Folia Geobot (2021) 56:81–95

https://doi.org/10.1007/s12224-021-09393-9

Plant diversity and conservation concerns

in a semi-deciduous rainforest in Cameroon: implications
for sustainable forest management
Jules Christian Zekeng & Jean Louis Fobane & Maginot Nganyo Heya & Reuben
Sebego & Wanda N. Mphinyane & Jean Michel Onana & Paul André Ebanga & Laurent
Florent Etoundi Menyene & Marie Marguerite Abada Mbolo

Received: 26 August 2020 / Revised: 15 April 2021 / Accepted: 6 May 2021 / Published online: 25 June 2021
# Institute of Botany, Czech Academy of Sciences 2021

Abstract Plant diversity conservation has become one sometimes neglected in Cameroon's communal forests
of the most critical objectives of forest management in because their botanical importance, and high-priority
an ecologically sustainable way. However, biodiversity species for conservation are poorly known. In the pres-
conservation and sustainable management issues are ent study, done in the Doume Communal Forest (DCF)
situated in eastern Cameroon, we explore its potential in
terms of plant diversity as well as conservation value.
J. C. Zekeng : J. L. Fobane : J. M. Onana : P. A. Ebanga : Forest inventories done in thirty 1-ha plots were
L. F. Etoundi Menyene : M. M. Abada Mbolo (*)
analysed using diversity indices, species and family
Department of Plant Biology, Faculty of Science, University of
Yaounde I, P.O. Box: 812, Yaounde, Cameroon importance values index as well as IUCN conservation
e-mail: [email protected] status. A total of 22,064 stems with a diameter ≥ 1 cm
were recorded within thirty 1-ha sampling plots, belong-
J. C. Zekeng : R. Sebego : W. N. Mphinyane
ing to 307 species, 194 genera and 72 families. The DCF
Department of Environmental Science, Faculty of Science,
University of Botswana, Private Bag UB, 0704 Gaborone, is particularly rich in members of the Malvaceae,
Botswana Annonaceae, Meliaceae and Apocynaceae. The list of
the plant species known from the DCF reaches 330
J. C. Zekeng : J. L. Fobane : L. F. Etoundi Menyene
species, with 290 identified at the species level. It was
Conservation and Sustainable Natural Resources Management found out that 49 species can be considered high-
Network, Yaounde, Cameroon priority species for conservation and that these species
include species that are rare and threatened in Camer-
oon, and even one species endemic to the country. This
J. L. Fobane
Department of Biology, Higher Teachers’ Training College, study highlights the botanical importance of communal
University of Yaounde I, P.O. Box 47, Yaounde, Cameroon forests in Cameroon and recommends that botanical
assessments of these forests are performed to contribute
to the sustainable management and conservation of their
M. Nganyo Heya
Facultad de Agronomía, Universidad Autónoma de Nuevo León biodiversity.
(UANL), Ex-Hacienda “El Canadá”, C.P. 66050 Escobedo,
Nuevo León, Mexico
Keywords biodiversity conservation . ecological
L. F. Etoundi Menyene values . endemism . rare species . threatened species .
Institute of Agricultural Research for Development (IRAD), high-priority species for conservation . Doume
Ministry of Scientific Research and Innovation (MINRESI), P.O. Communal Forest
Box: 2123, Yaounde-Messa, Cameroon
82 Zekeng et al.

Introduction institutional framework and adopting a forest law sup-

plemented through an implementing decree. Elsewhere,
Tropical forests are essential for human survival and with the forest code, the Republic of Cameroon aims to
well-being. They are at the centre of debates on climate promote sustainable management and conservation pur-
change and sustainable forest management (SFM) be- pose of its forests while encouraging all users' partici-
cause of their dual roles in climate change mitigation pation in the process of decentralized management of
and biodiversity conservation (Bodegom et al. 2009; the forest. Thus, the Republic of Cameroon decided to
Bele et al. 2015; Poorter et al. 2016; Arasa-Gisbert retrocede to the requested municipalities to manage and
et al. 2018). They also provide a wide range of other exploit forest areas for the benefit of their entire popu-
ecosystem services that contribute to people’s quality of lation. Such forests are called Communal Forests (CFs).
life (IPBES 2019). It is now globally recognized that In Cameroon, the concept of communal forestry has
forests must be managed sustainably (United Nations evolved, and in 2018, sixty-four communal forests were
1992; Kohm and Franklin 1997). Many forests are still representing an area of 1,812,150 ha or 8% of the
subject to deforestation and degradation, which has a country's total forest area (MINFOF 2018). Communal
significant negative impact on terrestrial biodiversity forests represent forest management centred on com-
and, thus, on ecosystem services closely linked to mercial wood resources with an obligation to deal with
biodiversity. the aspirations, interests and uses of local populations.
Because it is irreversible, extinction is the major Thus, the communal forest is a recent framework for
problem of the ongoing biodiversity crisis. Even though effective participatory forest management where
the conservation of biodiversity and reduction of its loss sustained tree harvesting must be combined at a local
has been reasserted by the Aichi targets for 2020 by the scale to enhance riparians' well-being (Poissonnet and
Parties to the United Nations Convention on Biological Lescuyer 2005). Like with forest management units
Diversity (CBD) after failing to meet the 2010 target (FMUs), the manager of each CF should elaborate a
(Butchart et al. 2010; CBD 2011), its loss does not seem management plan that is the keystone of sustainable
to be slowing down. Anthropogenic disturbances have forest management. It is generally revisable every fif-
resulted in a loss of species diversity, with the current teen years, depending on the legislation, whose deci-
rate of extinctions being at least 1,000 times higher than sions ratified by the company, and the administration in
natural extinction rates (De Vos et al. 2015). Therefore, charge of the forests will constitute the fundamental
the most obvious among them may be the lost opportu- elements of implementing these two key stakeholders'
nity for future resource use. Onana (2011) already noted forest policies. The management plan should be the
that some species lose their habitat and sometimes be- result of a thorough study of the forest's potential and
come extinct without being even known to science. the risks that may weigh on the forest and its functions,
With the loss of species, we lose the ultimate source and this study must have been guided by the develop-
and the basis of the ecosystems' structure and function ment possibilities of the forest (ATIBT 2007).
that support humans and all life on Earth (Mittermeier However, most of the studies carried out in CFs and
et al. 2011). Therefore, maintaining the biodiversity of FMUs of tropical forests concluded that conservation
forest ecosystems is essential to the supply of ecosystem issues and some SFM aspects are neglected to benefit
services and not less important to support their health woody resources assessment (Anonymous 2015;
and resilience (Butchart et al. 2010; Pereira et al. 2013). Tchouto et al. 2006). In most of these studies, floristic
In many respects, conservation and sustainable man- diversity is limited to tree species with a minimum
agement are local and people generally care more about diameter of exploitability for CFs and FMUs and even
the biodiversity in the place in which they live. Based on protected areas' management plans. Therefore, this ap-
this observation and since the summit of the earth in proach is not sufficient for plant diversity assessment
1992, Cameroon has made many efforts towards the because most components of the diversity of the eco-
path of conservation and SFM. These efforts has been systems, such as shrubs, small trees, lianas, herbaceous
evident through (i) the signing and ratification of the plants and epiphytic flora as well as their conservation
CBD treaty in 1994 and later the climate change treaty status, are not taken into account (Tchouto 2004). Most
such as REDD +; (ii) the implementation of internation- African countries like Cameroon have decided to estab-
al treaties signed at the national level by restructuring its lish large forest areas in protected areas as their
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 83

biodiversity conservation strategy (Mengue-Medou Cameroon's eastern region, 58 km from Abong-Mbang

2002; Muhumuza and Balkwill 2013). Terrestrial biodi- and 57 km from Bertoua. Spread over an area of 2,500
versity is too widely dispersed to allow its measurement km2, the municipality of Doume is bordered by Ndiang,
to be focused solely within strictly protected areas. Abong Mbang, Doumaintang and Angossas in the
Managed landscapes will continue to play vital roles north, south, east and southwest, respectively. Two
as buffer zones and corridors supporting protected areas blocks of the forest constitute the Doume Communal
and more generally as habitat for wild species, some of Forest (DCF) of 45,359 ha (Fig. 1).
which are likely never adequately represented within the Topographically, the region is covered by a succes-
protected area network (Dudley et al. 2005). Measure- sion of low hills with generally gentle slopes inter-
ment across the whole mosaic of land-cover types is spersed with small streams, or swampy depressions,
therefore essential, including in the area of sustainable sometimes vast (several hundred metres), but without a
use, particularly forests inside and outside protected distinct watercourse. The elevation varies from 540 to
areas. 676 m above sea level, with some particularly marked
The present study focuses on the Doume Communal summits, culminating at less than 700 m (Anonymous
Forest (DCF) situated in Cameroon's eastern region and 2015). The climate is tropical, with a wet and a dry
belonging to the Guineo-Congolese domain (Letouzey season. The mean annual temperature is 25°C and an-
1985). This forest is important for national development nual rainfall varies between 1,300 and 1,800 mm. The
and the livelihoods of about 22,763 locals (Anonymous soils are ferralitic with high sand content, high water
2015). It provides various ecosystem services, supports retention capacity and high humus content resulting
riparian people's livelihoods and harbours many endan- from the decomposition of plants and organic matter
gered animal and plant species populations. In addition (Zekeng 2020; Zekeng et al. 2020).
to supporting biodiversity, these forests also store and
absorb substantial but seldom measured quantities of Sampling design, botanical inventory
carbon (Zekeng et al. 2020). These forests are also and measurements
subject to intense pressure because of rapid population
growth, logging and hunting activities, which exert a Representative and homogeneous vegetation types were
diverse ecological impact on the forest ecosystems. selected based on physical features (e.g. topography and
Zekeng et al. (2019) have shown that the degradation altitude) and analyses of land use and land cover change
and conversion of the forests in Cameroon to other land of the study area (Zekeng et al. 2019). Thirty 1-ha (100
use types such as agroforestry systems is moving at a × 100 m) plots were established within the two blocks of
worrying speed. Therefore, with the increasing destruc- the DCF, avoiding rivers and swampy vegetation types.
tion of natural ecosystems, identifying species with high Each 1-ha (100 × 100 m) plot was subdivided into 25
conservation priorities and their habitat areas is an es- subplots of 20 m × 20 m. Five sub-quadrats of 5 m × 5 m
sential requirement for many environmental applica- were also installed at the four corners and the centre.
tions, including land use planning and landscape mon- Within each plot, complete inventories of all vascular
itoring. This study aimed to assess the plant diversity plants with a diameter at breast height (DBH) ≥ 10 cm,
and its conservation value in the DCF to demonstrate hereafter large trees, were realized. All the vascular
that communal forests deserve more attention in terms plants with DBH between 5.0 and 9.9 cm, hereafter
of biodiversity conservation and SFM. medium trees, were identified, and their DBH recorded
from thirteen plots of 20 m × 20 m of the 1-ha plot.
Stems of 1.0–4.9cm diameter, hereafter small stems,
Material and methods were recorded within the five 5 m × 5-m sub-quadrats.
If applicable, trees were measured 50 cm above the top
Study site of buttresses, or 2 cm above the deformities (Condit
1998).
The study was conducted in the moist, semi-deciduous During the fieldwork, direct identification was per-
communal forest managed by the municipality of formed at the time of tree mensuration by observing the
Doume (4°31′0″ S, 13°47′5″ W). The Doume munici- leaves, flowers, habit, slash and other diagnostic char-
pality is situated in the Upper Nyong division of acters. A unique code was attributed to unidentified
84 Zekeng et al.
960000 970000 980000 990000 1000000

Legend

500000

500000
H
! District of Doume
Villages Block 2
Road
Primary road

490000

490000
Secondary road
Communal forest
Toungrelo

Menyangwa
eléndom Kabili Mbaguempel
Konpatwa
Mbalengué Bayong V Kwèn
Ngomndouma Bayong I

480000

480000
Kongsimbang Tchandja

Oulbendamba
Kempong
Biziga
Goumbéjérong
DOUMÉ
Mbama !
H Kobila
Sibita Mala
Paki
470000

470000
Mendim
Boumpial
Bent
Nkoum

Block 1 Diaglassi
Bonando
460000

460000
Motcheboum
Spatial reference: WGS 1984, UTM zone 33 N

Source: Atlas forest, WRI, MINFOF 2019

450000

450000
0 4 125 8 250
Km

960000 970000 980000 990000 1000000

Fig. 1 Map and localization of the Doume Communal Forest, showing the two blocks (FC DOUME block 1 and 2) of it.

species for re-use during the inventory if encountered of species in a chosen community (Kent and Coker
later within the plot or locality. Also, herbarium speci- 1992) or a set of samples (Magurran 2004). Diversity
mens were collected for further checking and identifica- indices consider the number of species and whether
tion at the National Herbarium of Cameroon. A species are more or less equally abundant or if, on the
datasheet of each specimen was filled out describing contrary, one or more species dominate. Diversity was
its vegetative characters. The names of each species assessed using species richness, expressed as the
identified during the fieldwork were homogenized for Shannon-Weaver (Shannon and Weaver 1949),
synonymies and orthographic problems using several Simpson (Simpson 1949) and α-Fisher (Fisher et al.
plant databases such as the Plant List database (Hassler 1943) indices, which are the most widely used and thus
2018). The Angiosperm Phylogeny Group IV (APG; facilitate comparisons with other studies.
Byng et al. 2016) was used for families' botanical no- Species richness refers to the number of species pres-
menclature. For the Leguminosae subfamilies, the new ent, without any particular regard for the exact area or
subfamily classification based on a taxonomically com- number of individuals examined. Nevertheless, it is use-
prehensive phylogeny was used (LPWG 2017). ful to distinguish between numerical species richness, the
number of species present in a collection containing a
Data analysis specified number of individuals (Hurlbert 1971). To de-
termine whether sample size in this study was sufficient,
In ecology, diversity implies a measure of both species sample-based rarefaction curves were plotted.
number and evenness. To describe the diversity patterns The Shannon-Weaver index (Shannon and Weaver
across the Doume Communal Forest plots, alpha diver- 1949) assumes that individuals are randomly sampled or
sity indices were used, since they represent the number distributed from an indefinitely large population and that
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 85

all species are represented in the sample. The Shannon- which assumes that samples are a reasonable fit to a log-
Weaver index is computed from the following equation: series. It is independent of sample size and it describes
S
how the individuals are divided among the species. The
H 0 ¼ − ∑ pi ln pi ; α-Fisher index provides good discrimination between
i¼1 sites, it is not very sensitive to density fluctuations and it
where pi is the proportion of individuals belonging to the is normally distributed:
ith species found in a sample.
The Simpson index (Simpson 1949) is a measure of N ð1−xÞ S
the number of species present and also takes into ac- a¼ ; with x estimated from
x N
count the relative abundance of each species. It repre-
ð1−xÞ
sents the probability that two randomly selected indi- ¼ ;
viduals in the area belong to the same species. It mea- x½−lnð1−xÞ
sures how individuals were distributed among species of
a population and it is given by: where N represents the total number of individuals and S
the total number of species (Magurran 1988).
γ ¼ ∑i P2i ;
The species importance value index (IVI; Curtis and
where Pi is the proportion of individuals belonging to McIntosh 1951) and the family importance value index
the ith species found in a sample. Due to its reciprocal (FIV; Cottam and Curtis 1956) were used to describe the
character, it starts with the value of 1 and raises up to the ecological importance of species and families within the
total number of species. thirty 1-ha plots. The IVI was used to determine the
Trees diversity was also estimated using a common leading dominant taxa in the vegetation sample and
diversity index, the α-Fisher index (Fisher et al. 1943), computed using the following formulae:

Relative density ¼ ðnumber of species individuals or family individuals=total number of individualsÞ  100:

Relative frequency ¼ ðfrequency of a species=sum of all frequenciesÞ  100:

Relative dominance ¼ ðbasal area of the species or family=total basal areaÞ  100

Family relative diversity ¼ ðnumber of species in a family=total number of speciesÞ  100

IVI ¼ relative density þ relative dominance þ relative frequency;

FIV ¼ Relative dominance þ Relative density þ Relative diversity

High-priority species for conservation are defined as Onana 2013). For the establishment of this priority
species with high conservation value, such as Camer- species list, several steps were followed. Firstly, using
oon's endemic species, rare species or endangered spe- a management plant of the Doume Communal Forest
cies (Tchouto et al. 2006; Onana and Cheek 2011; and the thirty 1-ha plots field inventory, a plant checklist
86 Zekeng et al.

was generated. Secondly, to determine the threatened composition can be considered satisfactorily sampled
status of the species on this checklist, and to identify (Fig. 2). This study found that for the whole tree com-
endemic species, floras and monographs were consulted munity, the number of species per plot varied from 109
(Onana 2011; Onana and Cheek 2011), as were the to 131 species. Variation of between 93 and 116 species
IUCN (2020) red data list, the African Plant Database was found among the thirty 1-ha plot for the large tree
(2020) database and the Plant List database (Hassler diameter class. The areas sampled in each of the thirty 1-
2018). ha plots showed a variation of 43 to 61 species for
The notion of rarity is understood differently accord- medium trees and 19 to 29 species for small stems in
ing to the authors that worked on it, often according to the the DCF plots.
objectives sought (Tchouto et al. 2006; Gonmadje et al.
2012; Onana 2013). In this work, which aims to draw Diversity indices in the Doume Communal Forest
attention to policies on the management of communal
forests, it was decided to use the notion of a rarity as The Shannon-Weaver index for the whole tree
defined in Cameroon's forestry law. Indeed, the forest law community (4.05 ± 0.15) as well as for large trees
of 1994 and its application decree considered all species (3.99 ± 0.19) did not vary too much, in contrast to
with less than one tree per 100 ha rare. The major African medium trees (3.53 ± 0.27) and small stems (2.79
phytochoria (White 1979; White 1983) were used to ± 0.26). The same trends were found for the
determine the plant check-list's distribution patterns. Simpson and the Fisher-α index. However, for
Therefore, each species was assigned to one of the fol- small stems, it was found that the values of the
lowing categories: (1) Widespread (Ws) including pan- Simpson index were low (0.91 ± 0.04), with great-
African and paleotropical, (2) Guineo-Congolian (Gc), er variation (CV ≥ 3.96) among the plots than the
(3) Upper and Lower Guinea (Gu), (4) Lower Guinea other trees classes (CV ≤ 3.41). For the α-Fisher
(Lg), (5) Cameroon (Cam) and (6) Oriental Semi- index, it was found that small stems had low
deciduous Cameroon (OS-Cam). Based on the species values (17.55 ± 5.65) and more marked variation
information above, a list of high-priority species for (CV ≥ 32.17) among plots than the other tree size
conservation was produced, with preference given to taxa classes (Table 1).
that are endemic to Cameroon.
Floristic composition of Doume Communal Forest plots

Results The FIV average values of the most important

families within the whole thirty 1-ha plots are
Species richness in the Doume Communal Forest presented in Table 2. The families Malvaceae
(FIV = 28.16), Annonaceae (FIV = 24.95),
A total of 22,100 stems with a diameter of ≥ 1 cm were Meliaceae (FIV = 16.97) and Apocynaceae (FIV
recorded within the thirty 1-ha sampling plots, where = 16.92) are the four most important families with
22,064 stems belonging to 307 species were identified the highest FIV, considering the whole tree com-
and classified into 194 genera and 72 families. More munity of the 30 1-ha sample plots of DCF.
than 89% of morphospecies were identified at the spe- Moreover, these four families also appear as the
cies level, 6% at the generic level, 2% at the family families with the highest FIV in the large and
level, and 3% remained unidentified. The total number small trees classes (Table 2).
of stems registered and identified is distributed as fol- For the whole tree community, the most diverse
lows: 15,168 large trees (DBH ≥ 10 cm) belonging to families with more than fifteen species were of the
271 species, 4,567 medium trees (5 ≤ DBH < 10 cm) Malvaceae (24 species), Rubiaceae (20 species),
belonging to 242 species, and 2,010 small stems (1 ≤ Euphorbiaceae (19 species), Annonaceae (18 spe-
DBH < 5 cm) belonging to 167 species. Many species cies), Leguminosae-Detarioideae (17 species) and
were recorded to occur in more than one diameter class. Sapotaceae (17 species). However, these most di-
The rarefaction curve, showing the tree species rich- verse families are not the same when considering
ness in the thirty 1-ha sample plots rises only slowly trees by the three diameter classes. Because their
towards the end, suggesting that the forest trees species emergent or large canopy trees account for much
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 87

Fig. 2 Rarefaction curve showing species richness of the a – whole tree community and b – tree size classes for the thirty 1- ha plots of the
Doume Communal Forest.

of the basal area recorded in the plots, species of large trees (FIV = 11.38) and small stems level
the Malvaceae, Cannabaceae, Urticaceae and (FIV = 12.81; Table 2).
Leguminosae-Caesalpinioideae were the most dom- In general, among the species with the highest IVI
inant, contributing primarily to the floristic com- values for the whole tree community, some species
position of the canopy, thereby justifying their appear also at the top in one of the three diameter
high FIV at the level of large tree. The classes’ levels, a few species appear in any two of the
Annonaceae, Meliaceae, Apocynaceae and diameter classes, and some species even appear in all
Phyllanthaceae owed their high FIV values to their three diameter classes, for example Eriocoelum
density, contributing mostly to the medium trees macrocarpum and Trichilia dregeana (Table 3). The
classes with numerous individuals but exhibiting a four species with the highest IVI in the whole tree
rather low basal area; high FIV values were pro- community, which is also true for the tree census at
duced by medium trees and small stems. Other the level of large trees, were E. macrocarpum, Musanga
families, such as Leguminosae-Detarioideae, have cecropioides, Trichilia dregeana and Pycnanthus
approximately the same proportions of FIV for angolensis, belonging to the Sapindaceae, Urticaceae,

Table 1 Average values and coefficient of variation for species richness and diversity for the whole tree community and tree size classes
among the thirty 1-ha plots

Estimate parameters Whole tree community Large trees Medium trees Small stems

Average ± Standard error

Richness 121.03 ± 11.98 104.83 ± 11.96 51.63 ± 9.08 24.37 ± 4.81
Shannon-Weaver 4.05 ± 0.15 3.99 ± 0.19 3.43 ± 0.27 2.79 ± 0.26
Simpson 0.97 ± 0.01 0.97 ± 0.01 0.94 ± 0.03 0.91 ± 0.04
Fisher-α 42.24 ± 5.30 40.38 ± 6.24 30.97 ± 9.10 17.55 ± 5.65
Coefficient of variation
Richness 9.89 11.41 17.58 19.74
Shannon-Weaver 3.67 4.89 7.77 9.44
Simpson 0.84 1.27 3.41 3.96
Fisher-α 12.54 15.44 29.39 32.17
88 Zekeng et al.

Table 2 Average family importance value index of the most important families (in bold) for the whole tree community and the tree size
classes (by decreasing the whole tree community) within the 30 1-ha of the Doume Communal Forest plots

Families Average family importance value index

Whole tree community Large trees Medium trees Small stems

Malvaceae 28.16 29.14 9.74 22.22

Annonaceae 24.95 26.40 33.37 15.03
Meliaceae 16.97 16.23 21.15 26.23
Apocynaceae 16.92 16.18 20.22 34.27
Euphorbiaceae 15.96 16.18 12.20 14.15
Cannabaceae 14.72 16.18 9.94 7.71
Leguminosae-Detarioideae 11.65 11.38 8.43 12.81
Phyllanthaceae 11.64 12.94 6.14 2.76
Leguminosae-Caesalpinioideae 11.31 12.32 3.30 3.83
Urticaceae 11.15 11.92 5.52 6.26
Myristicaceae 10.26 10.69 11.23 9.48
Irvingiaceae 8.99 9.31 14.83 8.07
Sapindaceae 8.44 8.24 12.39 12.29
Moraceae 8.40 7.54 10.36 9.88
Rubiaceae 7.58 6.13 5.52 16.07
Salicaceae 6.96 6.19 10.42 22.92
Pandaceae 1.84 0.59 0.88 15.97

Meliaceae and Myristicaceae families, respectively. in various African phytochoria (16 %). Most species in
Musanga cecropioides, Sterculia rhinopetala, the list are Guineo-Congolian wide (69%) or restricted to
Terminalia superba and P. angolensis were the most the Upper and Lower Guinea (6%). However, 25 species
dominant species contributing to the canopy's floristic (9%) are only known from the Lower Guinean domain.
composition, where large trees result in high IVI values. Of these species, only Penianthus camerounensis
Whilst the species T. dregeana, E. macrocarpum, (Menispermaceae) is endemic to Cameroon (Table 4).
Homalium sp. (Salicaceae), Tabernaemontana crassa A list of 49 plant species considered species with
(Apocynaceae) and Microdesmis puberula owed their high-priority for conservation, including rare species,
high IVI values to their abundance, contributing mostly threatened species and Cameroon endemics, with infor-
to the medium class, high IVI values being provided by mation on their habit and chorology is provided. This
medium trees or small stems (Table 3). Across all the list included 38 species threatened at the global level.
thirty 1-ha sample plots, 16% of the species were rep- Moreover, it was found that heigh species threatened at
resented by only one or two individuals (51 species out the global level are not threatened at the national level. It
of all the 307 species). was also found that two species threatened at the nation-
al level were not at the global level (Table 4).
Conservation value and endemism

A plant checklist of 330 species including 23 species of Discussion

vascular plants (only trees) found in the forest manage-
ment plan and the 307 plant species encountered in the Diversity of the semi-deciduous Doume Communal
thirty 1-ha plots was made. Of these species, 291 species Forest
have been identified at the specific level.
Among the 291 species whose range was known, a A better floristic characterization of forest communities
small proportion consisted of widespread species found is based on the interpretation of diversity indices (Sonké
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 89

Table 3 Average importance value index of the most important species (in bold) for the whole tree community and the tree size classes
within the thirty 1-ha plots of the Doume Communal Forest

Species Average importance value index

Whole tree community Large trees Medium trees Small stems

Eriocoelum macrocarpum 8.39 8.15 12.72 11.57

Musanga cecropioides 8.33 9.00 2.43 1.67
Trichilia dregeana 6.95 6.35 14.91 18.92
Pycnanthus angolensis 6.60 6.90 7.12 6.19
Sterculia rhinopetala 6.59 7.22 4.11 2.43
Terminalia superba 6.31 6.96 0.78 –
Tabernaemontana crassa 5.78 5.25 9.96 19.87
Anonidium mannii 5.61 6.45 4.85 –
Celtis adolfi-friderici 5.12 5.82 3.02 1.24
Uapaca guineensis 4.95 5.54 2.92 1.44
Funtumia elastica 4.63 5.30 4.20 0.40
Desbordesia glaucescens 4.58 4.80 4.97 3.93
Homalium sp. 4.41 3.44 10.18 19.00
Polyalthia suaveolens 4.11 4.15 5.93 6.90
Annickia chlorantha 3.37 3.56 5.30 1.58
Microdesmis puberula 1.84 0.59 1.63 15.24
Rinorea sp. 1.70 1.09 3.12 7.33
Streblus usambarensis 1.63 0.47 7.78 5.87
Tabernaemontana pachysiphon 1.39 0.15 2.63 12.16
Thomandersia hensii 0.94 0.35 1.91 6.89
Octolobus spectabilis 0.83 0.31 1.47 7.40

2004; Mbolo et al. 2016), so a forest is considered rich if showed that diversity indices such as Shannon’s,
it is characterized by a Shannon diversity index greater Simpson’s and Pielou’s are only elements of measure-
than or equal to 3.5 (Kent and Coker 1992). Therefore, ment and biodiversity characterization.
the DCF plots, which at the levels of the whole tree This study showed that among the thirty 1-ha plots of
community and the large and medium tree groups had the DCF, species richness varied from 109 to 133
high values of Shannon diversity (H′ > 3.5) and species·ha−1 and that this species richness decreased
Fisher-α can, accordingly, be considered very diverse. with tree size groups. The species richness of large trees
The semi-deciduous DCF harbours a rich and diverse found in this study (104 ± 12 species·ha−1) was similar
trees flora. However, the Shannon-Weaver index of to the values of 119 ± 9 and 96 ± 10 species ha−1 found
small stems groups showed that it is not rich and diverse in Cameroon Atlantic forest of Okoroba and Yingui,
and the rarefaction species accumulation curve con- respectively (Fobane 2017), and the value of 110
firmed this. Indeed, this curve shows that the rate of species·ha−1 was found in the lowland evergreen forest
species increases with sampling effort had reached an of Ngovayang (Gonmadje 2012). However, the number
asymptote, indicating that the diversity of the DCF small of 271 species found in the thirty 1-ha plots of the semi-
stem group had been satisfactorily captured and that deciduous forest of east Cameroon was greater than the
even if the sample area increases, the diversity will not value of 207 species found in terra firme evergreen
increase. However, the Simpson index justifies the rep- forest in the Dja Biosphere Reserve in Cameroon
resentativeness of the flora by some species in terms of (Djuikouo et al. 2010) and the value of 205 species
their abundance (Sonké 2004). McElhinny et al. (2005) found in the same sites (Tabue et al. 2016). Moreover,
90 Zekeng et al.

Table 4 List of high-priority species for conservation found in the Doume Communal Forest

Family Species Chorology Habit IUCN local status IUCN global status
(Onana 2011) (IUCN 2020)

Anacardiaceae Antrocaryon micraster A. Chev. & tr LC VU

Guillaumin*
Burseraceae Dacryodes igaganga Aubrév. & Pellegr. Lg tr. sh VU VU
Chrysobalanaceae Maranthes gabunensis (Engl.) Prance* Gc tr LC LC
Clusiaceae Garcinia kola Heckel Gc tr. sh VU VU
Garcinia mannii Gc tr.sh VU VU
Combretaceae Terminalia ivorensis A. Chev.* Gu tr LC VU
Cordiaceae Cordia platythyrsa Baker Gc tr. sh LC VU
Ebenaceae Diospyros crassiflora Hiern Gc tr. sh NT VU
Leguminosae-Detarioideae Afzelia africana Pers. Ws tr VU VU
Afzelia bipindensis Harms Gc tr.sh VU VU
Afzelia pachyloba Harms Gc tr VU VU
Bikinia letestui (Pellegr.) Wieringa* Lg tr LC
Brachystegia cynometroides Harms* Lg tr LC LC
*
Brachystegia mildbraedii Harms Lg tr LC LC
Daniella ogea (Harms) Rolfe ex Holland* Gu tr LC LC
Didelotia unifoliolata J.Léonard Lg tr LC NT
Gossweilerodendron balsamiferum Gc tr. sh EN
(Vermoesen) Harms
Gossweilerodendron joveri Aubrév. Gc tr VU
Tetraberlinia bifoliolata (Harms) Gc tr LC LC
Hauman*
Leguminosae - Papilionoideae Pericopsis elata (Harms) Meeuwen* Gc tr LC EN
Pterocarpus mildbraedii Harms Ws tr. sh LC VU
Leguminosae-Caesalpinioideae Albizia ferruginea (Guill. and Perr.) Benth. Gc tr LC VU
Malvaceae Bombax brevicuspe Sprague Gc tr LC VU
Mansonia altissima A. Chevalier Gu tr. sh VU LC
Nesogordonia papaverifera (A. Cheval.) Gc tr.sh VU VU
Capuron
Pterygota bequaertii De Wild. Gc tr LC VU
Pterygota macrocarpa K. Schum. Gc tr. sh LC VU
Meliaceae Entandrophragma angolense (Welw.) C. Ws tr. sh VU VU
DC.
Entandrophragma candollei Harms Gc tr. sh LC VU
Entandrophragma cylindricum (Sprague) Gc tr. sh VU VU
Sprague
Entandrophragma utile (Dawe & Gc tr. sh VU VU
Sprague) Sprague
Khaya anthotheca (Welw.) C.DC. Gc tr VU
Khaya grandifoliola C. DC. Ws tr. sh VU
Khaya ivorensis A. Chev. Gc tr VU VU
Leplaea cedrata (A.Chev.) E.J.M.Koenen Gc tr. sh VU VU
& J.J.de Wilde
Leplaea thompsonii (Sprague & Hutch.) Gc tr. sh LC VU
E.J.M.Koenen & J.J.de Wilde
Turraeanthus africanus (Welw. ex C. Gc tr. sh LC VU
DC.) Pellegr.
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 91

Table 4 (continued)

Family Species Chorology Habit IUCN local status IUCN global status
(Onana 2011) (IUCN 2020)

Menispermaceae Penianthus camerounensis A.J.F.M. Cam sh LC

Dekker**
Ochnaceae Lophira alata Banks ex Gaertn. fil.* Gc tr VU VU
Ochnaceae Ochna calodendron Gilg & Mildbr. Gc tr.sh VU LC
Putranjivaceae Drypetes preussii (Pax) Hutch. Lg tr. sh VU VU
Rhizophoraceae Anopyxis klaineana (Pierre) Engl. Gc tr. sh NT VU
Rubiaceae Mitragyna ledermannii (K.Krause) Gc tr VU
Ridsdale
Nauclea diderrichii (De Wild. & Gc tr. sh VU VU
T.Durand) Merrill
Salicaceae Dovyalis cameroonensis Cheek & Ngolan Gc tr. sh CR
Sapotaceae Autranella congolensis (De Wild.) Gc tr LC CR
A.Chev.
Baillonella toxisperma Pierre Lg tr.sh VU VU
Chrysophyllum lacourtianum De Wild.* Gc tr LC
Tieghemella africana Pierre Gc tr EN EN

Chorology: Lg – Lower Guinea; Gc – Guineo-Congolian; Gu – Upper and Lower Guinea; IUCN status: categories of the threat as EN –
Endangered; VU – Vulnerable; LC – Least Concern and NT – Near Threatened; Habit: tr – trees and sh – shrubs; *rare species with less than
0.02 trees per ha and therefore excluded from logging in the DCF, **endemic species

this result is also higher than the value of 127 species included Eriocoelum macrocarpum, Musanga
obtained in a semi-deciduous forest of east Cameroon cecropioides, Trichilia dregeana, Pycnanthus
(Chimi et al. 2018). Previous studies in this area showed angolensis, Sterculia rhinopetala, Terminalia
that disturbance that has occurred a long time ago in- superba, Tabernaemontana crassa, etc. According to
creased species richness (Zekeng et al. 2020), and soil the importance value index, among the most species,
conditions and topography also drove this species rich- there was a high abundance of pioneer species that
ness (Zekeng 2020). Therefore, ecological factors (i.e. could indicate a more advanced level of forest degra-
rainfall, topography, disturbance and soil conditions, dation in the inventory plot. Effectively, the DCF was
etc.) might explain the difference in species richness of subject to normal and legal exploitation under the
the DCF plots compared to the studies mentioned licensing regime between 1971 and 1980. Also re-
above. ported has been the illegal exploitation in the form of
Elsewhere, it was found that the thirty 1-ha plots of wild sawing between 2009 and 2014 (Anonymous
the semi-deciduous forest host 242 medium tree species, 2015). Therefore, some plots have experienced log-
with an average of 52 ± 9 species·ha−1, and that is ging of varying intensity, mirroring the status of a
greater than the value of 24 species found in a semi- large fraction of forests in the Congo basin
deciduous forest of east Cameroon (Chimi et al. 2018). (Megevand et al. 2013). Long-term disturbance, vary-
Besides, the 167 small stems species with the average of ing between 0.00 and 8.31%, which has occurred in
24 ± 5 species·ha−1 found in the DCF was also greater this area, may also explain the abundance of pioneer
than the value of thirty species obtained in 3.4 ha of a species found in the plots inventoried (Zekeng 2020;
semi-deciduous forest (Chimi et al. 2018). Zekeng et al. 2020). The variation of species along
the different strata may be explained by the environ-
ment's quality (Zekeng 2020). Indeed, previous stud-
Floristic composition ies in this area have shown that different indicator
species were observed in the different strata, which
The different tree diameter classes each had a differ- was consequent to the soil composition and the to-
ent assemblage of ecologically dominant species. The pography of each habitat (Zekeng 2020).
most important species found in the three strata
92 Zekeng et al.

Throughout the sample area, the Malvaceae and deforestation, degradation and conversion trajectories
Cannabaceae were among the ten families with the (Zekeng et al. 2019). Therefore, given the need to pre-
highest FIV (Table 2). These two APG IV classification serve these fragile and endangered species, it becomes
families included the Sterculiaceae and Ulmaceae's con- essential to characterize and protect the habitat that
ventional systematic, respectively, the presence of shelters them to avoid a total and irreversible erosion
which is characteristic of semi-deciduous forests of this unique biodiversity. This study also highlights
(Letouzey 1985). The leading dominant families de- the necessity to determine the habitat and real threats
scribed in the Doume Communal forest are similar to facing these species considered high-priority species for
those characteristic of the West African semi-deciduous conservation. This observation is not unique to the DCF
forest belt (White 1983; Vooren and Sayers 1992; as it can also be observed in other communal forests
Bakayoko et al. 2001), which includes members of the whose results have not yet been published. It should be
Malvaceae, Cannabaceae, Fabaceae, Moraceae, noted that the notion of sustainable management and
Ebenaceae, Rubiaceae, Putranjivaceae and biodiversity conservation constitutes a major challenge
Euphorbiaceae. of communal forestry, just like that of community for-
estry. Total destruction of some forest plots in Bimbia
Implications for biodiversity conservation Bonadikombo community forest was noted (Ferenc
and sustainable forest management et al. 2018).
Elsewhere, this study found out that some high-
The present study found that species with high priority priority species for conservation were recorded in the
for conservation in the DCF terra-firme forest account understorey. Considering that this forest is being logged,
for about 16% of all the species. This result shows that it is urgent to ensure that it is managed sustainably.
communal forests require rigorous attention to the ap- Indeed, the loss of biodiversity in production forests is
plication of standards and rules for sustainable manage- most often linked to non-compliance with the operator's
ment as well as for biodiversity conservation. Even if management plan (Ferenc et al. 2018). Sometimes it is
similar comparisons cannot be made with other studies related to non-compliance with low-impact logging
carried out in protected areas (e.g. Kenfack et al. 2007; standards, which is occasionally unintentional because
Gonmadje 2012; Fobane 2017), it emerges from this of lack of knowledge. Therefore, the capacity of the
study that production forest areas should also receive at communal forestry unit's members to master reduced-
least minimum attention from the point of view of both impact logging standards should be strengthened to
biodiversity conservation and sustainable management. ensure its correct implementation in the field by each
Besides, it should be noted that herbaceous plants have logger.
not been evaluated, so an increase in sampling in the Zekeng et al. (2019) point out that agroforestry
Doume Communal Forest could result in some addition- systems, including agricultural plantations, are the
al high=priority species for conservation. Larger study forest's main drivers to non-forest land conversion.
areas usually contain greater numbers of endemic and They also note the urgency to stop this in this
threatened species than smaller areas (Brooks et al. communal forest to avoid the loss or disappear-
2002). Therefore, if adjacent forests were included in ance of the high-priority species for conservation
the present study, the number of such species may identified in the Doume Communal Forest. The
increase. This study provided the conservation status intrusion of local people into the DCF to convert
of the species at a national scale (Onana and Cheek forest land into agricultural land can be justified
2011) as well as the global scale (IUCN 2020). This because most local people say they do not feel the
study reveals that some species were threatened at the impact of forest exploitation on their well-being.
global level, while at the local level they were not Therefore, the mayor's office should ensure that
threatened and vice versa. the council's actions contribute to these popula-
Endemic species are at a great risk of extinction tions' well-being. Moreover, it is urgent to confine
because of their confinement in specific habitats and and connect the farmers to the agricultural land
their low density (Myers et al. 2000; Brooks et al. they already occupy. It will be essential to build
2002). Therefore, they are essential for conservation. the farmers' capacities to utilize sustainable agri-
Previous studies done in this area highlight cultural techniques and the manufacturing of
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 93

biofertilizers, which would also increase agricultur- technical support during soil analyses. We are also grateful to the
Doume municipality for their logistical support during the field-
al yields.
work. Specifically, we thank the Mayoress and the Doume mu-
It has also been reported that species richness in- nicipality Secretary Mrs Mpans Giselle Rose and Ayinda Yannick,
creased aboveground carbon in the Doume Communal respectively, for their administrative diligence and for providing us
Forest (Zekeng 2020). This is important because it has with field permits. We furthermore express our thanks to all those
involved in the fieldwork and data collection as well as community
been shown that these species could constitute a poten- members of the different village of Doume.
tial carbon sink. For this reason, management tech-
niques that enhance carbon storage would be fundamen- Authors’ contributions J.C.Z., R.S., W.N.M and M.M.M.A
tal for improving the forest’s capacity to store carbon designed the research; J.C.Z. and P.A.E. collected the data;
(Zekeng et al. 2020). These measures could contribute J.C.Z. analysed the data; J.C.Z., J.L.F. and M.N.H. wrote the first
draft; and all the author's comments and discussed the results.
to the effectiveness of sustainable forest management.
To reduce the riparian population's pressures on the
DFC, we suggest that its managers introduce incentives,
such as REDD+ or subsidies, to promote local food References
production.
African Plant Database 2020. Conservatoire et Jardin botaniques
de la Ville de Genève and South African National
Conclusion Biodiversity Institute, Pretoria. Available at http://www.
ville-ge.ch/musinfo/bd/cjb/africa/recherche.php
Anonymous (2015) Management plan of the Doume Communal
The importance of the Doume Communal Forest for the forest. Doume council, Doume, Cameroon
conservation of tree diversity within Cameroon has long Arasa-Gisbert R, Vayreda J, Román-Cuesta RM, Villela SA,
been overlooked. The present study has determined the Mayorga R, Retana J (2018) Forest diversity plays a key role
floristic diversity of understorey and overstorey strata of in determining the stand carbon stocks of Mexican forests.
Forest Ecol Managem 415–416:160–171
this communal forest belonging to Cameroon's semi-
ATIBT (2007) Etude sur le plan pratique d’aménagement des
deciduous forest. The research has shown that the forêts naturelles de production tropicales africaines.
Doume Communal Forest is rich and diverse. It docu- Application au cas de l’Afrique Centrale volet 1 « production
ments that most species inventoried in this forest have forestière ». Association Technique Internationale des Bois
Guineo-Congolian-wide distributions and that it is also Tropicaux, Paris, France
Bakayoko A, Kouamé FN, Tra Bi FH, Traoré D (2001) Quelques
home to endemic, rare and threatened species. This aspects floristiques et structuraux dela forêt de Bossematié
study begins to fill in a significant gap in the floristic dans l’est de la Côte d’Ivoire. Ann Bot Afrique Ouest 1:7–19
knowledge about Cameroon's communal forests. Its Bele MY, Sonwa DJ, Tiani A-M (2015) Adapting the Congo
results give a first idea of the Communal Forest's diver- Basin forests management to climate change: link-ages
among biodiversity, forest loss, and human well-being.
sity and endemism. Further studies characterizing plant
Forest Policy Econ 50:1–10
diversity and biogeography of endemic and rare species Bodegom V, Jan A, Savenije H, Wit M (2009) Forests and climate
in the entire Doume Communal Forest and other com- change: adaptation and mitigation. Tropenbos International,
munal forests are required to confirm these first conclu- Wageningen, The Netherlands
sions and gain the data needed to support informed Brooks TM, Mittermeier RA, Mittermeier CG, Da Fonseca GAB,
Rylands AB, Konstant WR, Flick P, Pilgrim J, Oldfield S,
decisions on conservation and more sustainable man- Magin G, Hilton-Taylor C (2002) Habitat loss and extinction
agement of the communal forests of Cameroon. in the hotspots of biodiversity. Conservation Biol 16:909–
923
Acknowledgements The lead author is grateful for the Ph.D. Butchart SHM, Walpole M, Collen B, van Strien A, Scharlemann
exchange scholarship granted by the Transdisciplinary Training JPW, Almond REA, Baillie JEM, Bomhard B, Brown C,
for Resource Efficiency and Climate Change Adaptation in Africa Bruno J, Carpenter KE, Carr GM, Chanson J, Chenery AM,
II (TRECCAFRICA II) project which is funded by the European Csirke J, Davidson NC, Dentener F, Foster M, Galli A,
Union. The research leading to these results has received financial Galloway JN, Genovesi P, Gregory RD, Hockings M,
funding from the British Ecological Society (EA17/1005), the Kapos V, Lamarque J-F, Leverington F, Loh J, McGeoch
Rufford Foundation (grant agreement N° 24895-1), and field MA, McRae L, Minasyan A, Morcillo MH, Oldfield TEE,
material funding from the IDEA WILD Foundation. The authors Pauly D, Quader S, Revenga C, Sauer JR, Skolnik B, Spear
thank all the technicians, especially Mr Oreeditse Kgosidintsi and D, Stanwell-Smith D, Stuart SN, Symes A, Tierney M,
Mr Macpherson T Kavouras of the soil laboratory in the Depart- Tyrrell TD, Vié J-C, Watson R (2010) Global biodiversity:
ment of Environmental Science at the University of Botswana, for indicators of recent declines. Science 328:1164–1168
94 Zekeng et al.

Byng JW, Chase MW, Christenhusz MJM, Fay MF, Judd WS, IUCN (2020) Red List of Threatened Species Available at http://
Soltis DE, Mabberley DJ, Sennikov AN, Soltis PS, Stevens www.iucnredlist.org
PF (2016) An update of the Angiosperm Phylogeny Group Kenfack D, Thomas DW, Chuyong G, Condit R (2007) Rarity and
classification for the orders and families of flowering plants: abundance in a diverse African forest. Biodivers &
APG IV. Bot J Linn Soc 181:1–20 Conservation 16:2045–2074
CBD (2011) Decision X/2, the strategic plan for biodiversity Kent M, Coker P (1992) Vegetation description and analysis: a
2011–2020 and the Aichi biodiversity targets. Nagoya, Japan practical approach. Belhaven press, London, UK
Chimi DC, Zapfack L, Djomo AN (2018) Diversity, structure and Kohm K, Franklin JF (1997) Forestry in the 21st century. Island
biomass (above and below) in a semi-deciduous moist forest Press, Covelo, California
of East Region of Cameroon. J Biodivers Environm Sci 12: Letouzey R (1985) Phytogeographic map of Cameroon at 1:500,
60–72 000, accompanied by: Notice of the phytogeographic map of
Condit R (1998) Tropical forest census plots. 1 edn. Springer Cameroon at 1:500,000. 4 TV: Domain of dense rainforest
Verlag, Berlin, Germany always green. Institute of the International Vegetation Map,
Cottam G, Curtis JT (1956) The use of distance measures in Toulouse, France
phytosociological sampling. Ecology 37:451–460 LPWG (2017) A new subfamily classification of the Leguminosae
Curtis JT, McIntosh RP (1951) An uppland forest continuum in based on a taxonomically comprehensive phylogeny: The
the prairie-forest border region of Wisconsin. Ecology 32: Legume Phylogeny Working Group (LPWG). Taxon 66:
476–496 44–77
De Vos JM, L. N. Joppa, J. L. Gittleman, P. R. Stephens, S. L. Magurran A (2004) Measuring biological diversity. Blackwell
Pimm (2015) Estimating the normal background rate of Publishing, Oxford, UK
species extinction. Conservation Biol 29:452–462 Magurran AE (1988) Diversity indices and species abundance
models. In Ecological diversity and its measurement.
Djuikouo MNK, Doucet J-L, Nguembou CK, Lewis SL, Sonké B
Springer Netherlands, Dordrecht, pp. 7–45
(2010) Diversity and aboveground biomass in three tropical
Mbolo AMM, Zekeng JC, Mala WA, Fobane JL, Djomo Chimi C,
forest types in the Dja Biosphere Reserve, Cameroon.
Ngavounsia T, Nyako CM, Menyene LFE, Tamanjong YV
African J Ecol 48:1053–1063
(2016) The role of cocoa agroforestry systems in conserving
Dudley N, Baldock D, Nasi R, Stolton S (2005) Measuring biodi-
forest tree diversity in the Central region of Cameroon.
versity and sustainable management in forests and agricul-
Agroforest Systems 90:577–590
tural landscapes. Philos Trans, Ser B 360:457–470
McElhinny C, Gibbons P, Brack C, Bauhus J (2005) Forest and
Ferenc M, Sedláček O, Tropek R, Albrecht T, Altman J, Dančák
woodland stand structural complexity: its definition and mea-
M, Doležal J, Janeček Š, Maicher V, Majeský Ľ, Motombi
surement. Forest Ecol Managem 218:1–24
FN, Murkwe M, Sáfián S, Svoboda M, Hořák D (2018)
Megevand C, Mosnier A, Hourticq J, , Sanders K, Doetinchem N,
Something is missing at the bottom: importance of coastal
Streck C (2013) Deforestation trends in the Congo Basin:
rainforests for conservation of trees, birds and butterflies in
reconciling economic growth and forest protection.
the Mount Cameroon area. African J Ecol 56:679–683
Directions in development. Environment and sustainable
Fisher RA, Corbet AS, Williams CB (1943) The relation between development. World Bank. Washington DC
the number of species and the number of individuals in a Mengue-Medou C (2002) Les aires protégées en Afrique: perspec-
random sample of an animal population. J Anim Ecol 12:42– tives pour leur conservation. VertigO 3:1–13
58 Mittermeier RA, Turner WR, Larsen FW, Brooks TM, Gascon C
Fobane JL (2017) Diversité floristique et biomasse des forêts (2011) Global biodiversity conservation: the critical role of
atlantiques du Cameroun. Université de Yaounde I hotspots. In Zachos FE, Habel JC (eds) Biodiversity hotspots:
Gonmadje CF (2012) Diversité et Biogéographie des forêts distribution and protection of conservation priority areas.
d'Afrique Centrale atlantique : le cas du massif de Springer Berlin Heidelberg, Berlin, Heidelberg, pp. 3–22
Ngovayang (Cameroun). Université de Montpellier 2 Muhumuza M, Balkwill K (2013) Factors affecting the success of
Gonmadje CF, Doumenge C, Sunderland TCH, Balinga MPB, conserving biodiversity in national parks: a review of case
Sonké B (2012) Analyse phytogéographique des forêts studies from Africa. Int J Biodivers 2013:7981011–20
d’Afrique Centrale: le cas du massif de Ngovayang Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB,
(Cameroun). Pl Ecol Evol 145:152–164 Kent J (2000) Biodiversity hotspots for conservation priori-
Hassler M (2018) World Plants: synonymic checklists of the ties. Nature 403:853
vascular plants of the world. In Roskov Y, Ower G, Orrell Onana JM (2011) The vascular plants of Cameroon. Ataxonomic
T, Nicolson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, checklist with IUCN assessments. In Onana JM (ed) Flore du
Decock W, Nieukerken E van, Zarucchi J, Penev L (eds) Cameroon 39. IRAD-National Herbarium of Cameroon,
Species 2000 & ITIS Catalogue of Life 2018 Annual London-UK, p 195
Checklist. Available at https://www.catalogueoflife. Onana JM (2013) Synopsis des espèces végétales vasculaires
org/annual-checklist/2018 (Accessed 30 October 2018) endémiques et rares du Cameroun. Check-liste pour la
Hurlbert SH (1971) The nonconcept of species diversity: a critique gestion durable et la conservation de la biodiversité. In
and alternative parameters. Ecology 52:577–586 Onana JM (ed) Flore du Cameroun 40. Ministère de la
IPBES (2019) Summary for policymakers of the global assessment Recherche et de l’Innovation, République du Cameroun,
report on biodiversity and ecosystem services of the Yaoundé, Cameroun, pp 1–277
Intergovernmental Science-Policy Platform on Biodiversity Onana JM, Cheek M (2011) Red data book of the flowering plants
and Ecosystem Services. IPBES secretariat, Bonn, Germany of Cameroon: IUCN global assessments. Royal Botanisc
Plant diversity and conservation concerns in a semi-deciduous rainforest in Cameroon: implications for... 95

Gardens and IRAD-National Herbarium of Cameroon, Kew assessment in an African protected forest: a case of the
Publishing London, UK eastern part of the Dja Wildlife Reserve in Cameroon. J Pl
Pereira HM, Ferrier S, Walters M, Geller GN, Jongman RHG, Sci 4:95–101
Scholes RJ, Bruford MW, Brummitt N, Butchart SHM, Tchouto MGP (2004) Plant diversity in central African rain forest.
Cardoso AC, Coops NC, Dulloo E, Faith DP, Freyhof J, Implications for biodiversity conservation in Cameroon. PhD
Gregory RD, Heip C, Höft R, Hurtt G, Jetz W, Karp DS, thesis, Wageningen University
McGeoch MA, Obura D, Onoda Y, Pettorelli N, Reyers B, Tchouto MGP, Yemefack M, De Boer WF, De Wilde JJFE, Van
Sayre R, Scharlemann JPW, Stuart SN, Turak E, Walpole M, Der Maesen LJG, Cleef AM (2006) Biodiversity hotspots
Wegmann M (2013) Essential biodiversity variables. Science and conservation priorities in the Campo-Ma’an rain forests,
339:277–278 Cameroon. Biodivers & Conservation 15:1219–1252
Poissonnet M, Lescuyer G (2005) Aménagement forestier et par- United Nations (1992) Agenda 21: the United Nations programme
ticipation: quelles leçons tirer des forêts communales du of action from Rio. United Nations, New York
Cameroun ? VertigO 6:1–6 Vooren F, Sayers J (1992) Côte d’Ivoire. In: Sayer JA, Harcourt
Poorter L, Bongers F, Aide TM, Almeyda Zambrano AM, CS, Collins NM (eds) The conservation Atlas of tropical
Balvanera P, Becknell JM, Boukili V, Brancalion PHS, forests: Africa. IUCN and Macmillan Publishers, pp 133–
Broadbent EN, Chazdon RL, Craven D, de Almeida-Cortez 142
JS, Cabral GAL, de Jong BHJ, Denslow JS, Dent DH, White F (1979) The Guineo-Congolian Region and its relation-
DeWalt SJ, Dupuy JM, Durán SM, Espírito-Santo MM, ships to other phytochoria. Bull Jard Bot Natl Belg 49:11–55
Fandino MC, César RG, Hall JS, Hernandez-Stefanoni JL, White F (1983) The vegetation of Africa: a descriptive memoir to
Jakovac CC, Junqueira AB, Kennard D, Letcher SG, Licona accompany the UNESCO/AETFAT/UNSOVegetation map of
J-C, Lohbeck M, Marín-Spiotta E, Martínez-Ramos M, Africa. UNESCO, Paris, France
Massoca P, Meave JA, Mesquita R, Mora F, Muñoz R, Zekeng JC (2020) Plant diversity, conservations concerns and
Muscarella R, Nunes YRF, Ochoa-Gaona S, de Oliveira carbon stocks in Doume Communal forest of Cameroon:
AA, Orihuela-Belmonte E, Peña-Claros M, Pérez-García implication for sustainable forests management. Ph.D thesis
EA, Piotto D, Powers JS, Rodríguez-Velázquez J, Romero- University of Yaounde I
Pérez IE, Ruíz J, Saldarriaga JG, Sanchez-Azofeifa A, Zekeng JC, Sebego R, Mphinyane WN, Mpalo M, Nayak D,
Schwartz NB, Steininger MK, Swenson NG, Toledo M, Fobane JL, Onana JM, Funwi FP, Mbolo MMA (2019)
Uriarte M, van Breugel M, van der Wal H, Veloso MDM, Land use and land cover changes in Doume Communal
Vester HFM, Vicentini A, Vieira ICG, Bentos TV, Forest in eastern Cameroon: implications for conservation
Williamson GB, Rozendaal DMA (2016) Biomass resilience and sustainable management. Modeling Earth Systems
of Neotropical secondary forests. Nature 530:211 Environm 5:1801–1814
Shannon CE, Weaver W (1949) The mathematical theory of Zekeng JC, Van der Sande MT, Fobane JL, Sebego R, Mphinyane
communication. University of Illinois Press, Urbana WN, Mbolo MMA (2020) Partitioning main carbon pools in
Simpson EH (1949) Measurement of diversity. Nature 163: 688 a semi-deciduous rainforest in eastern Cameroon. Forest
Sonké B (2004) Forêts de la Réserve du Dja (Cameroun): etudes Ecol Managem 457:1–13
floristiques et structurales.. Scripta Bot Belg 32:144
Tabue MRB, Zapfack L, Noiha Noumi V, Nyeck B, Meyan-Ya Publisher’s Note Springer Nature remains neutral with regard to
Daghela RG, Ngoma LR, Kabelong Banoho L-P, Chimi jurisdictional claims in published maps and institutional
Djomo C (2016) Plant diversity and carbon storage affiliations.