Growth Rates of Important East African Montane Forest Trees

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Growth Rates of Important East African Montane Forest TreesWith Particular Reference to Those of Mount KenyaAuthor(s) R W BussmannSource Journal of East African Natural History 88(1)69-78 1999Published By Nature KenyaEast African Natural History SocietyDOI httpdxdoiorg1029820012-8317(1999)88[69GROIEA]20CO2URL httpwwwbiooneorgdoifull1029820012-831728199929885B693AGROIEA5D20CO3B2

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Journal of East African Natural History 89 69-78 (1999)

GROWTH RATES OF IMPORTANT EAST AFRICAN MONTANEFOREST TREES WITH PARTICULAR REFERENCE TO THOSE OF

MOUNT KENYA

R W Bussmann

Lehrstuhl fUr Pflanzenphysiologie Universitat Bayreuth95440 Bayreuth Germany

ceja andinat-onlinede

ABSTRACT

The length increments of seedlings and branches and the extension growth ofspecimens of different age classes of 12 forest tree species were measured onMt Kenya between May 1992 and July 1995

Of all examined species the camphor tree Ocotea usambarensis showed thelowest growth rates The growth rates of other species of the primary forests namelyZanthoxyllum gillettii and Vitex keniensis were between 30 and 200 higher Theseedlings of Vitex outgrew even pioneer species such as Macaranga kilimandscharicaPax and Neoboutonia macrocalyx These trees showed growth rates at least twice ashigh as those of the primary species

Juniperus procera was found to be the fastest growing species in the cedar forestunderlining its success in forming dense stands after a fire Only young Podocarpuslatifolius showed a similar fast growth Olea europaea ssp cuspidata Olea capensisssp hochstetteri and Cassipourea malosana had nearly equal growth rates howeverconsiderably lower than those of Juniperus procera and Podocarpus latifolius

Hagenia abyssinica fell within the range of the fast growing species illustratingthe ability of this species to regenerate very fast under suitable conditions

INTRODUCTION

Today only about 2 of Kenyas land area are still covered with indigenous forests (Douteet al 1981) Mt Kenyas forests comprise nearly 2000 km2 or 20 of the total forestedarea (Beentje 1990) and is thus the most extended and coherent natural forest block in thecountry In addition to producing timber and firewood the forests of Mt Kenya representKenyas largest water catchment system supplying water for about 50 of the Kenyanpopulation In addition the National Park of Mt Kenya attracts up to 20000 visitors everyyear providing considerable financial revenue for the National Park Both the forest belt andalpine zone harbour an enormous variety of different conspicuous vegetation types with aunique flora and fauna including several endangered species eg the eastern bongo(Tragelaphus euryceros) black rhinoceros (Diceros bicomis) elephant (Loxodonta africana)black fronted duiker (Cephalophus nigrifrons) and giant forest hog (Hylochoerusmeinertzhageni) (Milner et al 1993) For all those reasons Mt Kenya has been included inthe UNESCO MAB 6 Program

70 RW Bussmann

Unfortunately the natural forests of Mt Kenya have been subjected to heavy tree fellingfor decades Moreover due to selective logging they also have undergone significantchanges of their species composition

For the total area of Kenya Myers (1979) estimated a former natural forest cover of15 which is equivalent to about 80000 km2bull Today even including plantations of exoticspecies only a quarter of this area still merits the term forest Although private tree plantingof exotic species has reduced the fuelwood deficit considerably the destruction of the naturalforests has increased drastically in the last decades Currently Kenya has an annualdeforestation rate of 15 which is in part driven by a human population growth rate of38 If current rates ofmiddotforest clearing continue no natural forest will remain by the year2040 (Barnes 1990)

Publications on the growth of tropical forest trees are still very rare Some growth rateshave been measured in cultivated species (eg Crow amp Weaver 1977 Schmidt amp Weaver1981) or have been reported from field observations (Breitsprecher amp Bethel 1990Chapman amp Chapman 1990 Lieberman 1885 Lieberman et ai 1985ab Lieberman ampLieberman 1987) Despite the enormous ecological and economical importance of themontane forests and the increasing demand for land and fuelwood of a rapidly increasingpopulation only very few studies have been carried out on the growth rates of the mostvaluable indigenous timber species (Kigomo 1980 1985ab 1987) and these were mainlyobtained from plantations Therefore information on the natural growth performance ofthese indigenous tree species is badly needed for the planning of sustainable forestry Thepresent study is a first step in filling this gap

MATERIAL AND METHODS

The growth of 12 tree species characteristic of the major types of primary and secondaryforests of Mt Kenya were determined between 1992 and 1995 All measurements were madeunder natural conditions Seeds of each species were germinated directly in the field inundisturbed stands of the mother trees The growth (elongation) of 50 seedlings of eachspecies was measured 90 days after germination

For the measurement of secondary growth 40 individuals of each species in three ageclasses (under five years 20-50 years over 50 years) were selected in the field and markedTo limit the influence of site differences these indiyiduals were selected always only in onerepresentative undisturbed stand per tree species In all 3 age classes trees of similar girthwidths at breast height (dbh) were selected to avoid growth differences due to different treeage (Struhsaker 1997) The dbh of these 40 marked specimens of each tree species weremeasured every 3 months Of 10 of these individuals of each species in the 3 age classesadditional samples were taken using a Mattson stem drill These samples were examinedmicroscopically As growth of the investigated species takes place mainly during and shortlyafter the wet seasons resulting in two distinct growth rings every year which were used asmarks Similar growth rings were observed in the Neotropics for periodically flooded forestsor dry seasons (Worbes 1989) The 50 values so obtained from the two methods were addedup to calculate the mean growth

Slices were cut from available timber samples of other individuals of the species andexamined in the same way to allow estimating the approximate age of specimens of a certaindbh However as these samples often could be only obtained from other sites these datawere not used in the final growth analysis to avoid errors due to site differences

Growth rates of important East African montane forest trees 71

To measure the elongation of branches 10 branches of 5 specimens of the markedindividuals of each species were marked Every month the last-unfolded pair of leaves at thetip of the branches was labelled At the end of the observation period the branches werecollected and the growth calculated from the distance between the labels

RESULTS

Growth of seedlings (Figure 1 table 1)Ocotea usambarensis by far the most important and endangered timber species of Kenyashowed the lowest growth rates of all investigated species Seedlings of this species onlyreached an average height of 46 cm 3 months after germination which is less than otherprimary forest species as Zanthoxyllum gillettii (53 cm) Podocarpus falcatus (69 cm) orOlea capensis and Olea europaea (88 and 75 cm) However there was another group ofprimary species characterised by a very fast growth of the seedlings eg Vitex keniensis(reaching 137 cm in the same time span) Podocarpus latifolius (101 cm) and Hageniaabyssinica (112 cm) Seedlings of secondary forest species as Macaranga kilimandscharica(1222 cm) and Neoboutonia macrocalyx (955 cm) elongated only a little faster than those ofthe named primary forest species This suggests that the influence of the early growth of theseedlings is of minor importance for the dynamics of the forest vegetation

25

20- E 15

i1 t I I ~~ f f f f f0

Fig 1 Mean height of seedlings after 90 days growth

Tab

le1

Mea

nan

nual

grow

thin

crem

ents

ofE

ast

Afr

ican

mon

tane

fore

sttr

ees

1992

-199

5(d

bhin

cm

xS

D

Ran

ge

cv)

lj

Hei

ghto

fse

edlin

gsaf

ter

Ann

ual

leng

thin

crem

ent

Ann

ual

diam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

90da

ysgr

owth

ofbr

anch

esof

tree

s1-

5ye

ars

oftr

ees

20-5

0ye

ars

oftr

eesgt

50ye

ars

N=

50504040

40In

crem

ent

(cm

)x

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SO

R

ange

cvF

amily

spe

cies

()

()

()

()

()

RH

ZO

PH

OR

AC

EA

EC

assi

pour

eam

aos

ana

772

371

273

-48

156

733

688

-47

073 011

042

-15

0470

090

29-

190

41 007

031

-17

128

629

51

096

061

053

RO

SA

CE

AE

Hag

enia

abys

sini

ca

112

34

385

21-

3919

48

1510

95-

42146

02

068

-14

0930

170

71-

180

72013

049

-18

172

331

44

171

125

095

CU

PR

ES

SA

CE

AE

Juni

peru

spr

ocer

a

886

54

254

-61

163

766

829

-47

154023

111

-15

087015

066

-17

0630

120

44-

1916

88

268

12

031

091

01E

UP

HO

RB

AC

EA

EM

acar

anga

kilim

ands

char

ica

122

23

676

89-

3035

29

1519

38-

26259

083

135

-32

186045

128

-24

136029

103

-21

195

749

26

322

257

187

Neo

bout

onia

mac

roca

yx

955

439

333

-46

391

821

269

1-21287

095

144

-33

192 052

097

-27

148031

098

-21

159

852

37

467

266

205

LAU

RA

CE

AE

Oco

tea

usam

bare

nsis

457

224

189

-49

183

421

121

6-23086

007

038

-8

052011

029

-21

025005

01-20

867

265

51

270

830

45O

LEA

CE

AE

Ole

aca

pens

is

879

319

329

-36

242

411

149

2-17077

019

025

-25

053 012

024

-23

045009

031

-20

156

632

25

113

092

062

Ole

aeu

ropa

eass

pcu

spid

ata

747

299

284

-40

206

618

106

6-30048

024

021

-50

036015

011

-42

032011

020

-34

132

233

93

092

059

055

PO

DO

CA

RP

AC

EA

EP

odoc

arpu

sfa

cat

us

688

255

412

-37

187

542

932

-29

085011

037

-13

032009

016

-28

028008

015

-29

132

927

11

129

052

041

Pod

ocar

pus

atif

oiu

s

101

33

555

91-

3522

4829

118

8-37127

017

099

-13

069014

032

-20

043014

019

-33

167

738

39

183

088

063

0V

ER

BE

NA

CE

AE

Vite

xke

nien

sis

136

59

812

99-

72no

tmea

sure

d1

870

221

26-

120

72019

057

-26

059014

031

-24

~21

24

294

103

074

tll

RU

TA

CE

AE

=

Zan

thox

yllu

mgi

lletti

i5

262

681

79-

51no

tmea

sure

d1

390

210

77-

151

02 016

078

-16

073 012

058

-21

933

205

167

097

I


Growth of branchesMacaranga and Neoboutonia outgrew all other investigated tree species (figure 2) Theirannual length increments of the branches (352 and 391 em respectively) was on averageapproximately about 75 higher than the growth rate of the primary species whichelongated by about 20 emyear on average

E~-cOJ

E~ltJ5pound-ClC~

50

45

40

35

30

25

20

15

10

5o ~I-~-~-~-~-~~-~-~-~-~~

Fig 2 Mean annual length increment of branches 1992-1995

Diameter increment by secondary growth (figures 3A B C)One-to-five-year old trees of all investigated species showed the highest rates of secondarygrowth Older trees reached only 25-60 of this rate

According to their rates of secondary growth the young trees could be grouped in threeclasses Ocotea usambarensis Podocarpus falcatus both Olea species and Cassipoureamalosana were the slowest growing species attaining an annual diameter increment ofbetween 05 and 09 em All other primary forest species (Zanthoxyllum gillettii Vitexkeniensis Podocarpus latifolius Juniperus procera and Hagenia abyssinica) grew abouttwice as fast (14-19 emyear) The third group consisted of the two Euphorbiaceae(Macaranga kilimandscharica and Neoboutonia macrocalyx which are typical for secondaryforests) reaching 26 and 29 emyear

The difference between primary and secondary forest species became even more evidentwhen comparing the secondary growth rates of trees older than 20 years Here the growthrates of all species decreased by approximately 30-50 however the observed decreasewas lowest for the secondary species

In individuals older than 20 years the percentage decrease in diameter increment wascomparable for all species The two pioneer species of the secondary forests however stillgrew about 4 times as fast as all primary species

74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I

Growth rates of important East African montane forest trees

4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75

Figure 3 Mean annual dbh increment of A young trees (1-5 years) 1992-1995 B 20-50 yearold trees 1992-1995 C mature trees (gt50 years) 1992-1995

DISCUSSION

The median dbh increments found are very well comparable to the ranges given for meanannual growth rates for African forest species (05-48 mm Struhsaker 1997) andneotropical rainforest species (eg 26-78 mm Chapman amp Chapman 1990 25-81 mmCrow amp Weaver 1977 08-115 mm Lang amp Knight 1983 04-134 mm Lieberman etat 1985a 11-65 mm Schmidt amp Weaver 1981) This is especially interesting asparticularly in contrast to the neotropical sites the study area shows very distinct wet and dryseasons In contrast to Connell et at (1984) and Alder (1992) the annual dbh increment

generally decreased with size and age of the measured individuals reaching the lowest ratesfor mature trees which is also reported by Struhsaker (1997) for Kibale forest whereasSwaine et at (1987a) found larger trees having higher growth rates In close correspondenceto the results of Felfili (1995) however light requiring canopy and pioneer species showedthe highest increment rates

The dbh growth rates for Ocotea Vitex Podocarpus and Juniperus measured in thenatural forests were similar to those obtained by Kigomo (1981 1985ab 1987) for trees ofroutinely thinned plantations

The growth rates of the species measured showed high differences between species butalso between individuals of the same species This corresponds closely to the observations of

76 RW Bussmann

Korning amp Balslev (1994) In contrast the growth of individual trees during the study periodshowed much less variation This phenomenon is also mentioned by Swaine (1989) Thecoefficient of variation (cvs) values calculated never reached more than 61 This is a veryclear contrast to the high individual growth variability found for species in other tropicalforest regions Especially in tropical lowland forests cvs rates of often much more than100 were reported (Felfili 1995 Gentry amp Terborgh 1990 Pires amp Prance 1977Swaine et al 1987b)

CONCLUSIONS

The results of the present study give valuable information for the management of naturalforests especially with respect to replanting After clearcutting or for fast reforestation ofagronomically used areas indigenous pioneer species as Macaranga kilimandscharica andNeoboutonia macrocalyx could be extremely useful Due to their fast growth these specieswould be valuable to limit the influence of erosion In their shade more valuable slow

growing primary species eg Ocotea usambarensis can be raised (Bussmann amp Beck1995) For the fast establishment of plantations or agroforestry several valuable and fastgrowing indigenous species as Juniperus procera and Vitex keniensis could easily be used toreplace the presently-favoured exotic softwoods

ACKNOWLEDGMENTS

I gratefully acknowledge the financial support of this work by the DeutscheForschungsgemeinschaft (DFG) and the Gesellschaft fUr technische Zusammenarbeit (GTZ)The author would also like to thank his colleagues Prof JO Kokwaro Nairobi and Dr JCOnyango Maseno for their untiring assistance Finally thanks are due to the NationalResearch Council of Kenya for granting permission for research on Mt Kenya and to BongoWoodley the Warden of Mt Kenya National Park and his staff for all their logistic support

REFERENCES

Alder D (1992) Simple methods for calculating mifiimum diameter and sustainable yield intropical forest In FR Miller amp KL Adam (eds) Wise management of tropical forestsProceedings of the Oxford Conference on Tropical Forests Oxford Forestry Institute pp189-200

Barnes RFW (1990) Deforestation trends in tropical Africa African Journal ofEcology 28 161-173

Beentje HJ (1990) The forests of Kenya (Proceedings of the twelfth plenary meeting ofaetfat symposium II) Mitteilungen des lnstituts fUr Allgemeine Botanik Hamburg 23a265-286

Breitsprecher A amp JS Bethel (1990) Stem-growth periodicity of trees in a tropical wetforest in Costa Rica Ecology 71 1156-1164

Bussmann RW amp E Beck (1995) Regeneration and cyclic processes in the ocotea-forests(Ocotea usambarensis Engl) of Mount Kenya Verhandlungen der Gesellschaft fUrOkologie 24 35-39


Chapman CA amp LJ Chapman (1990) Density and growth rate of some tropical dry foresttrees Comparison between successional forest types Bulletin of the Torrey BotanicalClub 117 226-231

Connell JA JG Tracey amp LJ Webb (1984) Compensatory recruitment growth andmortality as factors maintaining rain forest tree diversity Ecological Monographs 54(2)141-164

Crow TR amp PL Weaver (1977) Tree growth in a tropical moist forest of Puerto RicoUnited States Forest Service Research Paper ITF-22

Doute R N Ochanda amp H Epp (1981) Forest cover mapping in Kenya using remotesensing techniques KREMU Nairobi

Felfili JM (1995) Growth recruitment and mortality in the Gama gallery forest in centralBrazil over a six-year period (1985-1991) Journal of Tropical Ecology 11(1) 67-83

Gentry AH amp J Terborgh (1990) Composition and dynamics of Cocha Cashu maturefloodplain forest In AH Gentry (ed) Four neotropical rainforests Yale UniversityPress New Haven pp 542-564

Kigomo BN (1980) Crown-bole diameter relationships of Juniperus procera (Cedar) andits application to stand density control and production survey in natural stands EastAfrican Agriculture and Forestry Journal 46(2) 27-37

Kigomo BN (1981) Observations on the growth of Vitex keniensis TurrilI (Meru Oak) inplantation East African Agriculture and Forestry Journal 47(2) 32-37

Kigomo BN (1985a) Diameter increment and growth of Podocarpus trees in naturalforests Kenya Journal of Science and Technology Seies B 6(2) 113-121

Kigomo BN (1985b) Growth characteristics of natural regenerates of African PencilCedar (Juniperus procera) East African Agriculture and Forestry Journal 50(3) 54-60

Kigomo BN (1987) The growth of Camphor (Ocotea usambarensis Engl) in plantation inthe eastern Aberdare Range Kenya East African Agriculture and Forestry Journal 52(3)141-147

Korning J amp H Balslev (1994) Growth rates and mortality patterns of tropical lowland treespecies and the relation to forest structure in Amazonian Ecuador Journal of TropicalEcology 10(2) 151-166

Lang GE amp DH Knight (1983) Tree growth mortality recruitment and canopy gapformation during a lO-year period in a tropical moist forest Ecology 64 1075-1080

Lieberman M amp D Lieberman (1985) Simulation of growth curves from periodicincrement data Ecology 66(2) 632-635

Lieberman D amp M Lieberman (1987) Forest tree growth and dynamics at La Selva CostaRica Journal of Tropical Ecology 3 347-359

Lieberman D M Lieberman G Hartshorn amp R Peralta (1985a) Growth rates and ageshysize relationships of wet tropical forest trees in Costa Rica Journal of Tropical Ecology1 97-109

Lieberman D M Lieberman R Peralta amp G Hartshorn (1985b) Mortality patterns andstand turnover rates in wet tropical forest in Costa Rica Journal of Tropical Ecology 1915-924

Milner J M Litoroh amp M Gathua (1993) Mammals of Mt Kenya amp its forests-apreliminary survey (draft report) Nairobi KIFCON

Myers N (1979) The sinking Ark Pergamon Press OxfordPires JM amp GT Prance (1977) The Amazon forest a natural heritage to be preserved In

GT Prance amp BS Elias (eds) Extinction is forever Threatened and endangered

78 RW Bussmann

species of plants in the Americas and their significance in ecosystems today and in thefuture Symposium Proceedings New York Botanical Garden pp 158-194

Schmidt R amp PL Weaver (1981) Tree diameter increment in the subtropical moist lifezone of Puerto Rico Turrialba 31 261-263

Struhsaker TT (1997) Ecology of an African Rain Forest-Logging in Kibale and theConflict between Conservation and Exploitation University of Florida Press Gainesville

Swaine MD (1989) Population dynamics of tree species in tropical forests In LB HolmshyNielsen I Nielsen amp H Balslev (eds) Tropical forests-Botanical dynamics speciationand diversity Academic Press London pp 101-110

Swaine MD JB Hall amp 11 Alexande (1987a) Tree population dynamics at KadeGhana (1968-1992) Journal of Tropical Ecology 3 331-345

Swaine MD D Lieberman amp FE Putz (1987b) The dynamics of tree populations intropical forests a review Journal of Tropical Ecology 3 359-366

Worbes M (1989) Growth rings increment and age of trees in inundation forests savannasand mountain forest in the Neotropics IAWA Bulletin ns 10 109-122

Journal of East African Natural History 89 69-78 (1999)

GROWTH RATES OF IMPORTANT EAST AFRICAN MONTANEFOREST TREES WITH PARTICULAR REFERENCE TO THOSE OF

MOUNT KENYA

R W Bussmann

Lehrstuhl fUr Pflanzenphysiologie Universitat Bayreuth95440 Bayreuth Germany

ceja andinat-onlinede

ABSTRACT

The length increments of seedlings and branches and the extension growth ofspecimens of different age classes of 12 forest tree species were measured onMt Kenya between May 1992 and July 1995

Of all examined species the camphor tree Ocotea usambarensis showed thelowest growth rates The growth rates of other species of the primary forests namelyZanthoxyllum gillettii and Vitex keniensis were between 30 and 200 higher Theseedlings of Vitex outgrew even pioneer species such as Macaranga kilimandscharicaPax and Neoboutonia macrocalyx These trees showed growth rates at least twice ashigh as those of the primary species

Juniperus procera was found to be the fastest growing species in the cedar forestunderlining its success in forming dense stands after a fire Only young Podocarpuslatifolius showed a similar fast growth Olea europaea ssp cuspidata Olea capensisssp hochstetteri and Cassipourea malosana had nearly equal growth rates howeverconsiderably lower than those of Juniperus procera and Podocarpus latifolius

Hagenia abyssinica fell within the range of the fast growing species illustratingthe ability of this species to regenerate very fast under suitable conditions

INTRODUCTION

Today only about 2 of Kenyas land area are still covered with indigenous forests (Douteet al 1981) Mt Kenyas forests comprise nearly 2000 km2 or 20 of the total forestedarea (Beentje 1990) and is thus the most extended and coherent natural forest block in thecountry In addition to producing timber and firewood the forests of Mt Kenya representKenyas largest water catchment system supplying water for about 50 of the Kenyanpopulation In addition the National Park of Mt Kenya attracts up to 20000 visitors everyyear providing considerable financial revenue for the National Park Both the forest belt andalpine zone harbour an enormous variety of different conspicuous vegetation types with aunique flora and fauna including several endangered species eg the eastern bongo(Tragelaphus euryceros) black rhinoceros (Diceros bicomis) elephant (Loxodonta africana)black fronted duiker (Cephalophus nigrifrons) and giant forest hog (Hylochoerusmeinertzhageni) (Milner et al 1993) For all those reasons Mt Kenya has been included inthe UNESCO MAB 6 Program

70 RW Bussmann










RESULTS


25

20- E 15



Tab

le1

Mea

nan

nual

grow

thin

crem

ents

ofE

ast

Afr

ican

mon

tane

fore

sttr

ees

1992

-199

5(d

bhin

cm

xS

D

Ran

ge

cv)

lj

Hei

ghto

fse

edlin

gsaf

ter

Ann

ual

leng

thin

crem

ent

Ann

ual

diam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

90da

ysgr

owth

ofbr

anch

esof

tree

s1-

5ye

ars

oftr

ees

20-5

0ye

ars

oftr

eesgt

50ye

ars

N=

50504040

40In

crem

ent

(cm

)x

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SO

R

ange

cvF

amily

spe

cies

()

()

()

()

()

RH

ZO

PH

OR

AC

EA

EC

assi

pour

eam

aos

ana

772

371

273

-48

156

733

688

-47

073 011

042

-15

0470

090

29-

190

41 007

031

-17

128

629

51

096

061

053

RO

SA

CE

AE

Hag

enia

abys

sini

ca

112

34

385

21-

3919

48

1510

95-

42146

02

068

-14

0930

170

71-

180

72013

049

-18

172

331

44

171

125

095

CU

PR

ES

SA

CE

AE

Juni

peru

spr

ocer

a

886

54

254

-61

163

766

829

-47

154023

111

-15

087015

066

-17

0630

120

44-

1916

88

268

12

031

091

01E

UP

HO

RB

AC

EA

EM

acar

anga

kilim

ands

char

ica

122

23

676

89-

3035

29

1519

38-

26259

083

135

-32

186045

128

-24

136029

103

-21

195

749

26

322

257

187

Neo

bout

onia

mac

roca

yx

955

439

333

-46

391

821

269

1-21287

095

144

-33

192 052

097

-27

148031

098

-21

159

852

37

467

266

205

LAU

RA

CE

AE

Oco

tea

usam

bare

nsis

457

224

189

-49

183

421

121

6-23086

007

038

-8

052011

029

-21

025005

01-20

867

265

51

270

830

45O

LEA

CE

AE

Ole

aca

pens

is

879

319

329

-36

242

411

149

2-17077

019

025

-25

053 012

024

-23

045009

031

-20

156

632

25

113

092

062

Ole

aeu

ropa

eass

pcu

spid

ata

747

299

284

-40

206

618

106

6-30048

024

021

-50

036015

011

-42

032011

020

-34

132

233

93

092

059

055

PO

DO

CA

RP

AC

EA

EP

odoc

arpu

sfa

cat

us

688

255

412

-37

187

542

932

-29

085011

037

-13

032009

016

-28

028008

015

-29

132

927

11

129

052

041

Pod

ocar

pus

atif

oiu

s

101

33

555

91-

3522

4829

118

8-37127

017

099

-13

069014

032

-20

043014

019

-33

167

738

39

183

088

063

0V

ER

BE

NA

CE

AE

Vite

xke

nien

sis

136

59

812

99-

72no

tmea

sure

d1

870

221

26-

120

72019

057

-26

059014

031

-24

~21

24

294

103

074

tll

RU

TA

CE

AE

=

Zan

thox

yllu

mgi

lletti

i5

262

681

79-

51no

tmea

sure

d1

390

210

77-

151

02 016

078

-16

073 012

058

-21

933

205

167

097

I



E~-cOJ

E~ltJ5pound-ClC~

50

45

40

35

30

25

20

15

10

5o ~I-~-~-~-~-~~-~-~-~-~~






74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I


4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann








70 RW Bussmann










RESULTS


25

20- E 15



Tab

le1

Mea

nan

nual

grow

thin

crem

ents

ofE

ast

Afr

ican

mon

tane

fore

sttr

ees

1992

-199

5(d

bhin

cm

xS

D

Ran

ge

cv)

lj

Hei

ghto

fse

edlin

gsaf

ter

Ann

ual

leng

thin

crem

ent

Ann

ual

diam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

90da

ysgr

owth

ofbr

anch

esof

tree

s1-

5ye

ars

oftr

ees

20-5

0ye

ars

oftr

eesgt

50ye

ars

N=

50504040

40In

crem

ent

(cm

)x

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SO

R

ange

cvF

amily

spe

cies

()

()

()

()

()

RH

ZO

PH

OR

AC

EA

EC

assi

pour

eam

aos

ana

772

371

273

-48

156

733

688

-47

073 011

042

-15

0470

090

29-

190

41 007

031

-17

128

629

51

096

061

053

RO

SA

CE

AE

Hag

enia

abys

sini

ca

112

34

385

21-

3919

48

1510

95-

42146

02

068

-14

0930

170

71-

180

72013

049

-18

172

331

44

171

125

095

CU

PR

ES

SA

CE

AE

Juni

peru

spr

ocer

a

886

54

254

-61

163

766

829

-47

154023

111

-15

087015

066

-17

0630

120

44-

1916

88

268

12

031

091

01E

UP

HO

RB

AC

EA

EM

acar

anga

kilim

ands

char

ica

122

23

676

89-

3035

29

1519

38-

26259

083

135

-32

186045

128

-24

136029

103

-21

195

749

26

322

257

187

Neo

bout

onia

mac

roca

yx

955

439

333

-46

391

821

269

1-21287

095

144

-33

192 052

097

-27

148031

098

-21

159

852

37

467

266

205

LAU

RA

CE

AE

Oco

tea

usam

bare

nsis

457

224

189

-49

183

421

121

6-23086

007

038

-8

052011

029

-21

025005

01-20

867

265

51

270

830

45O

LEA

CE

AE

Ole

aca

pens

is

879

319

329

-36

242

411

149

2-17077

019

025

-25

053 012

024

-23

045009

031

-20

156

632

25

113

092

062

Ole

aeu

ropa

eass

pcu

spid

ata

747

299

284

-40

206

618

106

6-30048

024

021

-50

036015

011

-42

032011

020

-34

132

233

93

092

059

055

PO

DO

CA

RP

AC

EA

EP

odoc

arpu

sfa

cat

us

688

255

412

-37

187

542

932

-29

085011

037

-13

032009

016

-28

028008

015

-29

132

927

11

129

052

041

Pod

ocar

pus

atif

oiu

s

101

33

555

91-

3522

4829

118

8-37127

017

099

-13

069014

032

-20

043014

019

-33

167

738

39

183

088

063

0V

ER

BE

NA

CE

AE

Vite

xke

nien

sis

136

59

812

99-

72no

tmea

sure

d1

870

221

26-

120

72019

057

-26

059014

031

-24

~21

24

294

103

074

tll

RU

TA

CE

AE

=

Zan

thox

yllu

mgi

lletti

i5

262

681

79-

51no

tmea

sure

d1

390

210

77-

151

02 016

078

-16

073 012

058

-21

933

205

167

097

I



E~-cOJ

E~ltJ5pound-ClC~

50

45

40

35

30

25

20

15

10

5o ~I-~-~-~-~-~~-~-~-~-~~






74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I


4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann










RESULTS


25

20- E 15



Tab

le1

Mea

nan

nual

grow

thin

crem

ents

ofE

ast

Afr

ican

mon

tane

fore

sttr

ees

1992

-199

5(d

bhin

cm

xS

D

Ran

ge

cv)

lj

Hei

ghto

fse

edlin

gsaf

ter

Ann

ual

leng

thin

crem

ent

Ann

ual

diam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

90da

ysgr

owth

ofbr

anch

esof

tree

s1-

5ye

ars

oftr

ees

20-5

0ye

ars

oftr

eesgt

50ye

ars

N=

50504040

40In

crem

ent

(cm

)x

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SO

R

ange

cvF

amily

spe

cies

()

()

()

()

()

RH

ZO

PH

OR

AC

EA

EC

assi

pour

eam

aos

ana

772

371

273

-48

156

733

688

-47

073 011

042

-15

0470

090

29-

190

41 007

031

-17

128

629

51

096

061

053

RO

SA

CE

AE

Hag

enia

abys

sini

ca

112

34

385

21-

3919

48

1510

95-

42146

02

068

-14

0930

170

71-

180

72013

049

-18

172

331

44

171

125

095

CU

PR

ES

SA

CE

AE

Juni

peru

spr

ocer

a

886

54

254

-61

163

766

829

-47

154023

111

-15

087015

066

-17

0630

120

44-

1916

88

268

12

031

091

01E

UP

HO

RB

AC

EA

EM

acar

anga

kilim

ands

char

ica

122

23

676

89-

3035

29

1519

38-

26259

083

135

-32

186045

128

-24

136029

103

-21

195

749

26

322

257

187

Neo

bout

onia

mac

roca

yx

955

439

333

-46

391

821

269

1-21287

095

144

-33

192 052

097

-27

148031

098

-21

159

852

37

467

266

205

LAU

RA

CE

AE

Oco

tea

usam

bare

nsis

457

224

189

-49

183

421

121

6-23086

007

038

-8

052011

029

-21

025005

01-20

867

265

51

270

830

45O

LEA

CE

AE

Ole

aca

pens

is

879

319

329

-36

242

411

149

2-17077

019

025

-25

053 012

024

-23

045009

031

-20

156

632

25

113

092

062

Ole

aeu

ropa

eass

pcu

spid

ata

747

299

284

-40

206

618

106

6-30048

024

021

-50

036015

011

-42

032011

020

-34

132

233

93

092

059

055

PO

DO

CA

RP

AC

EA

EP

odoc

arpu

sfa

cat

us

688

255

412

-37

187

542

932

-29

085011

037

-13

032009

016

-28

028008

015

-29

132

927

11

129

052

041

Pod

ocar

pus

atif

oiu

s

101

33

555

91-

3522

4829

118

8-37127

017

099

-13

069014

032

-20

043014

019

-33

167

738

39

183

088

063

0V

ER

BE

NA

CE

AE

Vite

xke

nien

sis

136

59

812

99-

72no

tmea

sure

d1

870

221

26-

120

72019

057

-26

059014

031

-24

~21

24

294

103

074

tll

RU

TA

CE

AE

=

Zan

thox

yllu

mgi

lletti

i5

262

681

79-

51no

tmea

sure

d1

390

210

77-

151

02 016

078

-16

073 012

058

-21

933

205

167

097

I



E~-cOJ

E~ltJ5pound-ClC~

50

45

40

35

30

25

20

15

10

5o ~I-~-~-~-~-~~-~-~-~-~~






74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I


4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann








Tab

le1

Mea

nan

nual

grow

thin

crem

ents

ofE

ast

Afr

ican

mon

tane

fore

sttr

ees

1992

-199

5(d

bhin

cm

xS

D

Ran

ge

cv)

lj

Hei

ghto

fse

edlin

gsaf

ter

Ann

ual

leng

thin

crem

ent

Ann

ual

diam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

Ann

uald

iam

eter

incr

ease

90da

ysgr

owth

ofbr

anch

esof

tree

s1-

5ye

ars

oftr

ees

20-5

0ye

ars

oftr

eesgt

50ye

ars

N=

50504040

40In

crem

ent

(cm

)x

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SD

R

ange

cvx

SO

R

ange

cvF

amily

spe

cies

()

()

()

()

()

RH

ZO

PH

OR

AC

EA

EC

assi

pour

eam

aos

ana

772

371

273

-48

156

733

688

-47

073 011

042

-15

0470

090

29-

190

41 007

031

-17

128

629

51

096

061

053

RO

SA

CE

AE

Hag

enia

abys

sini

ca

112

34

385

21-

3919

48

1510

95-

42146

02

068

-14

0930

170

71-

180

72013

049

-18

172

331

44

171

125

095

CU

PR

ES

SA

CE

AE

Juni

peru

spr

ocer

a

886

54

254

-61

163

766

829

-47

154023

111

-15

087015

066

-17

0630

120

44-

1916

88

268

12

031

091

01E

UP

HO

RB

AC

EA

EM

acar

anga

kilim

ands

char

ica

122

23

676

89-

3035

29

1519

38-

26259

083

135

-32

186045

128

-24

136029

103

-21

195

749

26

322

257

187

Neo

bout

onia

mac

roca

yx

955

439

333

-46

391

821

269

1-21287

095

144

-33

192 052

097

-27

148031

098

-21

159

852

37

467

266

205

LAU

RA

CE

AE

Oco

tea

usam

bare

nsis

457

224

189

-49

183

421

121

6-23086

007

038

-8

052011

029

-21

025005

01-20

867

265

51

270

830

45O

LEA

CE

AE

Ole

aca

pens

is

879

319

329

-36

242

411

149

2-17077

019

025

-25

053 012

024

-23

045009

031

-20

156

632

25

113

092

062

Ole

aeu

ropa

eass

pcu

spid

ata

747

299

284

-40

206

618

106

6-30048

024

021

-50

036015

011

-42

032011

020

-34

132

233

93

092

059

055

PO

DO

CA

RP

AC

EA

EP

odoc

arpu

sfa

cat

us

688

255

412

-37

187

542

932

-29

085011

037

-13

032009

016

-28

028008

015

-29

132

927

11

129

052

041

Pod

ocar

pus

atif

oiu

s

101

33

555

91-

3522

4829

118

8-37127

017

099

-13

069014

032

-20

043014

019

-33

167

738

39

183

088

063

0V

ER

BE

NA

CE

AE

Vite

xke

nien

sis

136

59

812

99-

72no

tmea

sure

d1

870

221

26-

120

72019

057

-26

059014

031

-24

~21

24

294

103

074

tll

RU

TA

CE

AE

=

Zan

thox

yllu

mgi

lletti

i5

262

681

79-

51no

tmea

sure

d1

390

210

77-

151

02 016

078

-16

073 012

058

-21

933

205

167

097

I



E~-cOJ

E~ltJ5pound-ClC~

50

45

40

35

30

25

20

15

10

5o ~I-~-~-~-~-~~-~-~-~-~~






74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I


4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann










E~-cOJ

E~ltJ5pound-ClC~

50

45

40

35

30

25

20

15

10

5o ~I-~-~-~-~-~~-~-~-~-~~






74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I


4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann








74

4

35- 3

K -25

c Q) lI~E

~~rt

C)

1t

5c ~

0C

050

RW Bussmann

f f

3A

~bullbull-cQ)

E

~5c0C

38

4

35

3

25

2

15

1

05

o I I I I I I I I I


4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann









4

353

- E~ 251 Q) 2E

~15t

r

5r 1

rr t

0

rl- t

r

0

050

3C

75


DISCUSSION





76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann








76 RW Bussmann


CONCLUSIONS



ACKNOWLEDGMENTS


REFERENCES



























78 RW Bussmann





























78 RW Bussmann








78 RW Bussmann








Growth Rates of Important East African Montane Forest Trees

Documents