance by maximizing the coexistence offast growing pioneers and more competitive canopy speciesfollowed by delayed recovery of woody biomass [59 64 65]However the disturbance intensity of agricultural use may betoo high for fast regeneration dynamics The reestablishmentof mature forest species is a slow process which demandslonger time periods than the chronosequence used in ourstudy

42 Species Composition We recorded a total of 3157 indi-viduals of 51 tree species in a total area corresponding to 4 haThe total number of species encountered in our study area

is sharply far from that encountered in Zambian Miombowhere a total of 2761 individuals corresponding to 83 specieswere recorded for mature woodlands [44] and they are inthe range of values reported in Tanzania and Zimbabwe[66 67] despite using different plot size The total numberof individuals found in the early stages falls within the rangeencountered in theMiombowoodlands of Kitulangalo ForestReserve in Tanzania and in Miombo woodlands of Mozam-bique using the same plot size or size thresholds for treeinclusion [68 69] Significant differences found regardingstand density between early regrowth stages and maturestands may be explained by the fact that abandoned sites arecharacterized by a continuous decrease in dominance of earlysuccessional and fire resistant species and an increase in thepresence of slow-growing tree species commonly associatedwith mature forest habitats [64] A reverse J-shaped size classdistribution observed in the mature stands may indicate astable population and good recruitment of late successionalspecies

The woodlands of Cusseque area are dominated by thetypical tree species that characterize Miombo forests allover south-central Africa [70ndash72] The sole exception wasIsoberlinia angolensiswhich was surprisingly not found in thearea although it is cited as a common codominant speciesin many Miombo forests [73ndash75] The dominance of theFabaceae subfamily Caesalpinioideae has also been reportedin various studies [76ndash78]

In general the forest stands showed high variation interms of species composition which may imply that eachspecies has a different ecological importance [79] Strongenvironmental changes caused by slash-and-burn agricul-ture particularly in terms of soil nutrients and microcli-mate are slowly converging to the previous conditions withincrease of fallow age and may also influence the changes inspecies composition

The dominance of Combretum spp often characterizesareas with high land use pressure where the species becomesthe fastest growing and most dominant tree in early suc-cession stages [46] The occurrence of Combretum speciesafter disturbance was also reported in various other studieson early secondary succession of Miombo in south-centralAfrica [80 81] The early stages of regrowth are frequentlycharacterized by a high dominance of fire tolerant generasuch as Burkea Pterocarpus Pseudolachnostylis and Termi-nalia [82] and by light demanding genera such as AlbiziaStrychnos and Uapaca [44] Kwibisa [60] found Brachys-tegia spiciformis and Julbernardia paniculata as dominanttree species among the canopy of Zambian Miombo whilethe understory was dominated by Combretum spp Ochnaschweinfurthiana Pseudolachnostylis maprouneifolia Diplor-hynchus condylocarponTerminalia spp andBurkea africana

Erythrophleum africanum and Pericopsis angolensis wererecorded in all stand stages with the exception of the latterwhich is not found in mature forest stands This can beexplained by the fact that these tree species are frequently leftstanding on fields because of the hardness of their trunksThefew individuals of Guibourtia coleosperma and Pterocarpusangolensis found in the study area can be explained by theimportance of these woody species as timber and commercial


value are associated both species were referred to as themostexploited woody species in the Angolan part of the Okavangobasin at least in the colonial era [83]

The mature forest plots were dominated by BrachystegiaspiciformisBurkea africanaCryptosepalum exfoliatum subsppseudotaxus and Monotes africanus The absence of B spi-ciformis in the grass dominated stages of previously farmedareas may be due to the species being fire sensitive [82] TheMiombo species indicators showed low dominance in theearly stages of regrowth and increased in dominance withage with the exception of J paniculata which was moredominant in the intermediate stages of fallows than inmatureforest stands reinforcing the assumption that the maturestands in the study areas are characterized by the dominanceof C exfoliatum subsp pseudotaxus These findings are alsoin accordance with the biplot diagram displayed from thePrincipal Components Analysis

The dominance of C exfoliatum subsp pseudotaxus mayalso be related to sites which are unsustainable for agriculturetherefore rarely hit by fire according to local knowledgethese areas are normally used for other activities such ashoney production The long-term leaching processes causedby high precipitationmade some of the soils of mature forestsvery acidic with the lowest pH values encountered also in thestudy area [84 85] which may explain the unsuitability ofthese areas for agricultural purposes

The Miombo woodlands demonstrate remarkable capac-ity to return to mature stages after disturbance [72] Naturalrecovery from stumps and resprouts of suppressed saplingscould explain the increasing number of species with ageas observed in our study site and also reported in variousstudies in southern Africa (eg Chidumayo [86]) Thedemand of land for slash-and-burn agriculture means thatmuch of the newly cropped lands quickly degrades andbecomes unsuitable for agriculture Therefore efficiency ofthe agricultural systems needs to be enhanced to improve thelongevity of fields and reduce the consumption of forests byslash-and-burn agriculture

43 Principal Coordinates Similarity and Indicator SpeciesAnalysis ThePrincipal Coordinates Analysis (PCoA) delim-ited four different groups The group formed by the maturestands was consistent with the Analysis of Similarity andthe distinction of this group can be also explained by thedominance of Cryptosepalum exfoliatum subsp pseudotaxusas demonstrated by the Indicator Species Analysis Localsite-specific environmental characteristics such as relieftopography and physicochemical soil properties may playa secondary role as demonstrated in other studies in thewoodlands of southern Africa and savanna biome of SouthAmerica [87 88]

5 Conclusions

TheCusseque study area is characterized by a high diversity oftree species with the number of tree individuals and speciesincreasing with fallow age This trend shows an ongoingrecovery process of the mature forest ecosystems after slash-and-burn agriculture as long as the woodlands are not

subjected to others disturbances The recovery process maytake several decades to reachmature forests status Increasingpressure for land demand andother disturbances increasinglybecome a determining hindrance for forest recovery

Competing Interests

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

Acknowledgments

The authors are grateful to the Germany Federal Ministryof Education and Research (BMBF) for providing financialsupport for this study through the Future Okavango Project(Grant no 01LL0912A) Special thanks are due to the tra-ditional authorities local communities in general and localfield guides in Cusseque study area The authors would likealso to acknowledge the staff of ISCED Huıla for all ofthe support given during the project implementation thestaff at the Herbarium of Lubango and the working groupat Biocentre Klein Flottbek of the University of HamburgSpecial thanks are due to the Remote Sensing Team of theUniversity of Trier for performing the analysis of fallow ageusing LandTrendr

References

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[2] J Clarke W Cavendish and C Coote ldquoRural householdsand miombo woodlands use value and managementrdquo inThe Miombo in Transition Woodlands and Welfare in Africacharpter 5 CIFOR 1996

[3] J M Bila and N Mabjaia ldquoCrescimento e fitossociologia deuma floresta com Colophospermum mopane em MabalaneProvıncia de Gaza Mocambiquerdquo Pesquisa Florestal Brasileiravol 32 no 71 pp 421ndash427 2012

[4] M L Du Preez ldquoSouthern Africarsquos Dryland forest climatechange and the water-energy-food securityrdquo Occasional Paper189 South African Institute of International Affairs SADC2014

[5] N N P Junior R S S Murrieta and C Adams ldquoA agriculturade corte e queima um sistema em transformacaordquo Boletim doMuseu Paraense Emılio Goeldi Ciencias Humanas vol 3 no 2pp 153ndash174 2008

[6] K Pelzer ldquoLand utilization in the humid tropicsrdquo in Proceedingsof the 9th Pacific Science Congress vol 20 November-December1957

[7] E B Haney Jr The Nature of Shifting Cultivation in LatinAmerica The Land Tenure Center University of WisconsinMadison 1968

[8] S Dhakal ldquoAn anthropological perspective on shifting cul-tivation a case study of Khoriya cultivation in the ArunValley of Eastern Nepalrdquo Occasional Papers in Sociology andAnthropology vol 6 pp 93ndash111 2000

[9] T A Aththorick D Setiadi Y Purwanto and E GuhardjldquoVegetation stands structure and aboveground biomass afterthe shifting cultivation practices of Karo People in Leuser


Ecosystem North Sumatrardquo Biodiversitas Journal of BiologicalDiversity vol 13 no 2 pp 92ndash97 2012

[10] T Karthik G G Veeraswami and P K Samal ldquoForest recoveryfollowing shifting cultivation an overview of existing researchrdquoTropical Conservation Science vol 2 no 4 pp 374ndash387 2009

[11] R FWatters Shifting Cultivation in Latin America FAO RomeItaly 1971

[12] L Christanty ldquoShifting cultivation and tropical soils patternsproblems and possible improvementsrdquo in Traditional Agricul-ture in Southeast Asia A Human Ecology Perspective G GMarten Ed West View Press Boulder Colo USA 1968

[13] J Neelo D Teketay K Kashe and W Masamba ldquoStandstructure diversity and regeneration status of woody species inopen and exclosed dry woodlands sites aroundmolapo farmingareas of the Okavango Delta Northeastern Botswanardquo OpenJournal of Forestry vol 5 no 4 pp 313ndash328 2015

[14] E Spencer Shifting Cultivation in Southeast Asia vol 19 ofUniversity of California Publication in Geography University ofCalifornia Press Berkeley Calif USA 1966

[15] G Kowero B M Campbell and U R Sumaila Policies andGovernance Structures inWoodlands of SouthernAfrica CIFORBogor Indonesia 2003

[16] R F R Monteiro Alguns Elementos de Interesse Ecologicoda Flora Lenhosa do Planalto do Bie (Angola) Instituto deInvestigacao Cientıfica de Angola Luanda Angola 1970

[17] M E Durigan Florıstica dinamica e analise proteıca de umaFlorestaOmbrofilaMista emSao Joao doTriunfo-PR [Dissertacaode Mestrado em Manejo Florestal] Universidade Federal doParana Curitiba Brasil 1999

[18] J A Isango ldquoStand structure and tree species compositionof Tanzania miombo woodlands a case study from miombowoodlands of community based forest management in IringaDistrictrdquoWorking Papers of the Finnish Forest Research Instituteno 50 pp 43ndash56 2007

[19] M C Hansen P V Potapov R Moore et al ldquoHigh-resolutionglobal maps of 21st-century forest cover changerdquo Science vol342 no 6160 pp 850ndash853 2013

[20] A Grongroft J Luther-Mosebach L Landschreiber R Rever-mann M Finckh and A Eschenbach ldquoCussequemdashlandscaperdquoBiodiversity amp Ecology vol 5 pp 43ndash44 2013

[21] T Weber ldquoCussequemdashclimaterdquo Biodiversity and Ecology vol 5pp 45ndash46 2013

[22] M Finckh R Revermann and M P M Aidar ldquoClimaterefugees going undergroundmdasha response to Maurin et al(2014)rdquo New Phytologist vol 209 no 3 pp 904ndash909 2016

[23] R Revermann and M Finckh ldquoOkavango basin-vegetationrdquoBiodiversity amp Ecology vol 5 pp 29ndash35 2013

[24] H K A Shaw ldquoThe vegetation of Angolardquo Journal of Ecologyvol 35 no 1-2 pp 23ndash48 1947

[25] L A G Barbosa Carta Fitogeografica de Angola Instituto deInvestigacao Cientıfica de Angola Luanda Angola 1970

[26] C A Diniz Caracterısticas Mesologicas de Angola Descricao eCorrelacao dos Aspecto Fisiograficos dos Solos e da Vegetacao daZonas Agrıcolas Angolanas Missao de Inqueritos Agrıcolas deAngola Nova Lisboa Angola 1973

[27] B J Huntley and E M Matos ldquoBotanical diversity and itsconservation in Angolardquo in Botanical Diversity in SouthernAfrica Strelitzia 1 B J Huntley Ed pp 53ndash74 NationalBotanical Institute Pretoria South Africa 1994

[28] R Revermann F Maiato A Gomes F Lages and M FinckhldquoCusseque-Vegetationrdquo Biodiversity amp Ecology vol 5 pp 59ndash632013

[29] M Stellmes D Frantz M Finckh and R Revermann ldquoOka-vango basin-earth observationrdquo Biodiversity amp Ecology vol 5pp 23ndash27 2013

[30] S Domptail L M Gro120573e B Kolwalski and J BaptistaldquoCussequeCacuchimdashThe Peoplerdquo Biodiversity amp Ecology vol5 pp 73ndash80 2013

[31] J Wehberg and T Weinzierl ldquoOkavango Basinmdashphysicogeo-graphical settingrdquo Biodiversity amp Ecology vol 5 pp 11ndash13 2013

[32] R E Kennedy Z Yang and W B Cohen ldquoDetecting trendsin forest disturbance and recovery using yearly Landsat timeseries 1 LandTrendrmdashtemporal segmentation algorithmsrdquoRemote Sensing of Environment vol 114 no 12 pp 2897ndash29102010

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[36] A E Magurran ldquoMeasuring biological diversityrdquo Journal of theTorrey Botanical Society vol 131 no 3 pp 277ndash278 2004

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[38] M R Kanieski A C B Araujo and S J Longhi ldquoDiversityquantification in Mixed Ombrophilous Forest by differentAlpha Indexesrdquo Scientia Forestalis vol 38 no 88 pp 567ndash5772010

[39] T Leinster and C A Cobbold ldquoMeasuring diversity theimportance of species similarityrdquo Ecology vol 93 no 3 pp 477ndash489 2012

[40] M L Carranza A Acosta and C Ricotta ldquoAnalyzing landscapediversity in time the use of Renyirsquos generalized entropy func-tionrdquo Ecological Indicators vol 7 no 3 pp 505ndash510 2007

[41] J Oldeland N Dreber and D Weduls ldquoDiversity measures incomparative rangeland studies application and advantages ofspecies abundance distributions and diversity profilesrdquo Dinte-ria no 31 pp 50ndash66 2010

[42] Oslash Hammer PAST Paleontological Statistics Version 217 Ref-erence Manual Natural History Museum University of Oslo2012

[43] J T Curtis and R P McIntosh ldquoAn upland forest continuum inthe Prairie-forest border region of Wisconsinrdquo Ecology vol 32no 3 pp 476ndash496 1951

[44] F K Kalaba C H Quinn A J Dougill and R VinyaldquoFloristic composition species diversity and carbon storage incharcoal and agriculture fallows and management implicationsin Miombo woodlands of Zambiardquo Forest Ecology and Manage-ment vol 304 pp 99ndash109 2013

[45] P K T Munishi R A P C Temu and G Soka ldquoPlant com-munities and tree species associations in Miombo ecosystemsin Lake Rukwa basin Southern Tanzania Implications forconservationrdquo Journal of Ecology and the Natural Environmentvol 3 no 2 pp 63ndash71 2011

[46] E K K Jew A J Dougill S M Sallu J OrsquoConnell and TG Benton ldquoMiombo woodland under threat consequences fortree diversity and carbon storagerdquo Forest Ecology and Manage-ment vol 361 pp 144ndash153 2016


[47] D Mueller-Dumbois and H Ellenberg Aims amp Methods ofVegetation Ecology John Wiley amp Sons New York NY USA1974

[48] M Kent and P Coker Vegetation Description and Analysis APractical Approach John Wiley amp Sons 1992

[49] W K Freitas and L M S Magalhaes ldquoMetodos e Parametrospara Estudo da Vegetacao com Enfase no Estrato ArboreordquoFloresta e Ambiente no 19 pp 520ndash540 2012

[50] K R Gabriel ldquoThe biplot graphic display of matrices withapplication to principal component analysisrdquo Biometrika vol58 no 3 pp 453ndash467 1971

[51] F Udina ldquoInteractive biplot constructionrdquo Journal of StatisticalSoftware vol 13 no 5 pp 1ndash16 2005

[52] J W Huntley ldquoCharpter 2 exploratory multivariate techniquesand their utility for understanding ancient ecosystemsrdquo inQuantifying the Evolution of Early Life M Laflamme J DSchiffbauer and S Q Dornbos Eds vol 36 of Topics inGeobiology pp 23ndash48 Springer Dordrecht Netherlands 2011

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[55] RDevelopment Core Team R A Language and Environment forStatistical Computing R Foundation for Statistical ComputingVienna Austria 2015

[56] M De Caceres P Legendre and M Moretti ldquoImprovingindicator species analysis by combining groups of sitesrdquo Oikosvol 119 no 10 pp 1674ndash1684 2010

[57] M Dufrene and P Legendre ldquoSpecies assemblages and indi-cator species the need for a flexible asymmetrical approachrdquoEcological Monographs vol 67 no 3 pp 345ndash366 1997

[58] A J Hamilton ldquoSpecies diversity or biodiversityrdquo Journal ofEnvironmental Management vol 75 no 1 pp 89ndash92 2005

[59] C Uhl ldquoFactors controlling succession following slash-and-burn agriculture in Amazoniardquo Journal of Ecology vol 75 no2 pp 377ndash407 1987

[60] L Kwibisa The effects of indigenous cultivation practices on therecovery of dry Miombo woodland in Central Zambia [MSthesis] University of Zambia 2000

[61] J K K NrsquoDja and G Decocq ldquoSuccessional patterns of plantspecies and community diversity in a semi-deciduous tropicalforest under shifting cultivationrdquo Journal of Vegetation Sciencevol 19 no 6 pp 809ndash820 2008

[62] F C Lumbwe Modeling avifauna responses to Miombo wood-land degradation in Serenje District central province Zambia[PhD thesis] University of Zambia 2010

[63] T Banda M W Schwartz and T Caro ldquoWoody vegetationstructure and composition along a protection gradient in amiombo ecosystem of western Tanzaniardquo Forest Ecology andManagement vol 230 no 1-3 pp 179ndash185 2006

[64] I M McNicol C M Ryan andMWilliams ldquoHow resilient areAfrican woodlands to disturbance from shifting cultivationrdquoEcological Applications vol 25 no 8 pp 2330ndash2336 2015

[65] T H Mwampamba and M W Schwartz ldquoThe effects ofcultivation history on forest recovery in fallows in the EasternArc Mountain Tanzaniardquo Forest Ecology and Management vol261 no 6 pp 1042ndash1052 2011

[66] C J Mafupa Impact of human disturbances in Miombo Wood-lands of Igombe River Forest Reserve Nzega District Tanzania

[MS thesis] Sokoine University of Agriculture MorogoroTanzania 2006

[67] P Gandiwa E Chinoitezvi and E Gandiwa ldquoStructure andcomposition of woody vegetation in two important bird areasin southern Zimbabwerdquo Journal of Animal amp Plant Sciences vol23 no 3 pp 813ndash820 2013

[68] L Mbwambo S Valkonen and V Kuutti ldquoStructure anddynamics of miombo woodland stands at Kitulangalo ForestReserverdquoWorking Papers of the Finnish Forest Research Institutevol 98 pp 10ndash19 2008

[69] N S Ribeiro C N Matos I R Moura R A Washington-Allen and A I Ribeiro ldquoMonitoring vegetation dynamics andcarbon stock density in miombo woodlandsrdquo Carbon Balanceand Management vol 8 article 11 2013

[70] J O Oyugi J S Brown and C J Whelan ldquoEffects of humandisturbance on composition and structure of Brachystegiawoodland in Arabuko-Sokoke Forest Kenyardquo African Journalof Ecology vol 46 no 3 pp 374ndash383 2007

[71] I M Grundy B M Campbell and P G H Frost ldquoSpatial pat-tern regeneration and growth rates of Brachystegia spiciformisand Julbernardia globiflorardquoVegetatio vol 115 no 2 pp 101ndash1071994

[72] DD ShirimaO Totland P K TMunishi and S RMoe ldquoDoesthe abundance of dominant trees affect diversity of awidespreadtropical woodland ecosystem in Tanzaniardquo Journal of TropicalEcology vol 31 no 4 pp 345ndash359 2015

[73] E N Chidumayo ldquoSpecies structure in Zambian miombowoodlandrdquo Journal of Tropical Ecology vol 3 no 2 pp 109ndash1181987

[74] P V Desanker and I C Prentice ldquoMIOMBOmdasha vegetationdynamics model for the miombo woodlands on ZambezianAfricardquo Forest Ecology andManagement vol 69 no 1ndash3 pp 87ndash95 1994

[75] F White The Vegetation of Africa A Descriptive Memoir toAccompany the UNESCOAETFATUNSO Vegetation Map ofAfrica vol 20 of Natural Resources Research UNESCO ParisFrance 1983

[76] E J Francisco C G Africano M Sanfilippo Y G QuintanaI C Martinez and Y A Crespo ldquoEstructura y composiciondel bosque Miombo del Sector Norte de Canjombe AngolardquoRevista Forestal Baracoa vol 33 pp 306ndash316 2014

[77] N Muboko M R Mushonga N Chibwe C Mashapa andE Gandiwa ldquoWoody vegetation structure and compositionin Mapembe Nature Reserve Eastern Zimbabwerdquo Journal ofApplied Sciences and Environmental Management vol 17 no 4pp 475ndash481 2013

[78] E E Mwakalukwa H Meilby and T Treue ldquoFloristic composi-tion structure and species associations of dry Miombo Wood-land in Tanzaniardquo International Journal of Forestry Research vol2014 Article ID 153278 15 pages 2014

[79] H AWale T Bekele and G Dalle ldquoFloristic diversity regener-ation status and vegetation structure of woodlands in MetemaArea AmharaNational Regional State Northwestern EthiopiardquoJournal of Forestry Research vol 23 no 3 pp 391ndash398 2012

[80] P Stromgaard ldquoEarly secondary succession on abandonedshifting cultivatorrsquos plots in the Miombo of South CentralAfricardquo Biotropica vol 18 no 2 pp 97ndash108 1986

[81] I Backeus B Pettersson L Stromquist andCRuffo ldquoTree com-munities and structural dynamics in miombo (Brachystegia-Julbernardia) woodland Tanzaniardquo Forest Ecology andManage-ment vol 230 no 1-3 pp 171ndash178 2006


[82] T RMcClanahan andT P Young EdsEast African Ecosystemsand their Conservation Oxford University Press New York NYUSA 1996

[83] N Baptista ldquoLiterature study of the woody Miombo vegetationand forest management in southeastern Angola with focuson data from the colonial erardquo Polytechnic of Namibia inframework of The Future Okavango Project SP05-Task 6 2014

[84] E N Chidumayo ldquoUsing natural fertilizers in Miombo wood-landsrdquo Issues in African Biodiversity-The Biodiversity SupportProgram no 2 pp 1ndash8 1999

[85] A Grongroft ldquoCussequemdashsoilsrdquo Biodiversity and Ecology vol5 pp 51ndash54 2013

[86] E N Chidumayo ldquoDevelopment of Brachystegia-Julbernardiawoodland after clear-felling in central Zambia evidence forhigh resiliencerdquoApplied Vegetation Science vol 7 no 2 pp 237ndash242 2004

[87] S Syampungani Vegetation change analysis and ecologicalrecovery of the Copperbelt Miombo woodlands of Zambia [PhDthesis] Stellenbosch University Stellenbosch South Africa2009

[88] E Lenza J O Santos and L Maracahipes-Santos ldquoSpeciescomposition diversity and vegetation structure in a galleryforest-cerrado sensu stricto transition zone in eastern MatoGrosso Brazilrdquo Acta Botanica Brasilica vol 29 no 3 pp 327ndash338 2015

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of



EcosystemsJournal of


MeteorologyAdvances in

EcologyInternational Journal of


Marine BiologyJournal of


Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

Advances in


Environmental Chemistry

Atmospheric SciencesInternational Journal of



Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 2014

International Journal of

Geophysics


Geological ResearchJournal of

EarthquakesJournal of


BiodiversityInternational Journal of


ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of


The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014


ClimatologyJournal of


rarely being employed [6 7] The use of fire serves to clearthe field and enrich the soil with nutrients from the ash of thewoody biomass [8]

This agricultural system has been labeled as the mostserious land use problem and a major driver of deforestationin tropical areas ofAfrica [9 10] It has been however an idealsolution for soil fertility in the humid tropics at least in thepast when the human population density was low and fallowperiods were long enough to restore soil fertility [11 12] Therapid growth of human population in rural areas of Africaand increasing needs for food production have caused theshortening of fallow periods of abandoned fields and peopleto encroach into new areas that have not been subjected toagriculture before [12]The increasing demands of the rapidlygrowing human population have led to land scarcity andthe remaining forests are being increasingly over utilized farbeyond their capacity for regeneration [13] This process asa whole seriously impacts forest biodiversity patterns forexample species richness and composition [14]

TheAngolanMiombo which refers to dry tropical forestsin south-central Africa dominated by tree species of the gen-era Brachystegia Julbernardia and Isoberlinia [15] are poorlyknown in terms of woody species diversity compositionand resilience after abandonment Data on species diversityand composition are confined to sporadic and outdatedstudies (eg Monteiro [16]) Almost nothing is known onthe dynamics of forest recovery following disturbance whichhampers any systematic management approaches aimed atsustainable use of forest resources To obtain a holistic viewon forest resources of the Angolan Miombo it is crucial tounderstand variations in tree species composition to analyzepatterns of species distributions in the forests and to quantifythe relative contribution of different species to overarchingpatterns [17]

In order tomanage disturbed and undisturbed stands andto understand the provision of nontimber ecosystem servicesit is imperative to describe patterns of species composition[13] Additionally data on woody species composition serveas baseline information for projecting changes in vegetationover time which is fundamental to understand regenerationprocesses such as tree growth tree mortality understorydevelopment and the spread of disturbances [18]

The present study was conducted in the Cusseque areaof the Municipality of Chitembo in southern Bie provincewhich is situated in the center of theAngolanMiombo regionThis area is at the current front of deforestation of Africandry tropical forests [19] This study aims to assess the woodyspecies diversity and composition on abandoned agriculturalcrop fields and mature forest stands in the Cusseque area inorder to

(i) analyze changes on woody species composition dur-ing forest recovery

(ii) assess changes of woody species diversity during aslash-and-burn cycle

(iii) understand the temporal dynamics and timelineof forest regeneration from field abandonment tomature forest

2 Material and Methods

21 Study Area The study area is located on the southernslope of the Angolan Central Plateau in the Bie province(136985∘S 170382∘E) and it is one of the research sites ofthe Future Okavango Project in Angola (Figure 1) The meanelevation is 1560mwith amean annual temperature of 204∘C[20 21]

The study area has a tropical subhumid climate witha rainy season of approximately six months lasting fromOctober to April and a dry season corresponding to theremaining period of the year For the period 1971 to 2000the mean annual precipitation was 987mmyear Howeverthe annual rainfall in Cusseque shows a high interannualvariabilitywith a slightly decreasing trend since the 1970s [21]During the dry season frosts frequently occur especially inthe valleys [22]The soils of the study area correspondmainlyto ferralitic soils with a shallow sandy top horizon

The vegetation of the study site is dominated mainly bytypical Miombo woodlands species [23] Brachystegia sppJulbernardia paniculata Cryptosepalum exfoliatum subsppseudotaxus and Erythrophleum africanum are locally domi-nant tree species [16 24ndash28] The woodlands range from 4 to15m in height with sparse to moderate grass or shrub layersbelow the continuous canopy [27]While undisturbed forestsseem to bemostly single stemmedmany stands show signs ofearlier slash-and-burn agriculture with multistemmed treesabove knee height Fire also plays an important role in thecurrent forest dynamics [29] especially during the grassdominated early fallow stages

The human population of the study area mostly belongsto the ethnic group Cokwe Subsistence agriculture basedon shifting cultivation is their main source of livelihoodwith maize (Zea mays) cassava (Manihot esculenta) beans(Phaseolus vulgaris) and peanut (Arachis hypogaea) beingthe main crops Other sources of income include livestockkeeping wage labour and notably in the nearby townof Chitembo the retail of natural resources like charcoalbushmeat and honey [30]

22 Site Selection and Data Collection Forests and fallowswere sampled to represent a chronosequence of forest recov-ery in order to study the state of origin and the dynamicsof regrowth The site selection was done through informalmeetings with traditional authorities and the communityin general aiming to identify sites of former slash-and-burn agriculture and known age since abandonment Theinformation obtained regarding the fallows ages was con-firmed with remote sensing data obtained from LandTrendra programme that detects disturbance within Landsat time-series [32]

The selected areas were then grouped into four cate-gories according to fallow age young fallows (range 7-8 yrs)medium-aged fallows (range 10ndash14 yrs) old fallows (range 20-21 yrs) andmature forests (undisturbed forests orwoodlandsshowing structural features characteristic of later stages ofstand and succession development)

Field work was carried out during March May and June2013 and in September 2014 using 20m times 50m plots that



(km)0 8642

N

16∘56

㰀E 17∘0㰀E 17

∘4㰀E 17

∘8㰀E

13∘40

㰀S

13∘44

㰀S





1198671015840 = minussum119901119894ln (119901119894) (1)




119864 = 119890119867

119878 (2)








(3)
















3 Results





025 05 1 2 4 8 Inf0Alpha

0

1

2

3

H-a

lpha


2 4 6 8 10 12 140DBH classes (cm)

050

100150200250300350400450500

No

of s

tem

shaminus1





AntAgmAbmBpqBmd

BbkBsp

BafCbl

Ccl

Csp

CzhCba

Cef

DngDbt

Dcn

Ebn

Eaf

FrcGclHac

Jpn

MflMaf

Mkt

Ocn



Com

pone

nt 2

8

16

24

32

40

48

minus16


Component 1








woo

dyspeciesandIm

portance

Values

Indexof

four

forestrsquosstagesY

F=youn

gfallo

wsMF=medium

fallo

wsOF=oldfallo

ws

andMW

=mature

forests

woo

dlands

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Albiziaantunesia

namdash

147

mdash043

mdash064

mdash011

mdash018

mdash002

mdash217

mdash055

Albiziagummifera

258

mdashmdash

043

097

mdashmdash

023

043

mdashmdash

003

369

mdashmdash

067

Aniso

phyllea

boehmii

065

245

mdash043

024

090

mdash011

009

168

mdash005

092

391

mdash056

Baphiabequaertii

258

392

471

388

121

167

239

331

057

186

132

298

398

621

754

818

Bobgun

niamadagascarie

nsis

452

294

314

216

362

090

087

091

186

035

053

081

876

396

419

334

Brachyste

giabakeria

na19

414

7419

431

121

039

087

046

040

017

053

052

328

191

523

494

Brachyste

giaspicifo

rmis

581

490

524

431

362

437

380

1906

325

541

692

2740

1051

1108

1134

3251

Burkea

africana

516

490

471

388

628

463

423

354

652

205

309

212

1361

1021

997

813

Chrysophyllum

bangwe

olense

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdash19

0mdash

mdashCo

mbretum

collinu

m645

343

524

216

1304

244

651

126

745

144

694

062

2198

635

1406

362

Combretum

sp(139009)

mdashmdash

mdash388

mdashmdash

mdash17

1mdash

mdashmdash

090

mdashmdash

mdash589

Combretum

zeyh

eri

645

490

471

129

1039

617

651

046

799

1006

1493

019

1950

1443

1619

181

Copaifera

baum

iana

mdashmdash

mdash086

mdashmdash

mdash034

mdashmdash

mdash003

mdashmdash

mdash12

2Cr

yptosepa

lum

exfolia

tum

subsppseudo

taxu

s12

919

6471

431

121

090

141

1530

072

048

064

1076

274

302

634

2319

Dialiu

mengle

rianu

m323

441

314

172

242

219

076

080

316

242

136

070

670

740

435

276

Diospyros

batocana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash014

mdashmdash

mdash114

Diplorhyn

chus

cond

ylocarpo

n645

392

524

388

628

308

1020

605

214

336

845

565

1345

813

1825

1181

Ekebergiabenguelen

sismdash

196

mdashmdash

mdash051

mdashmdash

mdash030

mdashmdash

mdash257

mdashmdash

Erythrophleum

africanu

m645

490

524

431

1329

1195

1182

1347

1863

886

1413

1488

2595

1981

2177

2274

Faurea

rochetiana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash009

mdashmdash

mdash112

Guibourtiacoleo

sperma

258

049

052

259

097

013

011

126

023

004

004

325

362

063

064

493

Hym

enocardiaacida

452

441

471

388

314

398

271

251

675

363

177

097

991

961

801

671

Julberna

rdia

paniculata

581

490

524

431

483

3368

3221

285

372

2583

2700

660

1188

4719

4645

936

Mem

ecylon

flavovirens

mdashmdash

mdash216

mdashmdash

mdash12

6mdash

mdashmdash

117

mdashmdash

mdash380

Mon

otesafricanu

s19

4294

209

431

169

129

076

582

124

144

045

823

404

471

300

1287

Monoteskatangensis

258

245

157

345

242

077

076

126

161

027

033

191

553

331

244

534

Ochna

schw

einfurthiana

mdashmdash

419

345

mdashmdash

119

308

mdashmdash

029

073

mdashmdash

548

677


mdash049

209

043

mdash013

054

011

mdash004

030

003

mdash063

274

056

Parin

aricuratellifolia

mdash14

7mdash

259

mdash039

mdash10

3mdash

038

mdash036

mdash19

8mdash

373

Pericopsisan

golensis

129

441

524

mdash048

270

488

mdash334

809

355

mdash289

981

1130

mdashProtea

gaguedi

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

Protea

petio

laris

129

mdash10

5043

048

mdash022

011

034

mdash007

009

189

mdash12

9057

Protea

sp(139072)

mdash19

6mdash

mdashmdash

064

mdashmdash

mdash038

mdashmdash

mdash273

mdashmdash

Pseudolachnosty

lismaproun

eifoliavardekind

tii12

9245

419

302

048

090

152

148

029

220

286

134

187

408

666

495


Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227


387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070


194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439






Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics
















(km)0 8642

N

16∘56

㰀E 17∘0㰀E 17

∘4㰀E 17

∘8㰀E

13∘40

㰀S

13∘44

㰀S





1198671015840 = minussum119901119894ln (119901119894) (1)




119864 = 119890119867

119878 (2)








(3)
















3 Results





025 05 1 2 4 8 Inf0Alpha

0

1

2

3

H-a

lpha


2 4 6 8 10 12 140DBH classes (cm)

050

100150200250300350400450500

No

of s

tem

shaminus1





AntAgmAbmBpqBmd

BbkBsp

BafCbl

Ccl

Csp

CzhCba

Cef

DngDbt

Dcn

Ebn

Eaf

FrcGclHac

Jpn

MflMaf

Mkt

Ocn



Com

pone

nt 2

8

16

24

32

40

48

minus16


Component 1








woo

dyspeciesandIm

portance

Values

Indexof

four

forestrsquosstagesY

F=youn

gfallo

wsMF=medium

fallo

wsOF=oldfallo

ws

andMW

=mature

forests

woo

dlands

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Albiziaantunesia

namdash

147

mdash043

mdash064

mdash011

mdash018

mdash002

mdash217

mdash055

Albiziagummifera

258

mdashmdash

043

097

mdashmdash

023

043

mdashmdash

003

369

mdashmdash

067

Aniso

phyllea

boehmii

065

245

mdash043

024

090

mdash011

009

168

mdash005

092

391

mdash056

Baphiabequaertii

258

392

471

388

121

167

239

331

057

186

132

298

398

621

754

818

Bobgun

niamadagascarie

nsis

452

294

314

216

362

090

087

091

186

035

053

081

876

396

419

334

Brachyste

giabakeria

na19

414

7419

431

121

039

087

046

040

017

053

052

328

191

523

494

Brachyste

giaspicifo

rmis

581

490

524

431

362

437

380

1906

325

541

692

2740

1051

1108

1134

3251

Burkea

africana

516

490

471

388

628

463

423

354

652

205

309

212

1361

1021

997

813

Chrysophyllum

bangwe

olense

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdash19

0mdash

mdashCo

mbretum

collinu

m645

343

524

216

1304

244

651

126

745

144

694

062

2198

635

1406

362

Combretum

sp(139009)

mdashmdash

mdash388

mdashmdash

mdash17

1mdash

mdashmdash

090

mdashmdash

mdash589

Combretum

zeyh

eri

645

490

471

129

1039

617

651

046

799

1006

1493

019

1950

1443

1619

181

Copaifera

baum

iana

mdashmdash

mdash086

mdashmdash

mdash034

mdashmdash

mdash003

mdashmdash

mdash12

2Cr

yptosepa

lum

exfolia

tum

subsppseudo

taxu

s12

919

6471

431

121

090

141

1530

072

048

064

1076

274

302

634

2319

Dialiu

mengle

rianu

m323

441

314

172

242

219

076

080

316

242

136

070

670

740

435

276

Diospyros

batocana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash014

mdashmdash

mdash114

Diplorhyn

chus

cond

ylocarpo

n645

392

524

388

628

308

1020

605

214

336

845

565

1345

813

1825

1181

Ekebergiabenguelen

sismdash

196

mdashmdash

mdash051

mdashmdash

mdash030

mdashmdash

mdash257

mdashmdash

Erythrophleum

africanu

m645

490

524

431

1329

1195

1182

1347

1863

886

1413

1488

2595

1981

2177

2274

Faurea

rochetiana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash009

mdashmdash

mdash112

Guibourtiacoleo

sperma

258

049

052

259

097

013

011

126

023

004

004

325

362

063

064

493

Hym

enocardiaacida

452

441

471

388

314

398

271

251

675

363

177

097

991

961

801

671

Julberna

rdia

paniculata

581

490

524

431

483

3368

3221

285

372

2583

2700

660

1188

4719

4645

936

Mem

ecylon

flavovirens

mdashmdash

mdash216

mdashmdash

mdash12

6mdash

mdashmdash

117

mdashmdash

mdash380

Mon

otesafricanu

s19

4294

209

431

169

129

076

582

124

144

045

823

404

471

300

1287

Monoteskatangensis

258

245

157

345

242

077

076

126

161

027

033

191

553

331

244

534

Ochna

schw

einfurthiana

mdashmdash

419

345

mdashmdash

119

308

mdashmdash

029

073

mdashmdash

548

677


mdash049

209

043

mdash013

054

011

mdash004

030

003

mdash063

274

056

Parin

aricuratellifolia

mdash14

7mdash

259

mdash039

mdash10

3mdash

038

mdash036

mdash19

8mdash

373

Pericopsisan

golensis

129

441

524

mdash048

270

488

mdash334

809

355

mdash289

981

1130

mdashProtea

gaguedi

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

Protea

petio

laris

129

mdash10

5043

048

mdash022

011

034

mdash007

009

189

mdash12

9057

Protea

sp(139072)

mdash19

6mdash

mdashmdash

064

mdashmdash

mdash038

mdashmdash

mdash273

mdashmdash

Pseudolachnosty

lismaproun

eifoliavardekind

tii12

9245

419

302

048

090

152

148

029

220

286

134

187

408

666

495


Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227


387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070


194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439






Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics

















(3)
















3 Results





025 05 1 2 4 8 Inf0Alpha

0

1

2

3

H-a

lpha


2 4 6 8 10 12 140DBH classes (cm)

050

100150200250300350400450500

No

of s

tem

shaminus1





AntAgmAbmBpqBmd

BbkBsp

BafCbl

Ccl

Csp

CzhCba

Cef

DngDbt

Dcn

Ebn

Eaf

FrcGclHac

Jpn

MflMaf

Mkt

Ocn



Com

pone

nt 2

8

16

24

32

40

48

minus16


Component 1








woo

dyspeciesandIm

portance

Values

Indexof

four

forestrsquosstagesY

F=youn

gfallo

wsMF=medium

fallo

wsOF=oldfallo

ws

andMW

=mature

forests

woo

dlands

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Albiziaantunesia

namdash

147

mdash043

mdash064

mdash011

mdash018

mdash002

mdash217

mdash055

Albiziagummifera

258

mdashmdash

043

097

mdashmdash

023

043

mdashmdash

003

369

mdashmdash

067

Aniso

phyllea

boehmii

065

245

mdash043

024

090

mdash011

009

168

mdash005

092

391

mdash056

Baphiabequaertii

258

392

471

388

121

167

239

331

057

186

132

298

398

621

754

818

Bobgun

niamadagascarie

nsis

452

294

314

216

362

090

087

091

186

035

053

081

876

396

419

334

Brachyste

giabakeria

na19

414

7419

431

121

039

087

046

040

017

053

052

328

191

523

494

Brachyste

giaspicifo

rmis

581

490

524

431

362

437

380

1906

325

541

692

2740

1051

1108

1134

3251

Burkea

africana

516

490

471

388

628

463

423

354

652

205

309

212

1361

1021

997

813

Chrysophyllum

bangwe

olense

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdash19

0mdash

mdashCo

mbretum

collinu

m645

343

524

216

1304

244

651

126

745

144

694

062

2198

635

1406

362

Combretum

sp(139009)

mdashmdash

mdash388

mdashmdash

mdash17

1mdash

mdashmdash

090

mdashmdash

mdash589

Combretum

zeyh

eri

645

490

471

129

1039

617

651

046

799

1006

1493

019

1950

1443

1619

181

Copaifera

baum

iana

mdashmdash

mdash086

mdashmdash

mdash034

mdashmdash

mdash003

mdashmdash

mdash12

2Cr

yptosepa

lum

exfolia

tum

subsppseudo

taxu

s12

919

6471

431

121

090

141

1530

072

048

064

1076

274

302

634

2319

Dialiu

mengle

rianu

m323

441

314

172

242

219

076

080

316

242

136

070

670

740

435

276

Diospyros

batocana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash014

mdashmdash

mdash114

Diplorhyn

chus

cond

ylocarpo

n645

392

524

388

628

308

1020

605

214

336

845

565

1345

813

1825

1181

Ekebergiabenguelen

sismdash

196

mdashmdash

mdash051

mdashmdash

mdash030

mdashmdash

mdash257

mdashmdash

Erythrophleum

africanu

m645

490

524

431

1329

1195

1182

1347

1863

886

1413

1488

2595

1981

2177

2274

Faurea

rochetiana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash009

mdashmdash

mdash112

Guibourtiacoleo

sperma

258

049

052

259

097

013

011

126

023

004

004

325

362

063

064

493

Hym

enocardiaacida

452

441

471

388

314

398

271

251

675

363

177

097

991

961

801

671

Julberna

rdia

paniculata

581

490

524

431

483

3368

3221

285

372

2583

2700

660

1188

4719

4645

936

Mem

ecylon

flavovirens

mdashmdash

mdash216

mdashmdash

mdash12

6mdash

mdashmdash

117

mdashmdash

mdash380

Mon

otesafricanu

s19

4294

209

431

169

129

076

582

124

144

045

823

404

471

300

1287

Monoteskatangensis

258

245

157

345

242

077

076

126

161

027

033

191

553

331

244

534

Ochna

schw

einfurthiana

mdashmdash

419

345

mdashmdash

119

308

mdashmdash

029

073

mdashmdash

548

677


mdash049

209

043

mdash013

054

011

mdash004

030

003

mdash063

274

056

Parin

aricuratellifolia

mdash14

7mdash

259

mdash039

mdash10

3mdash

038

mdash036

mdash19

8mdash

373

Pericopsisan

golensis

129

441

524

mdash048

270

488

mdash334

809

355

mdash289

981

1130

mdashProtea

gaguedi

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

Protea

petio

laris

129

mdash10

5043

048

mdash022

011

034

mdash007

009

189

mdash12

9057

Protea

sp(139072)

mdash19

6mdash

mdashmdash

064

mdashmdash

mdash038

mdashmdash

mdash273

mdashmdash

Pseudolachnosty

lismaproun

eifoliavardekind

tii12

9245

419

302

048

090

152

148

029

220

286

134

187

408

666

495


Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227


387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070


194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439






Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics















025 05 1 2 4 8 Inf0Alpha

0

1

2

3

H-a

lpha


2 4 6 8 10 12 140DBH classes (cm)

050

100150200250300350400450500

No

of s

tem

shaminus1





AntAgmAbmBpqBmd

BbkBsp

BafCbl

Ccl

Csp

CzhCba

Cef

DngDbt

Dcn

Ebn

Eaf

FrcGclHac

Jpn

MflMaf

Mkt

Ocn



Com

pone

nt 2

8

16

24

32

40

48

minus16


Component 1








woo

dyspeciesandIm

portance

Values

Indexof

four

forestrsquosstagesY

F=youn

gfallo

wsMF=medium

fallo

wsOF=oldfallo

ws

andMW

=mature

forests

woo

dlands

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Albiziaantunesia

namdash

147

mdash043

mdash064

mdash011

mdash018

mdash002

mdash217

mdash055

Albiziagummifera

258

mdashmdash

043

097

mdashmdash

023

043

mdashmdash

003

369

mdashmdash

067

Aniso

phyllea

boehmii

065

245

mdash043

024

090

mdash011

009

168

mdash005

092

391

mdash056

Baphiabequaertii

258

392

471

388

121

167

239

331

057

186

132

298

398

621

754

818

Bobgun

niamadagascarie

nsis

452

294

314

216

362

090

087

091

186

035

053

081

876

396

419

334

Brachyste

giabakeria

na19

414

7419

431

121

039

087

046

040

017

053

052

328

191

523

494

Brachyste

giaspicifo

rmis

581

490

524

431

362

437

380

1906

325

541

692

2740

1051

1108

1134

3251

Burkea

africana

516

490

471

388

628

463

423

354

652

205

309

212

1361

1021

997

813

Chrysophyllum

bangwe

olense

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdash19

0mdash

mdashCo

mbretum

collinu

m645

343

524

216

1304

244

651

126

745

144

694

062

2198

635

1406

362

Combretum

sp(139009)

mdashmdash

mdash388

mdashmdash

mdash17

1mdash

mdashmdash

090

mdashmdash

mdash589

Combretum

zeyh

eri

645

490

471

129

1039

617

651

046

799

1006

1493

019

1950

1443

1619

181

Copaifera

baum

iana

mdashmdash

mdash086

mdashmdash

mdash034

mdashmdash

mdash003

mdashmdash

mdash12

2Cr

yptosepa

lum

exfolia

tum

subsppseudo

taxu

s12

919

6471

431

121

090

141

1530

072

048

064

1076

274

302

634

2319

Dialiu

mengle

rianu

m323

441

314

172

242

219

076

080

316

242

136

070

670

740

435

276

Diospyros

batocana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash014

mdashmdash

mdash114

Diplorhyn

chus

cond

ylocarpo

n645

392

524

388

628

308

1020

605

214

336

845

565

1345

813

1825

1181

Ekebergiabenguelen

sismdash

196

mdashmdash

mdash051

mdashmdash

mdash030

mdashmdash

mdash257

mdashmdash

Erythrophleum

africanu

m645

490

524

431

1329

1195

1182

1347

1863

886

1413

1488

2595

1981

2177

2274

Faurea

rochetiana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash009

mdashmdash

mdash112

Guibourtiacoleo

sperma

258

049

052

259

097

013

011

126

023

004

004

325

362

063

064

493

Hym

enocardiaacida

452

441

471

388

314

398

271

251

675

363

177

097

991

961

801

671

Julberna

rdia

paniculata

581

490

524

431

483

3368

3221

285

372

2583

2700

660

1188

4719

4645

936

Mem

ecylon

flavovirens

mdashmdash

mdash216

mdashmdash

mdash12

6mdash

mdashmdash

117

mdashmdash

mdash380

Mon

otesafricanu

s19

4294

209

431

169

129

076

582

124

144

045

823

404

471

300

1287

Monoteskatangensis

258

245

157

345

242

077

076

126

161

027

033

191

553

331

244

534

Ochna

schw

einfurthiana

mdashmdash

419

345

mdashmdash

119

308

mdashmdash

029

073

mdashmdash

548

677


mdash049

209

043

mdash013

054

011

mdash004

030

003

mdash063

274

056

Parin

aricuratellifolia

mdash14

7mdash

259

mdash039

mdash10

3mdash

038

mdash036

mdash19

8mdash

373

Pericopsisan

golensis

129

441

524

mdash048

270

488

mdash334

809

355

mdash289

981

1130

mdashProtea

gaguedi

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

Protea

petio

laris

129

mdash10

5043

048

mdash022

011

034

mdash007

009

189

mdash12

9057

Protea

sp(139072)

mdash19

6mdash

mdashmdash

064

mdashmdash

mdash038

mdashmdash

mdash273

mdashmdash

Pseudolachnosty

lismaproun

eifoliavardekind

tii12

9245

419

302

048

090

152

148

029

220

286

134

187

408

666

495


Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227


387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070


194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439






Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics
















woo

dyspeciesandIm

portance

Values

Indexof

four

forestrsquosstagesY

F=youn

gfallo

wsMF=medium

fallo

wsOF=oldfallo

ws

andMW

=mature

forests

woo

dlands

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Albiziaantunesia

namdash

147

mdash043

mdash064

mdash011

mdash018

mdash002

mdash217

mdash055

Albiziagummifera

258

mdashmdash

043

097

mdashmdash

023

043

mdashmdash

003

369

mdashmdash

067

Aniso

phyllea

boehmii

065

245

mdash043

024

090

mdash011

009

168

mdash005

092

391

mdash056

Baphiabequaertii

258

392

471

388

121

167

239

331

057

186

132

298

398

621

754

818

Bobgun

niamadagascarie

nsis

452

294

314

216

362

090

087

091

186

035

053

081

876

396

419

334

Brachyste

giabakeria

na19

414

7419

431

121

039

087

046

040

017

053

052

328

191

523

494

Brachyste

giaspicifo

rmis

581

490

524

431

362

437

380

1906

325

541

692

2740

1051

1108

1134

3251

Burkea

africana

516

490

471

388

628

463

423

354

652

205

309

212

1361

1021

997

813

Chrysophyllum

bangwe

olense

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdash19

0mdash

mdashCo

mbretum

collinu

m645

343

524

216

1304

244

651

126

745

144

694

062

2198

635

1406

362

Combretum

sp(139009)

mdashmdash

mdash388

mdashmdash

mdash17

1mdash

mdashmdash

090

mdashmdash

mdash589

Combretum

zeyh

eri

645

490

471

129

1039

617

651

046

799

1006

1493

019

1950

1443

1619

181

Copaifera

baum

iana

mdashmdash

mdash086

mdashmdash

mdash034

mdashmdash

mdash003

mdashmdash

mdash12

2Cr

yptosepa

lum

exfolia

tum

subsppseudo

taxu

s12

919

6471

431

121

090

141

1530

072

048

064

1076

274

302

634

2319

Dialiu

mengle

rianu

m323

441

314

172

242

219

076

080

316

242

136

070

670

740

435

276

Diospyros

batocana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash014

mdashmdash

mdash114

Diplorhyn

chus

cond

ylocarpo

n645

392

524

388

628

308

1020

605

214

336

845

565

1345

813

1825

1181

Ekebergiabenguelen

sismdash

196

mdashmdash

mdash051

mdashmdash

mdash030

mdashmdash

mdash257

mdashmdash

Erythrophleum

africanu

m645

490

524

431

1329

1195

1182

1347

1863

886

1413

1488

2595

1981

2177

2274

Faurea

rochetiana

mdashmdash

mdash086

mdashmdash

mdash023

mdashmdash

mdash009

mdashmdash

mdash112

Guibourtiacoleo

sperma

258

049

052

259

097

013

011

126

023

004

004

325

362

063

064

493

Hym

enocardiaacida

452

441

471

388

314

398

271

251

675

363

177

097

991

961

801

671

Julberna

rdia

paniculata

581

490

524

431

483

3368

3221

285

372

2583

2700

660

1188

4719

4645

936

Mem

ecylon

flavovirens

mdashmdash

mdash216

mdashmdash

mdash12

6mdash

mdashmdash

117

mdashmdash

mdash380

Mon

otesafricanu

s19

4294

209

431

169

129

076

582

124

144

045

823

404

471

300

1287

Monoteskatangensis

258

245

157

345

242

077

076

126

161

027

033

191

553

331

244

534

Ochna

schw

einfurthiana

mdashmdash

419

345

mdashmdash

119

308

mdashmdash

029

073

mdashmdash

548

677


mdash049

209

043

mdash013

054

011

mdash004

030

003

mdash063

274

056

Parin

aricuratellifolia

mdash14

7mdash

259

mdash039

mdash10

3mdash

038

mdash036

mdash19

8mdash

373

Pericopsisan

golensis

129

441

524

mdash048

270

488

mdash334

809

355

mdash289

981

1130

mdashProtea

gaguedi

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

mdashmdash

Protea

petio

laris

129

mdash10

5043

048

mdash022

011

034

mdash007

009

189

mdash12

9057

Protea

sp(139072)

mdash19

6mdash

mdashmdash

064

mdashmdash

mdash038

mdashmdash

mdash273

mdashmdash

Pseudolachnosty

lismaproun

eifoliavardekind

tii12

9245

419

302

048

090

152

148

029

220

286

134

187

408

666

495


Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227


387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070


194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439






Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics















Table1Con

tinued

Species

Relativefrequ

ency

Relatived

ensity

Relatived

ominance

Impo

rtance

ValueInd

exYF

MF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

YFMF

OF

MW

Ptele

opsis

aniso

ptera

065

049

052

302

024

013

011

160

007

002

001

082

091

063

064

489

Pterocarpu

sangolensis

387

147

209

302

217

051

108

194

057

011

152

104

624

202

368

530

Rothmanniaengle

riana

mdash049

052

172

mdash013

011

046

mdash002

002

027

mdash063

064

227


387

294

157

086

145

103

033

034

051

075

006

013

549

422

191

125

Securid

acalongepedun

culata

065

098

262

043

024

039

065

023

004

017

057

011

090

142

346

070


194

mdash209

129

097

mdash043

034

031

mdash030

016

301

mdash263

169

Strychnosp

ungens

mdash049

mdashmdash

mdash013

mdashmdash

mdash054

mdashmdash

mdash080

mdashmdash

Strychnosm

itis

mdashmdash

mdash17

2mdash

mdashmdash

068

mdashmdash

mdash021

mdashmdash

mdash248

Strychnosspinosa

194

049

105

129

097

013

022

034

074

002

004

011

315

062

128

167

Syzygium

benguellense

065

mdashmdash

086

024

mdashmdash

034

005

mdashmdash

008

090

mdashmdash

123

Syzygium

cordatum

mdash049

052

086

mdash013

011

046

mdash049

003

035

mdash078

064

143

Syzygium

guineense

323

392

209

mdash338

373

065

mdash114

428

033

mdash699

908

285

mdashTerm

inalia

brachyste

mma

645

490

262

129

1135

437

119

114

2547

845

112

031

2629

1209

418

254

Uapaca

kirkiana

mdash343

mdash043

mdash219

mdash011

mdash278

mdash003

mdash062

mdash056

Uapaca

nitid

avarnitid

a065

049

209

431

024

013

054

205

007

002

039

271

091

654

277

727

Unidentified

sp1

(139230)

mdash049

052

mdashmdash

013

011

mdashmdash

004

005

mdashmdash

063

065

mdashUnidentified

sp2

(139236)

mdash14

7mdash

mdashmdash

039

mdashmdash

mdash013

mdashmdash

mdashmdash

mdashmdash

Vitexmadien

sis12

9294

052

129

048

090

022

046

030

035

006

040

187

396

076

188

Warneckea

sapinii

mdash049

mdash302

mdash013

mdash114

mdash007

mdash070

mdash064

mdash439






Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics



















Mature forests




Coordinate 1

Coo

rdin

ate 2


minus040

minus032

minus024

minus016

minus008

008

016

024

032

040





Old

Medium

Young

Mature

10

20

1

1

2

2

0 0 0

3

3 3

3

7

4



4 Discussion




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics
































5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics




















5 Conclusions



Competing Interests


Acknowledgments


References


































































































Journal of












Volume 2014

Advances in









Geophysics






































































































Journal of












Volume 2014

Advances in









Geophysics































































Journal of












Volume 2014

Advances in









Geophysics


























Journal of












Volume 2014

Advances in









Geophysics


















Journal of












Volume 2014

Advances in









Geophysics














Documents

Tree Species Diversity and Composition of Miombo Woodlands ...downloads.hindawi.com/journals/ijfr/2017/6202093.pdf · The diversity profiles indicate that mature forests were more