Conservation of biodiversity in a relic forest in Benin
– an overview
Peter Nagel, Brice Sinsin, and Ralf Peveling
Der Lama-Wald ist einer der letzten Reste des westafrikanischen Wald/Savan
nen-Mosaiks, des ‘Dahomey Gap’. Er enthält natürlichen Wald sowie Plantagen
und hat den Schutzstatus einer “Forêt classée”. Im vorliegenden Artikel geben wir
einen Überblick über ein laufendes Gemeinschaftsforschungsprojekt, das sich auf
die Erhaltung der Biodiversität im Lama-Wald konzentriert. Das Projekt basiert
auf der Annahme, dass Erhaltungs- und Managementstrategien auf der Basis des
45/2 2004 S. 125-137
Adresse der Autoren: Prof. Dr. Peter Nagel and Privatdozent Dr. Ralf Peveling, Institut
für Natur-, Landschafts- und Umweltschutz (NLU) / Biogeographie (Institute of Environ
mental Sciences, Biogeography), Universität Basel, St. Johanns-Vorstadt 10, CH-4056
Basel. Prof. Dr. Brice Sinsin, Faculté des Sciences Agronomiques, Université d’Abo
mey-Calavi, 01 BP 526 Cotonou, Bénin
Lama forest is one of the last remnants of the West African forest/savannah mosaic
known as the Dahomey Gap. It comprises natural forest and forest plantations and
has the protection status of a classified forest. In the present article, we give an
overview of an ongoing research partnership project focusing on the conservation
of biodiversity in Lama forest. The project is based on the assumption that conser-
vation and management strategies must be founded on an understanding of both
structural and functional ecological traits. It comprises studies on the biodiversity
of arthropods, a group which has received little attention as yet in tropical biodiver-
sity assessments, as well as studies on key ecological processes such as the break-
down of litter. Emphasis is laid on the relationship and interaction between natural
forest and plantations. With respect to ecological as well as biogeographical pecu-
liarities, we found evidence of the importance of Lama forest for biodiversity con-
servation in Benin. Specifically, our study elucidated the role of anthropogenic fo-
rests as buffer zones, migration corridors and surrogate habitats for rare forest ani
mals. Important insight has been gained with regard to the management and con
servation of isolated biodiversity resources in Benin and other African countries.
Verständnisses der strukturellen und funktionalen ökologischen Merkmale entwi
ckelt werden müssen. Es umfasst Untersuchungen über die Biodiversität der Ar
thropoden, einer Tiergruppe, der bis jetzt in tropischen Biodiversitätsuntersuchun
gen eher weniger Aufmerksamkeit geschenkt wurde, sowie Untersuchungen über
ökologische Schlüsselprozesse wie den Streuabbau. Hauptgewicht wird auf die Be
ziehung und die Interaktion zwischen natürlichem Wald und Plantagen gelegt. In
Bezug auf ökologische wie biogeographische Besonderheiten fanden wirBelegefür
die Bedeutung des Lama-Waldes für die Erhaltung der Biodiversität in Benin. Ins
besondere klärte unsere Studie die Rolle der anthropogenen Wälder als Pufferzo
nen, Migrationskorridore und Ersatzlebensräume für seltene Waldtiere auf. Wich
tige Einblicke hinsichtlich des Managements und der Erhaltung isolierter Biodiver
sitätsressourcen in Benin und in anderen afrikanischen Ländern wurden gewon
The Dahomey Gap is a zone of low rainfall separating the western and eastern part
of the humid Guineo-Congolean evergreen and semi-evergreen forests of West and
West-central Africa, extending along the coast from Takoradi in Ghana to Cotonou
in Benin (L’Hôte & Mahé 1996). The annual precipitation ranges from 900 to
1,500 mm. With respect to the natural vegetation expected under the prevailing cli-
mate, Sayer (1992) postulated a spatial dominance of semi-deciduous forest, inter-
spersed with tracts of denser vegetation such as riverine forest as well as patches of
swamp forest and lowland evergreen forest. This implies that the present savannah
vegetation is of anthropogenic origin. During the humid phase in the Holocene, the
area was presumably covered with evergreen lowland rainforest (cf., Anhuf 1994).
There are several examples of disjunction of plants and animals along the Dahomey
Gap (Knapp 1973, Kingdon 1990, Schiøtz 1999). For some West and West-central
African species, the Dahomey Gap demarcates the eastern and western limit, re
spectively, of their present range. This suggests that extant patches of natural forest
in Southern Benin are remnants of the former semi-evergreen lowland forest that
has been largely destroyed by humans. For species distributed in the lowland rainfo
rest during the Holocene, these forest remnants are important refuges. Likewise, it
is likely that they serve as stepping stones for species moving between the western
and eastern rainforest belts. Of the forest remnants in southern Benin, Lama forest
represents one of the largest tracts.
With few exceptions (e.g., Sayer 1992), the possible role of Lama forest for
biodiversity conservation has not been acknowledged adequately by the interna
tional conservation community (IUCN 1987, 1991). Even though, national institu
tions (ONAB 1992) and development agencies (German Development Agency and
German Development Bank), consider Lama forest a priority conservation area, de
spite its small size. Preliminary biodiversity inventories were conducted under the
auspices and with the financial aid of these organizations (Emrich et al. 1999).
However, the studies were confined to natural forest in the centre, the so-called
Noyau central. While the importance of plantation forests for the conservation of
wildlife and as nuclei for the regeneration of natural forest has been demonstrated
elsewhere in the tropics, confirmation from West Africa is still pending.
The overall goal of the present, ongoing study is the conservation and enhance
ment of biodiversity through an improved management of natural and plantation
forests in Lama forest. A central theme in this research is the ecological and
biogeographical role of plantations within an overall conservation strategy for
Southern Benin. To this end, our study focuses on the functional importance of
biodiversity as well as on key ecological processes. The project is conducted within
the scope of a research partnership between the Faculté des Sciences Agrono
miques, University of Abomey-Calavi, Benin, and the Institut für Natur-,
Landschafts- und Umweltschutz (NLU) / Biogeographie, University of Basel.
2 Geographical and historical characteristics of the study site
Lama forest is situated at the northern limit of the central part of the Dépression
médiane (between 6°55.8–58.8’N and 2°4.2–10.8’E). The altitude ranges between
40–80 m above sea level, and the average annual rainfall is about 1,200 mm. A large
rainy season from April to July is followed by a short dry season from August to
September, followed by a second, shorter rainy season from October to November
and the large dry season from December to March. During the rainy season, the soil
may become temporarily waterlogged which restricts the agricultural use. The
Tab. 1 Recent history of the development of Lama forest (modified after Emrich et al.
Year Total forest cover (ha)
1946 16,250 11,000 Gazetted as a forêt classée;
colonization by autochthonous ethnic
groups (Fon, Aïzo)
mid 1950s Immigration of allochthonous ethnic
groups (Holli and Aïzo from other
parts of Benin)
1960 10,800 Afforestation with exotic timber
species, mainly teak
1972 6,700 Increased immigration of Holli
1987 4,500 1,900 Full ban on logging of natural forest;
reforestation in the Noyau central;
resettlement of Holli people in two
prevailing soil type is a black cotton soil (vertisol) rich in humus and young clay de
posits. The vernacular name Kô is synonymous with Lama, a word of Portuguese
origin meaning mud.
Of the different ethnic groups living at Lama forest, only the Holli tribe – immi
grants from eastern Benin – are specialized in the cultivation of black cotton soil.
Until their resettlement in 1988, about 1,200 families practiced small-scale shifting
agriculture within the Noyau central. During the preceding five years, the mean an
nual deforestation was 400 ha, compared to an overall annual loss of natural forest
of 300 ha between 1946–1988. The former Holli land use system is reflected in the
present structure of the Noyau central which is composed of a mosaic of fallow land
of different age and successional stage, secondary forest and patches of primary
Fig. 1 Outline map of the Lama forest (from Lachat et al. 2004a).
NC = Noyau central T = Teak plantations,
FP = Fuelwood plantations S = Settlements, small-scale agriculture,
IF = Isolated forest fragments
Except for teak forests planted in the early sixties (1963–1965) in the northern
and southern part of the area, teak plantations were established 1985–1996, enclos
ing the Noyau central nearly entirely. In the south-western part, fuelwood forests
were planted to satisfy the demand of the local population for firewood, and to re
duce the pressure on remnant natural resources. Open canopy forest patches, clear
ings and former farmland within the Noyau central are encroached with thickets of
Chromolaena odorata, an alien, invasive species of neotropical origin. Enrichment
plantings with native forest species were established in these open areas in order to
assist and accelerate the regeneration of natural forest.
In view of dramatically diminishing forest coverage and ever increasing defor-
estation rates, a new forest management and protection project was implemented in
1987/88. All settlers were banned from the Noyau central and resettled in neigh-
bouring agro-forestry schemes where they were granted housing and agricultural
land. Moreover, education and health services were provided, and new opportuni-
ties to work in the forest sector were created. The idea was to convert forest users
into forest conservationists. Other measures taken to support local communities
were an improvement of the infrastructure (road network), the establishment of tree
nurseries, and the construction of a saw-mill for the processing of teak and produc
tion of parquet floor tiles for the global market. For educational purposes, an eco
logical trail was established in the Noyau central in 1995.
3 Implementation of research
To evaluate the role of teak and fuelwood plantations for the protection of the natu
ral forest remnants, we focused initially on the following areas of research, (1) the
diversity and composition of arthropod assemblages, in particular detritivorous and
xylophagous species, (2) the niche differentiation of key ecological groups, and (3)
the breakdown of litter and dead wood in relation to forest type.
Access into the forest is possible by a system of parallel trails (layons) running
from East to West at a distance of one kilometre from each other. We selected four
replicate sites in each of nine different forest types. Within the Noyau central, these
Tab. 2 Forest cover in Lama forest in 2000 (modified after Emrich et al. 1999)
Area Total surface (ha) Composition
Noyau central 4,800 1,900 ha natural forest
1,200 ha degraded forest of different successional stages
1,400 ha fallow land of different successional stages,
covered mainly with Chromolaena odorata
300 ha teak (Tectona grandis) and grey teak
(Gmelina arborea) plantations
Plantations 9,000 5,600 ha teak plantations
1,400 ha grey teak plantations, partly mixed with teak,
Eucalyptus spp., Acacia sp. and other species
2,400 ha firewood plantations (mainly Senna siamea)
Other 2,400 Agriculture, settlements
comprise (1) semi-deciduous forest (primary forest), (2) seasonally flooded
Cynometra megalophylla lowland forest (primary forest), (3) Anogeissus leiocarpa
dry forest (secondary forest), (4) abandoned settlements (secondary forest) and (5)
perennial Chromolaena odorata thickets. Outside of the Noyau central, we studied
(6) old teak plantations, (7) young teak plantations, (8) fuelwood plantations and (9)
isolated small forest islands.
4.1 Geographical Information System
Specht (2002) integrated a geo-ecological catalogue of maps into a Geographical
Information System (GIS). Emphasis was put on the classification of the vegetation,
using multitemporal Landsat 7 data. The exact delimitation of the main vegetation
types is an important basis for both the interpretation of ecological and biodiversity
data as well as for the management of Lama forest. Moreover, possible dispersal
and migration corridors as well as suitable habitats of animals and plants can be de
Fig. 2 Dry season aspect of Lama forest (teak plantations, left, and semi-deciduous forest,
right). Photo: T. Lachat
4.2 Biodiversity studies
4.2.1 Vegetation and flora
The natural vegetation of the Lama depression is classified as a dense
semi-deciduous forest. Afzelia africana und Ceiba pentandra are dominant, emer
gent species of the uppermost stratum of this forest type, while Diospyros
mespiliformis, Dialium guineense and Mimusops andongensis represent the lower
tree stratum. Disturbed secondary forest is often characterized by Anogeissus
leiocarpa. The vegetation in seasonally waterlogged areas is dominated by large in
dividuals of Cynometra megalophylla (Emrich et al. 1999).
The most abundant trees are Dialium guineense, Diospyros mespiliformis,
Albizia zygia, Afzelia africana, Khaya senegalensis and Anogeissus leiocarpa. The
last three species have their main distribution in the drier Sudan zone further to the
North. Two tree species new to Benin were recorded in a recent study. More than 15
species have been listed in the Red List of threatened plants (Emrich et al. 1999).
4.2.2 Arthropod diversity and faunistics
Preliminary evidence was found that the Lama forest and probably other forest rem-
nants of the Dahomey gap are home to endemic Coleoptera and Lepidoptera species
(Goergen 2003). Furthermore, about half of the 83 butterfly species recorded so far
are new to Benin (Fermon & Schulze 1998).
By using a combination of different types of traps (pitfall, Malaise, window and
light traps) we found seven different myrmecophilous ant nest beetles (Carabidae:
Paussini), six of which are stenoecious, typical forest species (Nagel 2003). It is re-
markable that these are extremely rare species. Even though most specimens were
sampled in the Noyau central, we also found some specimens in old teak plantations
and isolated forest fragments.
Lachat et al. (2004a) demonstrated that arthropod diversity was similar among
natural, degraded and secondary forests of the Noyau central, whereas isolated for
est fragments differed from all other forest types. Species richness was lowest in
young teak and fuelwood plantations, highest in old teak plantations and forest is
lands and intermediate in the different forest types within the Noyau central. A rare
ground beetle, Hoplolenus obesus (Murray) (Carabidae: Oodini), was identified as
an indicator species of old teak plantations, suggesting that anthropogenic forest
ecosystems may have a role to play in biodiversity conservation.
Several mammals are frequently encountered in Lama forest, e.g., the common
cusimance (Crossarchus obscurus), Maxwell’s duiker (Cephalophus maxwelli),
red river hog (Potamochoerus porcus) and bushbuck (Tragelaphus scriptus). The
most conspicuous monkeys are the three guenons mona, red-bellied monkey and
vervet monkey (Kassa & Sinsin 2003). It has been suggested that colobus monkeys,
including Colobus vellerosus, have disappeared from Lama forest (Matsuda 1995).
However, new sightings have been recorded recently by our research team.
Rare and threatened forest ungulates include the sitatunga (Tragelaphus
spekei), the royal antelope (Neotragus pygmaeus), the black duiker (Cephalophus
niger) und the yellow-backed duiker (C. silvicultor)(Kassa & Sinsin 2003). It is
also possible that the Kintampo rope squirrel (Funisciurus substriatus), an endemic
of the Dahomey Gap, occurs in Lama forest (Refisch 1998).
The flagship species of the Lama forest is the red-bellied monkey
(Cercopithecus erythrogaster erythrogaster). This subspecies is endemic to the Da-
homey Gap and Benin. Its former range in Southern Benin extended from the valley
of the Couffo river to the Nigerian border. Due to hunting and habitat destruction,
surviving populations now seem to be restricted to sacred groves, equally small
swamp forests (e.g., Lokoli forest) and humid forest relics, of which Lama forest is
the largest (Sinsin et al. 2002, Sinsin & Assogbadjo 2002).
There are indications of dispersal movements among these habitat fragments. In
most cases, the red-bellied guenon is associated with mona monkeys. This affords
mutual benefits in terms of predator avoidance and forage location. During the dry
season, the diet of red-bellied guenons consists of immature fruits of the cotton tree,
providing not only food but also water which is a scarce resource during this time of
the year (Nobime & Sinsin 2003). A pilot study on the feasibility of radio telemetry
and habituation techniques for studying the behavioural biology and ecology of
red-bellied monkeys showed that both methods are difficult because of the timidity
and shyness of this species (Altherr 2003).
Fig. 3 Bush meat (duiker), confiscated by a forestry officer. Photo: P. Nagel
4.2.4 Birds and reptiles
Fifteen forest bird species in Benin have been observed only in the Lama forest thus
far, including the white-crested hornbill (Tockus albocristatus), the crested Guinea
fowl (Guttera pucherani edouardi), the western bronze-naped pigeon (Columba
iriditorques) and the purple-headed glossy starling (Lamprotornis purpureiceps).
Densities of species such as the bristle-bill (Bleda syndactyla) seem to be at their
lowest limit to maintain viable populations (Waltert 1998).
Among the reptiles, there are interesting species such as Python regius and Py
thon sebae (Daouda 1998). A more recent survey found 44 species, including a new
chameleon species and probably a new gecko species (Ullenbruch 2003; Ullen
bruch et al. submitted).
4.3 Functional ecological studies
The litterbag technique was used to study the breakdown of leaf litter in different fo
rest types (Attignon et al. 2004). Decay rates in semi-deciduous forest were signifi
cantly higher than in plantation forests. Moreover, natural leaf litter (afzelia, Afzelia
africana, and cotton tree, Ceiba pentandra) degraded faster than exotic litter (teak,
Tectona grandis, and cassia, Senna siamea). This was related to the activity of lit
ter-dwelling invertebrates, suggesting that their role in litter breakdown and nu
trient-cycling must be considered in forest management. Several studies have been
initiated to analyse decomposer communities and soil quality in different types of
natural and plantation forests. A synthesis of these ongoing studies, however, is still
Fig. 4 The flagship species of Lama forest: red-bellied monkey (Cercopithecus e. erythro
gaster). Photo: G. Altherr
pending. Thus far, there are indications that earthworm relative abundance is higher
in natural than in plantation forests (Weibel 2003), whereas the contrary is true for
termites (Attignon et al. submitted a,b).
From 2002–2004, emergence trap studies were conducted to study the associa
tion of beetles with dead wood (Cakpo 2003, Lachat et al. 2004b). Again, final re
sults cannot be presented as yet. Preliminary evidence suggests that the saproxylic
beetle fauna responds very specifically to the type of dead wood and the degree and
state of its decomposition. From these studies we expect to gain general insight into
the ecological significance of dead wood in these types of tropical forests.
All investigations of the composition and distribution of biodiversity resources in
Lama forest conducted so far demonstrated the unique biogeographical and ecolo
gical status of the Noyau central. Here, rare Dahomey Gap endemics such as the
red-bellied guenon found one of their last refuges in Benin. Moreover, numerous
rainforest-adapted species occur only in Lama forest, the largest continuous tract of
dense natural forest in southern Benin. For West African forest species such as the
royal antelope, Lama forest represents the easternmost limit of their current range.
These examples underline the importance of this last remnant of dense
semi-deciduous forest and its characteristic fauna for biodiversity conservation in
Due to a lack of surface water during the dry season, species in need of water
may have to conduct seasonal movements and migrations into more humid areas
such as the Lokoli swamp forest. These movements can be a risk to maintaining via-
ble populations. Moreover, there is a higher poaching risk for animals moving
through settled land, even though the overall poaching pressure seems to be rela-
tively low. The plantation forests encircling the Noyau central may serve as a Cor
don sanitaire, i. e., a protection forest reducing the risk of illegal logging of natural
The paucity of litter-dwelling invertebrates in plantation forests and the resul
tant reduction in litter breakdown clearly suggest that specific forest management
programmes should aim to prevent an impoverishment of decomposer communities
in order to maintain productivity.
The present study focuses on the role of forest plantations for biodiversity con
servation in the Dahomey Gap. All evidence collected so far suggests that old teak
plantations in particular (about 40 years old) may provide suitable habitats even for
stenoecious rainforest insects. This seems to be due to the dense and species-rich
undergrowth resembling secondary forest undergrowth. It is noteworthy that even
extremely rare species were found in these plantations, as well as in isolated forest
islands. It follows that conservation programmes should aim to include certain
types of forest plantations as habitats for rare and/or threatened species. Nature con
servation and forest production – even monocultures – are not necessarily mutually
exclusive goals. Rather, both can be combined in an ecologically and economically
Few studies have investigated the role of forest plantations as wildlife habitats and
migration corridors. This even holds for mammals and birds, the best studied taxa.
Future studies should therefore aim to identify dispersal routes between Lama forest
and other forest remnants or forest-like habitats. These studies should establish the
scientific basis for the development of a protected area system. The degree of isola
tion of fragmented populations should be investigated using molecular genetic
methods, and the analysis be based on metapopulation models. Aspects of the func
tional importance of biodiversity in natural forests and plantations are as yet grossly
understudied and should receive more attention in future research in order to predict
possible risks timely.
The Lama forest reserve represents the most important element in the existing
network of remnant forest biotopes in southern Benin. It may even serve as a step
ping stone across the Dahomey Gap, even though more field evidence has to be col
lected to support this assumption. However, the existing database clearly exempli
fies the potential of Lama forest as a national heritage of regional importance. The
data collected so far even suggest the establishment of a Lama Biosphere Reserve.
Scientifically, this claim is founded on the outstanding importance of the Noyau
central, old teak plantations and forest remnants for biodiversity conservation. In
the past twenty years, the local population has been affected tremendously by con-
servation as well as forest production activities of governmental authorities and de-
velopment agencies - sometimes detrimentally, sometimes beneficially. It therefore
certainly must be integrated into a community conservation process inherent to the
designation of biosphere reserves.
The Swiss National Science Foundation (SNF) and the Swiss Agency for Develop
ment and Cooperation (SDC) are acknowledged for their financial support. We are
also very grateful for the technical support of the Office National du Bois, Cotonou,
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