Diet of forest elephants and their role in seed dispersal in the ...

Diet of forest elephants and their role in seed dispersal 

in the Bossematié Forest Reserve, Ivory Coast 

by J. THEUERKAUF 1 , W. E. WAITKUWAIT 2 , Y. GUIRO 2 , H. ELLENBERG 3 

and S. POREMBSKI 4 

1 Department of Biology, Philipps University, 35032 Marburg, Germany 

2 Projet forestier SODEFOR-GTZ, B.P. 878, Abengourou, Côte d’Ivoire 

3 Institute for World Forestry, Federal Research Centre for Forestry and Forest Products, 

Leuschnerstr. 91, 21031 Hamburg, Germany 

4 Institute for Biodiversity Research, Department of Botany, University of Rostock, 

Wismarsche Str. 8, 18051 Rostock, Germany 

Present address of corresponding author : 

Jörn Theuerkauf, Am Schäperkamp 3, 27711 Osterholz-Scharmbeck, Germany 

e-mail : Theuerkauf.Joern@t-online.de 

Abstract. – We followed fresh tracks of forest elephants (Loxodonta africana cyclotis Matschie, 

1900) in the heavily exploited semi-deciduous Bossematié Forest Reserve (southeastern 

Ivory Coast) and documented the elephants’ diet. The aim was to learn which ecological types of 

plants elephants eat in the very disturbed forest, as well as estimate the role of forest elephants in 

the dispersal of seeds in the study area. When comparing the 147 food plants with the flora of 

the study area, it appeared that elephants selected trees (P < 0.001) and avoided shrubs (P < 

0.001) as well as herb and grass species (P = 0.002). In contrast to results of studies in more 

natural forests, the elephants preferred shade tolerant (P = 0.024) and avoided light tolerant species 

(P = 0.008). They preferred the fruits (P = 0.043) and bark (P < 0.001) of shade tolerant 

species and avoided the bark of light tolerant species (P = 0.008). We estimated that in our study 

area elephants dispersed seeds at a mean distance of 5 to 12 km and were responsible for the dispersal 

of at least 66 species (more than 10 % of identified spermatophytes in the forest). 

Résumé. –Dans la Forêt Classée de Bossematié (sud-est de la Côte d’Ivoire), une forêt 

dense humide semi-décidue et fortement exploitée, nous avons suivi des traces fraîches d’éléphants 

de forêt (Loxodonta africana cyclotis Matschie, 1900) et documenté le régime alimentaire 

des éléphants. Le but était de découvrir les types écologiques des plantes que les éléphants 

consomment dans la forêt très ouverte et d’évaluer le rôle des éléphants dans la dissémination 

des graines sur l’aire de la forêt. Si l’on compare les 147 espèces de plantes fourragères avec la 

flore du terrain d’étude, les éléphants ont préféré les arbres (P < 0,001) et évité les arbustes 

(P < 0,001) et les herbacées et graminées (P = 0,002). En contraste avec les résultats de 

recherches dans des forêts plus naturelles, les éléphants ont préféré les espèces sciaphiles 

(P = 0,024) et évité les espèces héliophiles (P = 0,008). Ils ont préféré les fruits (P = 0,043) et 

l’écorce (P < 0,001) des espèces sciaphiles et évité l’écorce des espèces héliophiles (P = 0,008). 

Nous avons estimé une distance de dissémination des graines de 5 à 12 km en moyenne. Les éléphants 

disséminent dans la forêt étudiée au moins 66 espèces (plus que 10 % des spermatophytes 

déterminés dans la forêt). 

KEY WORDS : Loxodonta africana cyclotis, Ivory Coast, food choice, forest regeneration, 

germination. 

___________ 

Mammalia, t. 64, n° 2, 2000 : 447-459.

448 

MAMMALIA 

INTRODUCTION 

Forest elephants are reported to prefer secondary to primary parts of rain forests 

(Barnes et al. 1991 ; Prins and Reitsma 1989 ; Struhsaker et al. 1996). Exploited 

forests can bear a higher elephant density than primary forests (Barnes et al. 1991 ; 

Ekobo 1993) provided that elephants are not disturbed by human activity (Barnes et al. 

1991 ; Hall et al. 1997). The selection of secondary forests should be reflected in the 

elephants’ diet, thus elephants should prefer light tolerant to shade tolerant plant species. 

However, this was never tested in the studies that documented the diet of ele- 

phants (Alexandre 1978 ; Merz 1981 ; Short 1981 ; Martin 1982 ; Tchamba and Seme 

1993 ; White et al. 1993 ; Feer 1995 ; Maurois et al. 1997). In West African forests 

where there are usually no large clearings with grass, elephants feed mainly on trees 

and eat the fruits of 55-72 species (Alexandre 1978 ; Short 1981 ; Martin 1982 ; White 

et al. 1993 ; Feer 1995). In regions where forest elephants eat mainly grass, the number 

of fruit species eaten is much lower at 22-33 species (Tchamba and Seme 1993 ; Maurois 

et al. 1997). Therefore, primary forest parts with many fruiting trees may be more 

important in West African forests. We tested this hypothesis in the heavily exploited 

Bossematié Forest Reserve (Southeast Ivory Coast) where, because of past timber 

extractions, the canopy cover is much more open than in the forests where the above 

mentioned studies on habitat selection were conducted. 

Forest elephants disperse seeds of many species (e.g. Alexandre 1978 ; Merz 

1981 ; Gautier-Hion et al. 1985 ; Lieberman et al. 1987). The transit through the elephant 

intestine can even increase the germination rate of seeds, as found for Balanites 

wilsoniana (Chapman et al. 1992). However, probably only few species completely 

depend on elephant dispersal (Hawthorne and Parren 2000). Besides seed dispersal, 

elephants also influence the forest structure by browsing and trampling, especially in 

forests that have been exploited (Struhsaker et al. 1996). To evaluate the effect of seed 

dispersal on forest regeneration, it is essential to know about the germination success 

and the survival rate of seedlings in the elephant droppings. Brahmachary (1980) and 

Lieberman et al. (1987) studied the germination success of seeds in elephant dung in 

cultivated plots. It is however not known if the germination success under artificial 

conditions is representative of natural conditions and could be used to evaluate the role 

of forest elephants in the rehabilitation of forests. 

The objectives of this article are to study (1) which ecological type of food species 

forest elephants prefer in the heavily exploited study area, (2) if the germination 

success under artificial conditions is representative of natural conditions, and (3) the 

importance of forest elephants as seed dispersal agents in the regeneration of the Bossematié 

Forest Reserve. 

METHODS 

Study area 

The Bossematié Forest Reserve (220 km²) is a heavily exploited moist 

semi-deciduous rain forest in the lowland zone of southeastern Ivory Coast reaching 

from 3°24’ to 3°35’ W and from 6°22’ to 6°33’ N. The annual distribution of rainfall is 

bimodal with peaks in September-October and April-July. August and December to 

February are dry. Mean annual rainfall for 1961-1990 was 1330 mm but varied significantly 

from year to year (Schroth 1992). The northern part of the Bossematié is classi-

DISPERION DES GRAINES PAR LES ELEPHANTS DE FORET 

fied as « Celtis spp. forest with Triplochiton scleroxylon » whereas the southern part is a 

moister « variant with Nesogordonia papaverifera and Khaya ivorensis » (Guillaumet 

and Adjanohoun 1971). When we undertook this research, 526 species of spermatophytes 

were identified in the Bossematié Forest Reserve (Mühlenberg et al. 1993 and 

this study). For the 30 years up to 1988, the forest was exploited several times for timber, 

resulting in a mosaic of degraded forest with few old fruit trees and many small 

clearings, which are mainly covered by the neophytic Asteraceae Chromolaena odorata 

(Wöll 1992). Only 16 % of the forest has a canopy cover over 30 %, the canopy cover 

in 60 % of the forest is between 10 % and 30 %, and about 8 % of its surface are mostly 

illegal cocoa and coffee plantations (Wöll 1992). Elephant poaching was commonplace 

in the 1980’s until 1990, but when a rehabilitation project for the Bossematié Forest 

Reserve was implemented, elephant poachers were arrested and poaching stopped. 

Elephant diet 

On 28 days from August to December 1993, we followed fresh tracks of forest 

elephants to identify from which plant species the elephants ate fruits, bark, leaves or 

twigs. Their feeding signs permitted us to identify or collect the plants they had eaten. 

We classified the 526 known spermatophytes of the Bossematié Forest Reserve according 

to their life forms and light requirements as done by Hutchinson and Dalziel 

(1954-1972) and Aubréville (1959). We then calculated the percentage of species of a 

given life form or light requirement for the study area (expected values), which we 

compared to percentages in the elephants’ diet. 

Seed germination in elephant dung 

In order to quantify the success of germination in elephant droppings under different 

conditions, we observed seedlings in 90 elephant droppings of a known age. These 

droppings had been excreted by a group of 6 elephants during 12 hours on the 

18/9/1993 and by a group of 5 elephants during 6 hours on the 5/10/1993. We assumed 

that the diversity and abundance of seeds in the different droppings were comparable 

as elephant groups fed together. We took 30 of the droppings to a clearing overgrown 

with Chromolaena odorata in which we had cut 1-m wide strips. We cleared the vegetation 

around 10 droppings monthly, around 10 droppings every two months and never 

cleared around the last 10 droppings. We pooled the data for artificial conditions, 

because the clearing of vegetation had no effect on the germination success (one-wayanova, 

P = 0.448). We left another 60 droppings where they had fallen (30 in the shade 

and 30 in the sun) as a control of germination under natural conditions. We controlled 

all droppings once a month until April 1994, counted the number of seedlings and 

determined the species when possible. We added data about germination of seeds in 

elephant droppings obtained by Mühlenberg et al. (1993), who collected droppings 

monthly in 1992. They spread the droppings on beds in clearings were they kept the 

droppings free of invading vegetation and counted the total number and species of 

seedlings that germinated. 

RESULTS 

Forest elephants in the Bossematié Forest ate the leaves or twigs of 112 species, 

the bark of 21, the fruits of 54 and the bulbs of 3 species, which accounts for a total of 

449

450 

MAMMALIA 

147 plants eaten (Table 1). These 147 plants represented 30 % of the 526 spermatophytes 

identified in the Bossematié Forest. 

TABLE 1. – Frequency of food plants in the diet and frequency of germination in elephant droppings of the 

Bossematié Forest Reserve (1 : rarely; 2 : mean; 3 : often; in brackets seeds in dropping but germi- 

nation not observed ; * : only found by Mühlenberg et al. (1993) ; + : wind distributed species that germinated 

in elephant droppings). Scientific names following Hutchinson and Dalziel (1954-1972). 

Trees of forestry interest 

Leaves/ 

Bark Tubers 

twigs 

Fruits 

Seedlings Light 

in dropping requirements 

Life 

form 

Albizia ferruginea 3 light tree 

Albizia zygia 3 light tree 

Alstonia boonei 1 + intermediate tree 

Aningeria robusta 3 shade tree 

Antiaris africana 1 intermediate tree 

Bombax brevicuspe 3 intermediate tree 

Ceiba pentandra 1 + light tree 

Celtis adolfi-friderici 1 1* shade tree 

Celtis mildbraedii 3 1 1 intermediate tree 

Celtis zenkeri 3 intermediate tree 

Chlorophora excelsa et regia 3 3* both shade. trees 

Chrysophyllum subnudum 1 intermediate tree 

Chrysophyllum sp. 1 (1) tree 

Combretedendron macrocarpum 1 3 shade tree 

Copaifera salikounda 1 shade tree 

Detarium senegalense 1 (1) light tree 

Discoglypremna caloneura 3 1 1* light tree 

Distemonanthus benthamianus 1 shade tree 

Entandrophragma angolense 1 1 shade tree 

Fagara macrophylla 1 intermediate tree 

Funtumia africana 1 + intermediate tree 

Guarea cedrata 2 shade tree 

Guibourtia ehie 3 shade tree 

Homalium aylmeri 1 shade tree 

Khaya anthotheca 1 1 shade tree 

Khaya grandifoliola 1 1 shade tree 

Khaya ivorensis 1 1 shade tree 

Klainedoxa gabonensis 3 (1) shade tree 

Lannea welwitschii 1 3 1 1 light tree 

Lovoa trichilioides 1 shade tree 

Nesogordonia papaverifera 3 + intermediate tree 

Parinari excelsa 3 3* shade tree 

Parkia bicolor 3 shade tree 

Piptadeniastrum africanum 2 1 shade tree 

Pterygota macrocarpa 2 intermediate tree 

Pycnanthus angolensis 3 1 (1) intermediate tree 

Ricinodendron heudelotii 3 3 2 light tree 

Sterculia oblonga 1 shade tree 

Sterculia rhinopetala 1 (1) intermediate tree 

Sterculia tragacantha 3 intermediate tree 

Terminalia ivorensis 1 3 shade tree 

Terminalia superba 2 1 + shade tree 

Tieghemella heckelii 1 1 (1) shade tree 

Triplochiton scleroxylon + tree


Trees without forestry interest Leaves/ 

Bark Tubers 

twigs 

Fruits 

Seedlings 

in dropping 

Light 

requirements 

Life 

form 

Albizia adianthifolia 3 1 light tree 

Alchornea cordifolia 3 light tree 

Anthocleista nobilis 1 light tree 

Antiaris welwitschii 1 intermediate tree 

Antrocaryon micraster 1 intermediate tree 

Balanites wilsoniana 1 3 3* intermediate tree 

Baphia nitida 2 light tree 

Blighia sapida 1 intermediate tree 

Blighia unijugata 2 intermediate tree 

Bosqueia angolensis 1 1 shade tree 

Breviea leptosperma 1 3 1* intermediate tree 

Bridelia grandis 3 3 intermediate tree 

Calpocalyx brevibracteatus 1 intermediate tree 

Carapa procera 2 1 light tree 

Cecropia peltata 3 light tree 

Cleistopholis patens 3 1 1* light tree 

Cola gigantea 1 1* intermediate tree 

Craterispermum caudatum 3 shade tree 

Diospyros viridicans 1 intermediate tree 

Duboscia viridiflora 3 3 shade tree 

Elaeis guinuensis 1 1 light tree 

Erythrina mildbraedii 1 light tree 

Ficus capensis 3 light tree 

Ficus exasperata 3 3 1 1* light tree 

Ficus goliath 1 1* tree 

Ficus mucuso 3 3 1* tree 

Grewia malacocarpa 2 light tree 

Grewia sp. 1 (1) tree 

Guarea thompsonii 2 shade tree 

Hannoa klaineana 1 intermediate tree 

Hymenostegia afzelii 3 light tree 

Hymenostegia aubrevillei 1 tree 

Irvingia gabonensis 3 2 shade tree 

Irvingia grandifolia 3 (3) shade tree 

Kigelia africana 1 1* light tree 

Macaranga spinosa 3 light tree 

Maesobotrya barteri 1 light tree 

Maesopsis eminii 1 1* light tree 

Majidea fosteri 1 intermediate tree 

Mammea africana 2 (2) shade tree 

Microdesmis keayana 1 shade tree 

Musanga cecropioides 2 2 1 1 light tree 

Myrianthus arboreus 3 3 3 light tree 

Omphalocarpum elatum 1 1 (1) intermediate tree 

Panda oleosa 2 (3) shade tree 

Placodiscus boya 3 light tree 

Tetrapleura tetraptera 3 3 3 light tree 

Treculia africana 3 3 shade tree 

Trichilia martineaui 1 intermediate tree 

Trichilia megalantha 1 intermediate tree 

Trichilia monadelpha 2 light tree 

Xylopia quintasii 1 light tree 

451

452 

MAMMALIA 

Other plants 

Leaves/ 

Bark Tubers 

twigs 

Fruits 

Seedlings Light 

in dropping requirements 

Life 

form 

Acacia pennata 3 1 light shrub 

Aframomum exscapum 1 1 light herb 

Aframomum sceptrum 1 1 light herb 

Ancistrophyllum secundiflorum 1 light liana 

Caloncoba gilgiana 1 light shrub 

Calycobolus africanus 1 light liana 

Capsicum frutescens 1 1* light herb 

Carica papaya 3 3 1 light herb 

Cephaelis peduncularis 1 light shrub 

Chromolaena odorata 1 light herb 

Cissus sp. 1 2 liana 

Clerodendrum formicarum 1 light liana 

Cola caricaefolia 1 intermediate shrub 

Commelina capitata 1 light herb 

Cordia vignei 1 1* shade shrub 

Costus afer 2 2 1 light herb 

Cucumis sp. 1 3 3 3 liana 

Cucumis sp. 2 1 1 liana 

Desplatsia chrysochlamys 3 3 light shrub 

Dioscorea minutiflora 3 light liana 

Dioscorea praehensilis 1 intermediate liana 

Dioscorea preussii 1 light liana 

Griffonia simplicifolia 3 light shrub 

Hippocratea vignei 3 light liana 

Lagenaria breviflora 1 3 3 light liana 

Landolphia hirsuta 2 1 1* light liana 

Mallotus oppositifolius 2 light shrub 

Maranthochloa leucanta 1 light herb 

Massularia acuminata 1 light shrub 

Mezoneurum benthamianum 1 3 light liana 

Millettia zechiana 1 shrub 

Momordica angustisepala 1 light liana 

Musa sp. 3 3 herb 

Oncinotis gracilis 1 light liana 

Palisota hirsuta 3 light herb 

Parquetina nigrescens 1 light liana 

Paullinia pinnata 2 light liana 

Phyllanthus discoideus 3 light shrub 

Platysepalum hirsutum 1 light liana 

Premna angolensis 2 light shrub 

Salacia erecta 1 light shrub 

Solanum erianthum 2 1 1 herb 

Solanum torvum 1 light herb 

Sorghum arundinaceum 1 1 1 light grass 

Strophanthus gratus 1 intermediate liana 

Strophanthus hispidus 1 light liana 

Strychnos aculeata 1 light liana 

Strychnos sp. 1 1 liana 

Telfairia occidentalis 1 (1) intermediate liana 

Trema orientalis 3 3 3 3 light shrub 

Urera obovata 1 light liana


Elephants in the Bossematié Forest Reserve selected leaves or twigs, fruits, and 

bark from trees and avoided these parts from shrubs (Fig. 1). They also consumed 

leaves or stems from herbs and grass less than expected whereas they did not prefer or 

avoid lianas. The elephants only consumed one species of grass (Sorghum arundinaceum), 

probably because the 20 grass species in our study area do not cover extensive 

areas. The plants from which elephants ate the bulbs were light tolerant liana species 

(Table 1). Light tolerant species dominate in the Bossematié Forest Reserve but forest 

elephants ate more fruits and bark of shade tolerant species than their occurrence in the 

forest led us to expect (Fig. 2). However, we found no selection or avoidance of leaves 

or twigs of shade tolerant or light tolerant species. 

Percentage of species 

80% 

60% 

40% 

20% 

0% 

P

454 

Percentage of species 

60% 

40% 

20% 

0% 

P=0.604 

P=0.043 

P

Seedlings per dropping 

5 

4 

3 

2 

1 

0 


0 2 4 6 8 

Age of dropping in months 

artificial sunny natural sunny natural shady 

Fig. 3. – Mean numbers of seedlings per elephant dropping under artificial sunny conditions, 

natural sunny conditions and natural shady conditions in relation to the age of a dropping. 

There is a seasonal pattern in the number of species that germinate in elephants 

droppings (Fig. 4), which seems to follow more the pattern of precipitation than the 

pattern of trees that fruit in each month. Most species germinate in droppings that elephants 

excreted in March-May and September. As most seedlings germinate 2 months 

after the excretion of droppings, the maximum germination was from May-July and in 

November, just after the months of maximal precipitation. 

Number of species 

50 

40 

30 

20 

10 

0 

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 

Month 

plants dispersed rainfall fruiting trees 

Fig. 4. – Seasonal distribution of seedling germination in elephant droppings in relation to the 

monthly numbers of trees fruiting and the monthly precipitation for 1992 and 1993 (pooled 

data of Mühlenberg et al. (1993) and this study). 

250 

200 

150 

100 

50 

0 

Mean rainfall in mm 

455

456 

MAMMALIA 

DISCUSSION 

It is generally admitted that forest elephants prefer secondary to primary parts of 

rain forests (Prins and Reitsma 1989 ; Barnes et al. 1991 ; Struhsaker et al. 1996). In 

our heavily exploited study area, however, elephants selected shade tolerant trees from 

primary forests for their diet, probably because of the shortage of primary forest species, 

which provide fruits. These results suggest that the Bossematié Forest Reserve 

offered a higher proportion of light tolerant species than that sought by the forest elephants. 

Vanleeuwe and Gautier-Hion (1998) also observed that forest elephants intensively 

used forest parts with a closed canopy and a high density of fruit trees whereas 

they avoided Marantaceae forests with an open canopy and low numbers of fruit trees. 

It therefore appears that the canopy cover in the Bossematié Forest Reserve was too 

open for the needs of forest elephants. The assumption that forest elephants would 

always prefer secondary forest may not be valid for all parts of Africa. We assume that 

in West African forests, where elephants eat considerably more fruit species than in 

Central Africa (Alexandre 1978 ; Short 1981 ; Martin 1982 ; Tchamba and Seme 1993 ; 

White et al. 1993 ; Feer 1995 ; Maurois et al. 1997), elephants would select primary 

forest parts more than they do in Central Africa. The effect of logging and habitat fragmentation 

may therefore be more detrimental to forest elephant populations in West 

than in Central Africa. Barnes (1999) predicted that West African elephants would 

soon only survive in protected areas. This emphasises the need to protect and support 

the natural regeneration of West African forests in which elephants still live. 

Our study of the germination success under natural conditions was small scale, so 

its results can only provide some hints. The germination rate under natural conditions 

in our study was lower than under artificial conditions in other studies : 14.5 young 

plants per dropping in Ghana (Lieberman et al. 1987) and 6.2 young plants per drop- 

ping in Zaire (Brahmachary 1980). We expect that predation of the seeds by rodents 

and other animals is an important factor in the forest (especially in the dense, shady 

forest). In addition, under natural conditions, the numbers of species were lower than 

under artificial conditions, maybe because seeds became exposed to light during the 

transport of the droppings. We conclude that artificial conditions should not be used to 

estimate a natural germination rate. 

The germination rate was higher under sunny conditions than under shady conditions. 

We expect therefore that the more open a forest is, the higher the germination 

success should be. In the Bossematié Forest, the number of species germinating in elephant 

droppings is at its highest in droppings excreted in March-May and September 

(Fig. 4) and the number of young plants in the droppings is higher in April and September 

than during other months (Mühlenberg et al. 1993). The germination rate found 

during our study should therefore be similar to that found in droppings left between 

March and May. The survival rate of the young plants was low after 9 months (9 %) 

because of the long dry season during our study. Seedlings of species that fruit from 

March to May after the dry season probably have higher survival rates. Each of the 

estimated 30 forest elephants in the Bossematié Forest Reserve (Theuerkauf et al. 

2000) produces 17.5 droppings per day (Theuerkauf and Ellenberg 2000), amounting to 

about 200,000 droppings per year in the study area (220 km²). When using the germination 

rates found in this study of 0.9 seedlings per dropping in the shade and 4.8 seedlings 

per dropping in the sun, about 800-4200 seedlings per km² would germinate per 

year in elephant droppings. The digestion of African elephants lasts between 21 and 

46 hours (Rees 1982) and its daily movements in the Bossematié Forest are about 6 km


(Theuerkauf and Ellenberg 2000), which allows us to estimate a maximal distance of 

seed dispersal of 5 to 12 km. Van Hoven et al. (1981) discovered that grass has a faster 

transit time in the intestine than leaves. We assume that the distance over which elephants 

disperse seeds in Bossematié is longer as they eat little grass. As the reserve is 

small (diameter is about 15 km), it is likely that the elephants are able to disperse seeds 

in all areas of the forest. Provided enough seedlings survive, this would accelerate the 

natural regeneration of the heavily exploited forest. 

Although only 39 identified species germinated in elephant droppings in the Bossematié 

Forest Reserve, we assume that the elephants can disseminate at least 66 species. 

This assumption is based on the fact that the seeds of 12 species, known to germinate 

in elephant droppings from other forests, were present in elephant droppings 

(Table 1) and that another 15 undetermined species germinated in elephant droppings 

(Mühlenberg et al. 1993). These 66 species, potentially dispersed by the elephants in 

the Bossematié Forest, represent more than 10 % of spermatophytes identified in the 

study area. Forest elephants can even disperse 30-50 % of fruit bearing plant species 

(Alexandre 1978 ; Gautier-Hion et al. 1985 ; Feer 1995). Feer (1995) assumed that the 

forest elephants are the exclusive disperser of ten species. Many species, which are dispersed 

by forest elephants, are also dispersed by other animal groups like birds, monkeys 

or rodents (Gautier-Hion et al. 1985). It may be that only a few species would 

disappear from a forest as direct consequence of the loss of elephants (Hawthorne and 

Parren 2000). However, elephants play an irreplaceable role as no other species dis- 

perse as many plant species in such large quantities. The role of elephants in the study 

area should now be even greater as the elephants population grew in the five years 

after this study at a rate of 13-21 % yearly (Theuerkauf et al. 2000). We therefore 

consider the forest elephant as important in the speeding up of natural regeneration and 

rehabilitation of rare species in the Bossematié Forest Reserve. 

ACKNOWLEDGEMENTS 

This study was part of the pilot project « Réhabilitation de la Forêt Classée de Bossematié » 

under the programme « Réhabilitation des forêts à l’est de la Côte d’Ivoire », a collaboration between 

the « Société de Développement des Plantations Forestières (SODEFOR)», the « Deutsche 

Gesellschaft für Technische Zusammenarbeit (GTZ) » and the « Kreditanstalt für Wiederaufbau 

(KfW)». We thank H. J. Wöll, H. Fickinger, N. Seabé and the SODEFOR team who took care of 

financial and logistical aspects, P. Ehouman, V. Bamba, K. Koffi, A. Koffi Kouamé and 

J. Somda for identifying some of the plants, L. Aké Assi (University of Abidjan) for confirming 

doubtful identifications, M. Mühlenberg, J. Slowik, H. Plachter and the late H. Remmert for cooperation, 

and S. Rouys for proofreading. 

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