Research Article

Journal of Orthoptera Research 2020, 29(1): 25-34

Effect of anthropogenic pressure on grasshopper (Orthopiera: Acridomorpha)
species diversity in three forests in southern Cameroon

CHARLY OuMAROU NGoute!, SEVILOR KEKEUNOU!, MICHEL LECOo@2, ARMAND RICHARD NzoKo FIEMAPONG!,
PHILENE CoringE AUDE Um Nyose!, CHARLES FELIX BILONG BILONG?

1 Laboratory of Zoology, Faculty of Science, University of Yaoundé 1, Cameroon.

2 CIRAD, Montpellier, France.

3 Laboratory of Parasitology and Ecology, Faculty of Science, University of Yaoundé 1, Cameroon.

Corresponding author: Charly Oumarou Ngoute (coumaroungoute@yahoo.fr)

Academic editor: Alina Avanesyan | Received 25 January 2019 | Accepted 4 May 2019 | Published 3 February 2020

http://zoobank.org/OAB48AF9-COD3-408A-87A2-F7148C727C6C

Citation: Oumarou Ngoute C, Kekeunou S, Lecoq M, Nzoko Fiemapong AR, Um Nyobe PCA, Bilong Bilong CF (2020) Effect of anthropogenic pressure
on grasshopper (Orthoptera: Acridomorpha) species diversity in three forests in southern Cameroon. Journal of Orthoptera Research 29(1): 25-34.

https://doi.org/10.3897/jor.29.33373

Abstract

Grasshoppers are highly diversified in tropical rainforests and considered
of both ecological and conservation importance. The population dynamics of
central African grasshoppers, however, and the structure of their communities
remain poorly studied. We report here on the impact of human activities on
the diversity of grasshopper species from three localities in southern Camer-
oon: Ongot, more anthropized forest; Zamakoe, moderately anthropized for-
est; and Ngutadjap, less anthropized forest. Data were collected using sweep
nets, quadrats, and pitfall traps. We analyzed how pressures from human
activities affected the grasshopper species compositions using five statistical
methods: (1) two non-parametric estimators for specific richness, (2) abun-
dance, (3) abundance distribution model, (4) a diversity index, and (5) B
diversity index. The results showed no significant differences in species rich-
ness between the sites (nine species at Zamakoe, seven each at Ongot and
Negutadjap). Among these species, one was specific to Ongot and Zamakoe,
while one, two, and three species, respectively, were found only in Ongot,
Negutadjap, and Zamakoe. Abundance and species diversity of grasshoppers
increased with anthropogenic pressure on the forests. We noticed a great sim-
ilarity between the grasshopper communities of the two localities under the
greatest anthropogenic pressure (Ongot and Zamakoe) compared to that of
the less anthropized locality of Ngutadjap. The most common grasshopper
species, Mazea granulosa, was most abundant where deforestation was high-
est. Species diversity was highest in the more and moderately anthropized
forests, and the diversity index showed greater similarity between these two
grasshopper communities compared with that of the less anthropized forest.
This work enables us to better understand how the parameters of these insect
communities reflect the degree of forest degradation in southern Cameroon.

Keywords

biodiversity, degradation rate, grasshopper communities, tropical rainforest

Introduction

Tropical rainforests shelter an important part of the world’s
biodiversity and represent an important stake for all countries, in

particular with regard to the effects of anthropogenic disturbances
and climate change. Forest biodiversity remains poorly studied
throughout the African continent (Basset et al. 2001), where these
ecosystems are heavily deforested, particularly in the Congo Ba-
sin. The rate of deforestation doubled in the Congo Basin between
1990 and 2005 (Tchatchou et al. 2015). These forests are subject
to growing anthropogenic pressures leading to their fragmenta-
tion and progressive destruction (de Wasseige et al. 2012). The
direct causes of deforestation include intensification of mining,
population expansion, intensive agricultural practices, and con-
struction of dams that severely alter the structure of the forest
and its dependent biodiversity. As the primary means of liveli-
hood for semi-subsistence farmers in the Congo Basin, shifting
cultivation uses forest resources for agricultural production and
as a source of non-wood products (Brown 2006). Cameroon loses
about 140,000 hectares of forest per year (Ndoye and Kaimowitz
2000). In its southern part, industrial wood production has in-
creased from 2.3 million m? in 1991 to more than 3 million m? in
2000 (de Wasseige et al. 2012). The destruction of these forests has
altered the biophysical structure of the natural environment and
leads to the breakdown of ecosystem equilibrium and the extinc-
tion of species as well as the modification of the structure of floral
and faunal communities. The faunal composition is known to be
negatively affected by this clearing, with reduction of canopy cover
being the major factor of these losses (Scott et al. 2006, Steer et al.
2009). The habitat loss is predicted to greatly impact invertebrates’
species diversity (Chinery 1993); these organisms are less mobile
than vertebrates, have short life cycles, and are more specialized in
micro-habitats due to their specificity to host plants.
Grasshoppers are a common and diverse invertebrate group
worldwide (Gangwere et al. 1997, Song 2010, Zhang 2011). They
are a dominant group of herbivorous insects with up to 20-30%
of all arthropod biomass (Soliman et al. 2017) and occasionally
constituting as much as half of the biomass in an environment
(Gillon 1983). This group plays an important role in terrestrial

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

26 Cc. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.F BILONG BILONG

food webs and is known to be a good source of protein for other
animals such as amphibians, small reptiles, birds, and small mam-
mals; therefore, their scarcity may disturb the trophic structure in
an ecosystem (Schmidt et al. 1991, Soliman et al. 2017). Grass-
hoppers are important bioindicators of threatened environments
because of their specific microhabitat preferences, functional im-
portance in ecosystems, sensitivity to the modification of biotic
and abiotic factors of their habitats, and the ease with which they
can be sampled (Armstrong and van Hensbergen 1997, Samways
1997, Andersen et al. 2001, Guido and Gianelle 2001, Soliman
et al. 2017). Diversity and community structure of grasshoppers
as they relate to anthropogenic activities, types of vegetation, and
climate change have been studied in many regions of the world
(Otte 1976, Kemp et al. 1990, Clayton 2002, Torrusio et al. 2002,
Gebeyehu and Samways 2003, Steck et al. 2007, Saha and Haldar
2009, Sirin et al. 2010, Branson 2011, Chen et al. 2011, Kekeunou
et al. 2017). However, despite the high rate of deforestation ob-
served, the bioindicator potential of grasshoppers in the Congo
Basin area, and particularly in Cameroon, has been largely ne-
glected. Apart from the recent works of Seino et al. (2013) and
Kekeunou et al. (2017) on the diversity of acridoids in higher
mountains in West Cameroon, abundance and grasshopper diver-
sity have been poorly studied. The present article is a contribution
to the understanding of the effect of anthropogenic pressure and
forest degradation on the abundance and diversity of grasshopper
species in southern Cameroon.

Materials and methods

Study sites. Grasshoppers were collected over a year in three lo-
calities (Ongot, Zamakoe, and Ngutadjap; Fig. 1) in the forest area
located on the margins of southern Cameroon plateau (3°27'N,
11°32'E and 4°10'N, 11°49'E). This area, about 650-700 m in el-
evation, is a part of the plateau that forms the northern and west-
ern edges of the Congo Basin (Westphal et al. 1981). The climate

is typical of the Guinean zone with four seasons comprised of a
long dry season (mid-November to mid-March), a short rainy sea-
son (mid-March to June), a short dry season (July to mid-Septem-
ber), and a long rainy season (mid-September to mid-November).
Precipitation ranges from 1500-2000mm per year (Amou’ou et
al. 1985, Santoir and Bopda 1995). The southern Cameroonian
forest is dominated by Sterculiaceae and Ulmacae, and its under-
growth is made up of herbaceous plants such as Maranthaceae and
Acanthaceae (Westphal et al. 1981). In this ecosystem, the natural
vegetation is regularly degraded by the economic exploitation of
wood and the practice of slash-and-burn agriculture (Santoir and
Bopda 1995). The resulting bushy vegetation after degradation is
less diversified and dominated by Chromolaena odorata, Ageratum
conizoides, Synedrella nodiflora, and Imperata cylindrica. Plantain,
cassava, yam, maize, and groundnut are the main food crops, while
industrial crops include cocoa, coffee, sweet banana, and palm oil.
Grasshoppers were sampled in three forest ecosystems, each with
different levels of anthropogenic pressure and degradation: Ongot
forest, 14 to 88 inhabitants/km? located in the division of Mefou
and Akono, near Yaoundé; Zamakoe forest, 10 to 41 inhabitants/
km? in the division of Nyong and So’o, near Mbalmayo; and Ngut-
adjap forest, 2 to 15 inhabitants/km/? in the division of Ntem Valley,
near Ebolowa (Gockowski 1996). Plant species richness is higher
in the less degraded Ngutadjap forest, lower in the Zamakoe for-
est, and lowest in the Ongot forest (Suppl. material 1). Gockowski
(1996) showed that the residents of Ongot draw more income from
paid work and extensive agriculture. In Ngutadjap, people depend
more on hunting and fishing activities, while Zamakoe is a transi-
tion zone between the conditions of Ongot and Ngutadjap forests.

Grasshopper sampling.—The grasshopper species were sampled
every month from the forests of Ongot, Zamakoe, and Ngutad-
jap using sweep nets, quadrats, and pitfall traps. Samples by net
were made randomly for 30 min; grasshoppers were also captured
by hand on the litter in 22 movable iron quadrats of 1 m? each.

Vegetation

MM Dense forest

Mixed forest
Other

Fig. 1. Study sites in relation to vegetation types in Cameroon (see Mertens et al. 2012).

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.E BILONG BILONG 27

These quadrats were placed every 10m, on two parallel transects of
110m, separated from each other by 10m. Other specimens were
collected in 10 pitfall traps (of 8cm diameter each), 1/3 filled with
5% formalin as a preservative; each trap was laid every 20m in the
same transects after quadrat exploration.

Grasshopper identification.—The collected specimens were identi-
fied using keys from Dirsh (1956, 1961, 1965, 1966, 1970), Jago
(1967), Kevan (1975), Hollis (1975), and Lecog (1980).

Data analysis

Species richness, sampling efforts and species accumulation curves.—
Species richness (S) is the number of species reported from each
sampling site. We have estimated these theoretical values by the
non-parametric estimators viz. Chaol and Abundance-based
Coverage Estimator (ACE) (Marcon 2015) using the software Esti-
mateS (Colwell 2013). The plots of cumulative species number per
sample were generated using the same software with data random-
ized 100 times. We estimated the sampling effort as the ratio of
observed species richness to theoretical species richness. Average
efforts were compared using a Kruskal-Wallis H-test in the soft-
ware PAST (Hammer et al. 2001).

Relative abundances.—The average relative abundances (Marcon
2015) were calculated using the following formula:

Del+nx2+-+nal7
N

fx= 100
Ynx1+nx2+...+nx17 is the sum of abundances of species x from
the first to the seventeenth month in a given site; N is the sum of
abundances of all the species in the three sites. Mean abundance
between the different sites and between species were compared by
the Kruskal-Wallis H-test while comparisons of mean abundances
for two samples were made by the Wilcoxon W-test using PAST.

Abundance distribution models.—The abundance distributions of
the reported species were compared to the geometric distribution

10

model of Motomura, the broken stick model of Mac-Arthur, and
the log series model of Fisher (Carlo et al. 1998, Cielo Filho et
al. 2012, Havyarimana et al. 2013, Marcon 2015) to find the one
that fits most to our dataset. These models provide information
on how species are distributed and on how they share the avail-
able resources in the ecosystem (Havyarimana et al. 2013). PAST
software automatically generates the results from the row data.
The x test was used in PAST to compare the observed abundance
distribution to the expected for the three types of theoretical dis-
tributions tested.

Diversity. —Species diversity of grasshoppers was calculated in PAST
and expressed as dominance (D), Shannon diversity (H), and
evenness (H/Hmax) indexes (Carlo et al. 1998, Tadu et al. 2013,
Marcon 2015, Kekeunou et al. 2017, Mbenoun Masse et al. 2017,
Raghavender and Vastrad 2017). The Shannon index for two sam-
ples were compared using the Student t-test (Hutcheson 1970).

Similarity. —Similarities between the grasshopper communities
were assessed by the Bray Curtis index (C,) (Bray and Curtis 1957,
Tadu et al. 2013, Tadu and Djiéto-Lordon 2014, Raghavender and
Vastrad 2017) and the correspondence analysis of the species to
the different communities (Yelland 2010). Cluster analysis was per-
formed using the Paired Group Method (UPGMA) in PAST. PAST
graphically generates the Euclidean distances between rows (spe-
cies) and columns (sites/forests) for the correspondence analysis.

Results

Species richness.—A total of 12 grasshopper species were identified
belonging to two families: Pyrgomorphidae (two species) and Ac-
rididae (10 species) (Fig. 2A). The subfamily Catantopinae was the
most diverse with six species following by the Oxyinae and Pyr-
gomorphinae (two species each), and Acridinae and Coptacrinae
with only one species each (Fig. 2B).

Ten of the 12 identified species were collected by net, six
species were collected in quadrats, and only two species in pitfalls
(Table 1). Two species were collected only from the least disturbed
forest of Ngutadjap (Gemeneta opilionoides and Parapetasia femorata).

B

Wn
8 ov
Wn
() 7)
am 6 5
= -
Oo oT)
- we}
2 4 E
= s
= 2
0
.@)
<%
om

®@ Acrididae

m Pyrgomorphidae

mOngot MZamakoe ®Ngutadjap

Fig. 2. Species richness from each study site. A. Families; B. Subfamilies.

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

28 C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.F BILONG BILONG

Table 1. Species richness according to the different sampling methods in the forests.

Family Subfamily Species Ongot Zamakoe Ngutadjap
net quadrat pitfall net quadrat pitfall net quadrat pitfall
Acrididae Acridinae H. gerstaeckeri + + + + +
Catantopinae A. degener + +
G. opilionoides +
G. terrea -
M. granulosa + + + + + + + +
P. carnapi +
S. opacula + =f + %
Coptacrinae C. hopei +
Oxyinae D. fasciata + + + +
P. apicalis + +
Pyrgomorphidae Pyrgomorphinae P. femorata
T. ferruginea +
Number of taxa 7 4 1 7 5 2 6 2 1

+ indicates the presence of the species at the site for the collection method used.

Gemeneta terrea was collected from the two more disturbed forests
of Ongot and Zamakoe, while Apoboleus degener was collected
only from the most disturbed forest of Ongot. Pteropera carnapi,
Cyphocerastis hopei, and Taphronota ferruginea were collected only
from the moderately anthropized forest of Zamakoe (Table 1).
The remaining five species were common to all three localities.

Sampling effort and species accumulation curves.—Sampling captured
almost the entire estimated species assemblage (95.3 + 1.42%).
No significant difference (H = 2, P = 0.36) was observed between
the localities: Ngutadjap (97.0 + 3%), Ongot (96.5 + 3.5%), and
Zamakoe (92.5 + 2.5%) (Table 2). The species accumulation curve
of each forest started to flatten out towards the end of the sam-
pling period (Fig. 3).

Relative abundance.—A total of 465 individuals were collected from
the target localities (Appendix 1). We did not observe great differ-
ences in abundance between seasons (Appendix 1). Among these

Table 2. Sampling effort and diversity of grasshopper species from
the study sites. The values in brackets represent the theoretical spe-
cies richness; a and b: the results of Shannon diversity index test
for two samples.

Diversity/Estimator Ongot Zamakoe Ngutadjap
Taxa S i 9 7
Individuals 167 226 72
Dominance D 0.54 0.71 0.47
Shannon H 0.97% 1.18° 0.73?
Evenness H/Hmax 0.38 0.23 0.46

ACE 93% (7.52) 90% (10.00) 94% (7.40)

Chaol 100% (7.00) 95% (9.47) 100% (7.00)

Mean of Estimators 96.543.5% 92.542.5% 97+3%
(7.26+0.26) (9.73+0.26) (7.2+0.2)

The same letter between two sites shows no significant difference between
the values.

12

10
n
ov
9 8
S.
n
6 6
o
2
£ 4
=)
z

2

0

0 200 400 600 800 1000 1200 1400
Number of samples
Global Ongot Zamakoe Ngutadjap

Fig. 3. Species accumulation curves of the study sites.

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.F. BILONG BILONG 29

Table 3. The mean relative abundance (%) of species between the different study sites. Each value is: mean + standard error; H-value:
Kruskal Wallis test; P-value: probability; a, b and c: the results of the comparisons, with the Wilcoxon test, for two samples.

Family Subfamily/ Species Ongot Zamakoe Ngutadjap H- value P-value Total
Acrididae Acridinae
H. gerstaeckeri 1.10.4 2.07+0.9 1.65+0.8 0.14 0.91 4.82+1.7
Catantopinae
A. degener 0.44+0.2 0 0 0.64 0.13 0.44+0.29
G. opilionoides 0 0 0.45+0.4 0.16 0.36 0.45+0.45
G. terrea OA7t0n1 0.45+0.3 0 0.51 0.32 0.62+0.03
M. granulosa 2G ALED SP 41.05+3.2° 11662252" 22.02 <0.0001 CEASE.
P. carnapi 0 0.77+0.4 0 1.45 0.05 0.77+0.4
S. opacula 4.8+1,9% 0.82+0.3> 1.2+0.6 5.28 0.03 6.82+1.8
Coptacrinae
C. hopei 0 0.37+0,3 0) 0.16 0.36 0.37+0.3
Oxyinae
D. fasciata 0.85+0.4 1.14+0.6 0.35+0.3 0.65 0.51 2.34+0.09
P. apicalis 1.7+0.6 0.86+0.4 0.92+0.4 0.76 0.56 3.48+1.07
Pyrgomorphidae Pyrgomorphinae
P. femorata 0 0 0.57+40.4 0.64 0.12 0.57+40.4
T. ferruginea 0 0.17+0.1 0 0.17+0.1
H-value 62.5 50.02 37.62 85.58
P-value <0.0001 <0.0001 <0.0001 <0.0001
Total 35.522.8? 47.7+3.0° 1682253" 23.49 <0.0001 100

The same letter between two sites shows no significant difference between the values.

A

120
100 200 50
c
i) oY oO
¢ 60 < 120 E 30
re rue 3 20
< <
20 AO 10
0
) 0

LZ «4. SEG 87

1a 22. A SRG 7 89
Rank of abundance

Rank of abundance

Mmmm Observed abundance
— — Log series (Fisher)

B C

1 2 534.40 5.726
Rank of abundance

eeeeee Geometric (Motomura)
@---@ Broken stick (Mac-Arthur)

Fig. 4. Abundance distribution model of species in the different forests. A. Ongot; B. Zamakoe; C. Ngutadjap.

465 individuals, 72 (16.8 + 2.3%) were collected from the low an-
thropized forest of Ngutadjap, 167 (35.5 + 2.8%) from the more
anthropized forest of Ongot, and 226 (47.7 + 3.0%) from the
moderately anthropized forest of Zamakoe (Table 3). The mean
abundances were significantly higher (H = 23.49, P < 0.0001) in
the grasshopper community from Zamakoe, and significantly low-
er in that of Ngutadjap. Mazea granulosa reported from all locali-
ties was the most abundant species (79.2%) (Table 3). The abun-
dance of this species significantly differed among the three sites
(H = 22.02, P < 0.0001): 11.7% in Ngutadjap, 26.4% in Ongot,
and 41.1% in Zamakoe (Table 3). The common species Holopercna
gerstaeckeri and Serpusia opacula were less abundant than M. granu-
losa. All other species were present with very low abundances in
the different sites studied (Table 3).

Abundance distribution models.—The grasshopper species collected
during this study were distributed into seven abundance ranks in
the Ongot and Ngutadjap forests and in nine abundance ranks
in Zamakoe forest. The distribution models of species abundance
from the target localities were very different from the geometric
model of Motomora: Ongot (7? = 53.3; P < 0.001; Fig. 4A), Za-
makoe (x? = 562.2; P < 0.001; Fig. 4B), and Ngutadjap (7? = 30.6;
P < 0.001; Fig. 4C); the broken stick of MacArthur model: Ongot
(x? = 88.6; P < 0.001; Fig. 4A), Zamakoe (x? = 290.8; P < 0.001;
Fig. 4B), and Ngutadjap (7 = 27.4; P < 0.001; Fig. 4C). All the
observed abundance distribution models were closer to, though
slightly different from, Fisher’ log-series distribution model: On-
got (xy? = 17.1; P = 0.002; Fig. 4A), Zamakoe (x? = 110.5; P < 0.001;
Fig. 4B), and Ngutadjap (v7? = 11.1; P = 0.011; Fig. 4C). The rare

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

30 ¢. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.E BILONG BILONG

Similarity

os 9 9 2S 9

Ngutadjap

Zamakoe

Ongot

Fig. 5. Similarity between the different forest grasshopper com-
munities using the Bray-Curtis index.

species (average relative abundance <1%) accounted for 58% of
the species collected in all three of the studied forests.

Diversity.—The dominance index showed that there are fewer
dominant species in Zamakoe (0.71) than in Ongot (0.54) and
Ngutadjap (0.47) (Table 2). The Shannon diversity index (H) was
higher at Zamakoe (H = 1.18, H/Hmax = 0.23) followed by Ongot
(H = 0.97, H/Hmax = 0.38) and lower at Ngutadjap (H = 0.73, H/
Hmax = 0.46) (Table 2). The Shannon diversity index of grasshop-
per communities of Zamakoe and Ngutadjap forests were signifi-
cantly different (t = 14.3, P= 0.005) (Table 2). All the above shows
that the species diversity has increased with the level of forest deg-
radation in the study site.

Similarity.—The cluster analysis based on species composition per-
formed on the basis of Bray-Curtis index revealed that the grass-
hopper communities of Zamakoe and Ongot forests are more
similar to each other (Fig. 5). The disposition of species by cor-

Zamakoe

C. hopei, P. carnapi

0.6 T. ferruginea
0.3 4 G. teNXea All
e °M. granulosa
0.0 D. fasciata
N
cB)
we -0.3
£
2 06 S. onaeuia
S
-0.9
42 A. degener

oH. gerstaeckeri

e P. apicalis

respondence analysis shows that most of the species studied were
closer to these two most degraded forests of Ongot and Zamakoe
(Fig. 6). A. degener was specific to Ongot; P. carnapi, C. hopei, and
T. ferruginea were specific to Zamakoe; and G. opilionoides and P.
femorata were specific to Ngutadjap (Fig. 6).

Discussion

Species richness and sampling effort.—The sampling efforts were high,
varying between 87% and 93% in the forests studied, with no signif-
icant difference, which is consistent with the statement of Branson
(2011) that evaluation and comparison of grasshopper diversities
requires that all regions and ecosystems be studied in the same way.
The species accumulation curve of each forest started to flatten out
towards the end of the sampling period; this shows that almost all
the species had been collected: all the common species were sam-
pled. The missing species are likely to be all rare taxa corresponding
to the expected low abundance nature of tropical forest faunas.

Overall, 12 species were identified: seven in Ngutadjap and
Ongot and nine in Zamakoe. Seino et al. (2013) and Kekeunou
et al. (2017) have identified, respectively, 28 and 27 species in
the mountainous area of West Cameroon. This considerable dif-
ference in species richness can be explained by the fact that (1)
previous studies collected grasshoppers in both fallows and for-
ests and (2) the works cited were conducted in the upland area
of western Cameroon with two climatic seasons, while we carried
out the present work in the southern Cameroon plateau with four
climatic seasons. The structure, biology, and ecology of the grass-
hopper communities are logically expected to be different in the
two different habitats.

Grasshoppers are indeed recognized as abundant insects in
open environments, which may explain the low species richness
observed in our work. Joubert et al. (2016) recently reported that
grasshoppers constitute a significant proportion of invertebrate
diversity in grasslands; their abundance increases with burning,
cattle grazing, and short vegetation. Spungis (2007) and Arya et

Ngutadjap

G. opilionoides
P. femorata

-0.8

-0.4 0.0

0.4 0.8

12 16 2.0 2.4

Coordinate 1

Fig. 6. Disposition of species between the different study sites using correspondence analysis.

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.F. BILONG BILONG 3]

al. (2015) also carried out studies in forests and found 14 and
12 species of grasshoppers identified, respectively, in the Western
Himalaya in India and in the Ziemupe Nature Reserve forest in
Latvia; the number of species of grasshoppers collected in these
studies are similar to ours. Nevertheless, Raghavender and Vastrad
(2017) report a high species richness, 30 species, in the forest of
Dharwad in India. This difference may be explained by differences
in the climate, types of vegetation, and grasshopper communities
between southern Cameroon and the Dharwad region in India. In
our study, the families were Acrididae (10 species) and Pyrgomor-
phidae (two species). Dirsh (1965, 1966, 1970) and Mestre and
Chiffaud (2009) showed that these two taxa are the main acridid
families in the fauna of both Cameroon and Congo Basin.

In the same way, Seino et al. (2013) and Kekeunou et al.
(2017) found that Acrididae (18 and 22 species, respectively)
and Pyrgomorphidae (four and six species, respectively) are the
more speciose families in West Cameroon. The same results were
given by Almeida and Camara (2008) in Brazil, and by Arya et al.
(2015), More and Nikam (2016), and Raghavender and Vastrad
(2017) in India. The Catantopinae was the richest subfamily in
the study areas with three species in Ngutadjap and four species
in Zamakoe and Ongot. Seino et al. (2013) reported this sub-
family as most speciose in West Cameroon. After the Oedipo-
dinae, the Catantopinae was also the richest subfamily in both
agriculture and forest ecosystems of Dharwad, India (Raghaven-
der and Vastrad 2017). The above results are consistent with the
findings by Dirsh (1965) more than fifty years ago in Cameroon
and in Africa.

Relative abundance and abundance distributions.—The abundance
of grasshoppers in the three study sites increased with human
pressure. In fact, it is already known that grasshopper abundance
increases in dry grassland habitats and forests used by humans
(Latchininsky and Gapparov 1996, 2011, Spungis 2007, Latchi-
ninsky 2008). These results contrast with those of Soliman et al.
(2017) who reported higher species richness, abundance, and di-
versity in the less disturbed sites in South Cairo, Egypt. We can
therefore assume that the behavior of grasshoppers in response to
the environmental disturbances is influenced by the eco-climatic
zone and the structure of plant and even animal communities.

In fact, ecosystem changes strongly affect behavior, especially
of poikilotherms such as grasshoppers that feed on plant materi-
als (Bronwyn 2013). The increase in abundance as the forest is
opened up by human agency that was observed in our work is not
due to an invasion by grassland or forest edge species, but of for-
est forms due to increased light penetration and, thus, a change in
understory vegetation. The positive correlations that exist between
the population density of grasshoppers and plant species diversity
can be explained by both feeding and sheltering requirements of
grasshoppers (Spungis 2007).

Disturbed and new habitats can be important for the spreading
of some grasshopper forms (Samways et al. 1997, Sergeev 1998,
Latchininsky et al. 2011). At the same time, some grasshopper spe-
cies are threatened by anthropogenic pressures, such as overgrazing
and ploughing (Latchininsky and Gapparov 1996, 2011, Sergeev
1998). The abundance distribution of the species observed in this
work were most similar to, though slightly different from, Fisher’
log-series distribution model. Therefore, species with low abun-
dance were the most numerous in the forests studied compared
to the most abundant ones or those with intermediate abundance
(Havyarimana et al. 2013). This distribution model shows that
although they had different levels of utilization and degradation,

these three forest ecosystems are disturbed by human activities
(Hughes 1986). Under these conditions, the available resources
may be immobilized by a small number of species that develop
strategies of resistance to human disturbances (Ramade 2009, Cie-
lo Filho et al. 2012, Havyarimana et al. 2013); this was the case of
M. granulosa, S. opacula, and H. gerstaeckeri in the forests studied.
The other species are relegated to the unfavorable areas (Ramade
2009), as was the case with G. opilionoides and P. femorata, two very
rare species found only in the less degraded forest of Ngutadjap. It
therefore seems necessary to reconstitute and conserve these dif-
ferent ecosystems in order to protect this forest biodiversity and its
trophic structure.

Diversity and similarity.—In this work, species diversity increased
with the level of human activity and use of forest resources: it was
higher in the more anthropized forests of Zamakoe and Ongot and
lower in the less anthropized forest of Ngutadjap. This result is pre-
sented by our cluster analysis based on species composition. Steer
et al. (2009) also observed an increase in the invertebrate diversity,
especially of diurnal Lepidoptera, with the level of forest degrada-
tion in Madagascar. Recently, Soliman et al. (2017) also reported
significant differences between grasshopper community structures
in moderately and highly disturbed sites in India, using one-way
analysis of similarity. We therefore assume that invertebrate com-
munities, especially of insects, are strongly influenced by increased
human activities in forest ecosystems around the world; these in-
vertebrates are recognized worldwide as indicative of the levels of
natural habitat degradation (Clayton 2002, Gebeyehu and Sam-
ways 2003, Steck et al. 2007, Sirin et al. 2010, Chen et al. 2011).

Acknowledgements

This work was financially supported by a Rufford Small Grant (ID
application: 19665-1) from the Rufford Foundation. We thank Dr.
C.H.E. Rowell for proofreading the manuscript.

References

Almeida AV, Camara CAG (2008) Distribution of grasshoppers (Orthop-
tera: Acridoidea) in the Tapacura ecological station (Sao Lourenco da
Mata, PE/Brazil). Brazilian Journal of Biology 68: 21-24. https://doi.
org/10.1590/S1519-69842008000100004

Amou’ou JP, Melingui A, Mounkam J, Tchepannou A (1985) Le Cameroun.
Armand Colin, Paris.

Andersen AN, Ludwig JA, Lowe LM, Rentz DCF (2001) Grasshopper bio-
diversity and bioindicators in Australian tropical savannas: Responses
to disturbance in Kakadu National Park. Austral Ecology 26: 213-222.
https://doi.org/10.1046/j.1442-9993.2001.01106.x

Armstrong AJ, van Hensbergen HJ (1997) Evaluation of afforestable mon-
tane grasslands for wildlife conservation in the north-eastern Cape,
South Africa. Biological Conservation 81: 179-190. https://doi.
org/10.1016/$0006-3207(96)00034-1

Arya M K, Joshi PC, Vinod PB (2015) Species composition, abundance, den-
sity and diversity of grasshoppers (Insecta: Orthoptera) in a protected
forest ecosystem in the Western Himalayas, India. International Journal
of Fauna and Biological Studies 2: 42-46. http://www.faunajournal.
com/vol2Issue5/pdf/2-5-6.1.pdf

Basset Y, Aberlenc HP, Barrios H, Curletti G, Berenger JM, Vesco JP, Causse
PA, Haug A, Hennion AS, Lesobre L, Marques E O’Meara R (2001)
Stratification and diel activity of arthropods in a lowland rainforest in
Gabon. Biological Journal of the Linnean Society 72: 585-607. https://
doi.org/10.1111/j.1095-8312.2001 .tb01340.x

Branson DH (2011) Relationships between plant diversity and grasshop-
per diversity and abundance in the Little Missouri National Grassland.
Psyche 2011: 748635. https://doi.org/10.1155/2011/748635

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

32 C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.F BILONG BILONG

Bray JR, Curtis JT (1957) An ordination of the upland forest communities
of Southern Wisconsin. Ecological Monographs 27: 325-349. https://
doi.org/10.2307/1942268

Bronwyn AE (2013) Culturally and Economically Significant Insects in the
Blouberg Region, Limpopo Province, South Africa. Dissertation, Uni-
versity of Limpopo, South Africa. http://hdl.handle.net/10386/1002

Brown DR (2006) Personal preferences and intensification of land use:
Their impact on southern Cameroonian slash-and-burn agroforestry
systems. Agroforestry Systems 68: 53-67. https://doi.org/10.1007/
$10457-006-0003-9

Carlo HR, Help P, Herman MJ, Soetaert K (1998) Indices of diversity and
evenness. Oceanis 24: 61-87.

Chen Y, Li Q, Chen Y, Chen Z (2011) Effects of different land use on grass-
hopper diversity in lac agroecosystems. Terrestrial Arthropod Reviews
4: 255-269. https://doi.org/10.1163/187498311X601704

Chinery M (1993) Insects of Britain and Northern Europe. Harper Collins
Publishers, New York.

Cielo Filho R, Martins FR, Gneri MA (2012) Fitting abundance distribution
models in tropical arboreal communities of SE Brazil. Community
Ecology 3: 169-180. https://doi.org/10.1556/ComEc.3.2002.2.4

Clayton JC (2002) The effects of clear cutting and wildfire on grasshop-
pers and crickets (Orthoptera) in an intermountain forest eco-
system. Journal of Orthoptera Research 11: 163-167. https://doi.
org/10.1665/1082-6467(2002)011[0163:TEOCAW]2.0.CO;2

Colwell RK (2013) EstimateS: Statistical estimation of species richness and
shared species from samples. Version 9. http://viceroy.eeb.uconn.
edu/estimates

de Wasseige C, de Marcken P, Bayol N, Hiol Hiol F Mayaux Ph, Desclée
B, Nasi R, Billand A, Defourny P, Eba’a Atyi R (2012) Les Foréts du
Bassin du Congo - Etat des Foréts 2010. Office des publications de
l'Union européenne, Luxembourg. https://doi.org/10.2788/48830

Dirsh VM (1956) The phallic complex in Acridoidea (Orthoptera) in rela-
tion to taxonomy. Transactions of the Royal Entomological Society
of London 108: 223-356. https://doi.org/10.1111/j.1365-2311.1956.
tb02270.x

Dirsh VM (1961) A preliminary revision of the families and subfamilies
of Acridoidea (Orthoptera, Insecta). Bulletin of the British Museum
(Natural History) Entomology 10: 351-419. https://doi.org/10.5962/
bhl.part.16264

Dirsh VM (1965) The African Genera of Acridoidea. Cambridge University
Press, Cambridge.

Dirsh VM (1966) Acridoidea of Angola. Publicacgdes Culturais da Com-
panhia de Diamantes de Angola 74: 1-727.

Dirsh VM (1970) Acridoidea of the Congo (Orthoptera). Annales du Musée
Royal d'Afrique Centrale, Tervuren, Serie in-8-Science Zoologiques-n °
182: 1-605.

Gangwere SK, Muralirangan MC, Muralirangan M (1997) The Bionomics
of Grasshoppers, Katydids and Their Kin. CAB International, Wall-
ingford.

Gebeyehu S, Samways MJ (2003) Responses of grasshopper assemblages
to long-term grazing management in a semi-arid African savanna.
Agriculture, Ecosystems & Environment 95: 613-622. https://doi.
org/10.1016/S0167-8809(02)00178-0

Gillon Y (1983) The invertebrates of the grass layer. In: Bourliére F (Ed)
Ecosystems of the World (Vol. 13). Tropical Savannas. Elsevier, Am-
sterdam, 289-311. http://www.documentation.ird.fr/hor/fdi:17364

Gockowski JJ (1996) Quelques données de l’enquéte agricole dans les vil-
lages de recherche. Programme EPHTA en collaboration avec l'IRAD/
ASB. IITA-Yaoundé, Cameroon.

Guido M, Gianelle D (2001) Distribution patterns of four Orthoptera
species in relation to microhabitat heterogeneity in an ecotonal
area. Acta Oncologica 22: 175-185. https://doi.org/10.1016/S1146-
609X(01)01109-2

Hammer @, Harper DAT, Ryan PD (2001) PAST: Paleontological statis-
tics software package for education and data analysis. Palaeontolo-
gia Electronica 4: 1-9. http://palaeo-electronica.org/2001_1/past/
issuel_01.htm

Havyarimana F, Bigendako MJ, Masharabun T, Bangirinama F, Lejoly J,
Barima YSS, De Canniére C, Bogaert J (2013) Diversité et distribution
d’abondances des plantes d’un écosysteme protégé dans un paysage
anthropisé: cas de la Réserve Naturelle Forestiére de Bururi, Burundi.
Tropicultura 31: 28-35. http://hdl.handle.net/2268/160257

Hollis D (1975) A review of the subfamily Oxyinae (Orthoptera: Acridoi-
dea). Bulletin of the British Museum (Natural History) Entomology
31: 189-234. https://doi.org/10.5962/bhl.part.29486

Hughes RG (1986) Theories and models on species abundance. The Amer-
ican Naturalist 128: 879-899. https://doi.org/10.1086/284611

Hutcheson K (1970) A test for comparing diversities based on the Shan-
non formula. Journal of Theoretical Biology 29:151-154. https://doi.
org/10.1016/0022-5193(70)90124-4

Jago ND (1967) A key to the grasshopper species (Orthoptera: Acridoidea)
recorded from Ghana. Transactions of the Royal Entomological Society
of London 119: 235-266. https://doi.org/10.1111/j.1365-2311.1967.
tb00511.x

Joubert L, Pryke JS, Samways MJ (2016) Positive effects of burning and
cattle grazing on grasshopper diversity. Insect Conservation and Di-
versity 9: 290-301. https://doi.org/10.1111/icad.12166

Kekeunou S, Membou DT, Tamesse JL, Oumarou Ngoute C (2017)
Acrididea diversity in degraded areas of higher mountains in
West Cameroon. African Entomology 25: 239-243. https://doi.
org/10.4001/003.025.0239

Kemp WP, Harvey SJ, O'Neill KM (1990) Patterns of vegetation and grass-
hopper community composition. Oecologia 83: 299-308. https://
doi.org/10.1007/BF00317552

Kevan DKM (1975) A revision of the genus Taphronota Stal, 1873 (Orthop-
tera; Acridoidea; Pyrgomorphidae). Publicades Culturais da Com-
panhia de Diamantes de Angola (1974) 88: 79-150.

Latchininsky AV (2008) Grasshopper outbreak challenges conservation
status of a small Hawaiian Island. Journal of Insect Conservation 12:
343-357. https://doi.org/10.1007/s10841-008-9143-8

Latchininsky A, Gapparov FA (1996) Les conséquences du desséchement
de la mer d’Aral sur la situation acridienne dans la région. Sécheresse
7: 109-113.

Latchininsky A, Sword GA, Sergeev MG, Cigliano MM, Lecog M (2011) Lo-
custs and grasshoppers: Behavior, ecology and biogeography. Psyche
2011: 578327. https://doi.org/10.1155/2011/578327

Lecog M (1980) Clés de détermination des acridiens des zones sahélienne
et soudanienne en Afrique de l’Ouest. Bulletin de I’Institut Fonda-
mental d'Afrique Noire (1979) 41/A/3: 531-595.

Marcon E (2015) Mesures de la biodiversité. UMR Ecologie des foréts de
Guyane, Kourou. http://hal-agroparistech.archives-ouvertes.fr/cel-
01205813v4

Mbenoun Masse PS, Nzoko Fiemapong AR, VandenSpiegel D, Golovatch
IS (2017) Diversity and distribution of millipedes (Diplopoda) in the
Campo Ma’an National Park, southern Cameroon. African Journal of
Ecology 56: 73-80. https://doi.org/10.1111/aje.12418

Mertens B, Neba SG, Steil M, Tessa B, Mendomo Biang JD, Leach A,
Mbouna D, Mboua P, Feuzing A, Methot P, Minnemeyer S, Douard
P, Ngilambi H (2012) Atlas Forestier Interactif du Cameroun. World
Resources Institute, Washington DC.

Mestre J, Chiffaud J (2009) Acridiens du Cameroun et de République cen-
trafricaine (Orthoptera Caelifera). Supplément au catalogue et atlas
des acridiens d'Afrique de l’Ouest. http://acrida.info/PDF2009/Cata-
logue-Acridiens-2009.pdf

More SV, Nikam KN (2016) Studies on grasshoppers (Orthoptera) in Ti-
lari forest, Chandgad, Kolhapur district of Maharashtra (India). Inter-
national Journal of Recent Scientific Research 7: 9457-9460. http://
www.recentscientific.com/sites/default/files/4578._0.pdf

Ndoye O, Kaimowitz D (2000) Macro-economics, markets and the humid
forests of Cameroon, 1967-1997. Journal of Modern African Studies
38: 225-253. https://doi.org/10.1017/S0022278X00003347

Otte D (1976) Species richness patterns of new world desert grasshoppers
in relation to plant diversity. Journal of Biogeography 3: 197-209.
https://doi.org/10.2307/3038010

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.F. BILONG BILONG 33

Raghavender B, Vastrad AS (2017) Changing scenario of short horned grass-
hopper diversity in agriculture and forest ecosystems in Dharwad. Jour-
nal of Entomology and Zoology Studies 5: 268-272. http://www.en-
tomoljournal.com/archives/2017/vol5issue2/PartD//5-1-138-321.pdf

Ramade F (2009) Eléments d’écologie: Ecologie fondamentale (4™ edn).
Dunod.

Saha HK, Haldar P (2009) Acridids as indicators of disturbance in dry de-
ciduous forest of West Bengal in India. Biodiversity and Conservation
18: 2343-2350. https://doi.org/10.1007/s10531-009-9591-9

Samways MJ (1997) Conservation biology of Orthoptera. In: Gangwere
SK, Muralirangan MC, Muralirangan M (Eds) The Bionomics of
Grasshoppers, Katydids and their Kin. CAB International, Walling-
ford, 481-496.

Santoir C, Bopda A (1995) Atlas régional Sud-Cameroun. Orstom/Minrest,
Paris/Yaoundé. http://www.documentation.ird.fr/hor/fdi:010004189

Schmidt GH, Ibrahim NM, Abdallah MD (1991) Toxicological stud-
ies on the long-term effects of heavy metals (Hg, Cd, Pb) in soil on
the development of Aiolopus thalassinus (Fabr.) (Saltatoria: Acridi-
dae). Science of the Total Environment 107: 109-133. https://doi.
org/10.1016/0048-9697(91)90254-C

Scott DM, Brown D, Mahood S, Denton B, Silburn A, Rakotondraparany
F (2006) The impacts of forest clearance on lizard, small mammal
and bird communities in the arid spiny forest, southern Madagascar.
Biological Conservation 127: 72-87. https://doi.org/10.1016/j.bio-
con.2005.07.014

Seino RA, Dongmo TI, Ghogomu RT, Kekeunou S, Chifon RN, Manjeli
Y (2013) An inventory of short horn grasshoppers in the Men-
oua Division, West Region of Cameroon. Agriculture and Biology
Journal of North America 4: 291-299. https://doi.org/10.5251/ab-
jna.2013.4.3.291.299

Sergeev MG (1998) Conservation of orthopteran biological diversity rela-
tive to landscape change in temperate Eurasia. Journal of Insect Con-
servation 2: 247-252. https://doi.org/10.1023/A:1009620519058

Sirin D, Eren O, Ciplak B (2010) Grasshopper diversity and abundance
in relation to elevation and vegetation from a snapshot in Medi-
terranean Anatolia: Role of latitudinal position in altitudinal dif-
ferences. Journal of Natural History 44: 1343-1363. https://doi.
org/10.1080/00222930903528214

Soliman MM, Haggag AA, El-Shazly MM (2017) Assessment of grasshop-
per diversity along a pollution gradient in the Al-Tebbin region, South
Cairo, Egypt. Journal of Entomology and Zoology Studies 5: 298-306.

Song H (2010) Grasshopper systematics: Past, present and fu-
ture. Journal of Orthoptera Research 19: 57-68. https://doi.
org/10.1665/034.019.0112

Spungis V (2007) Fauna and ecology of grasshoppers (Orthoptera) in the
coastal dune habitats in Ziemupe Nature Reserve, Latvia. Latvijas en-
tomologs 44: 58-68. http://leb.daba.lv/44-sp1.pdf

Steck CE, Biirgi M, Bolliger J, Kienast F Lehmann A, Gonseth Y (2007)
Conservation of grasshopper diversity in a changing environment.
Biological Conservation 138: 360-370. https://doi.org/10.1016/j.bio-
con.2007.05.001

Steer MD, Vater A, McCann-Wood S (2009) The effect of forest degra-
dation on the species richness and diversity of a diurnal Lepidop-
tera community in Northern Madagascar. Frontier - Madagascar
Environmental Research, Report 22. The Society for Environ-
mental Exploration, London. http://frontier.ac.uk/Publications/
Files 2010s 1393913551512 35 pdt

Tadu Z, Djiéto-Lordon C, Babin R, Yede, Messop-Youbi EB, Fomena A
(2013) Influence of insecticide treatment on ant diversity in tropi-
cal agroforestry system: Some aspect of the recolonization process.
International Journal of Biodiversity and Conservation 5: 832-844.
https://agritrop.cirad.fr/572130/1/document_572130.pdf

Tadu Z, Djiéto-Lordon C (2014) Ant diversity in different cocoa agroforest
habitats in the centre region of Cameroon. African Entomology 22:
388-404. https://doi.org/10.4001/003.022.0219

Tchatchou B, Sonwa DJ, Ifo S, Tiani AM (2015) Deforestation and Forest
Degradation in the Congo Basin: State of Knowledge, Current Causes
and Perspectives. Occasional paper 144, Center for International For-
estry Research, Bogor. https://doi.org/10.17528/cifor/005894

Torrusio S, Cigliano MM, De Wysiecki ML (2002) Grasshopper (Orthop-
tera: Acridoidea) and plant community relationships in the Argen-
tine pampas. Journal of Biogeography 29: 221-229. https://doi.
org/10.1046/j.1365-2699.2002.00663.x

Westphal E, Embrechts J, Mbouemboue P, Westphal-Stevels JMC, Mou-
zong-Boyomo (1981) L’agriculture autochtone au Cameroun; les
techniques culturales, les séquences de culture, les plantes alimen-
taires et leur consommation. Miscellaneous papers 20-Landbouwho-
gescool, Wageningen.

Yelland PM (2010) An Introduction to Correspondence Analysis. The
Mathematica Journal 12: 1-23. https://doi.org/10.3888/tmj.12-4
Zhang ZQ (2011) Animal biodiversity: An outline of higher-level classifica-

tion and survey of taxonomic richness. Zootaxa 3148: 1-237.

Supplementary material 1

Author: Charly Oumarou Ngoute, Sévilor Kekeunou, Michel
Lecoq, Armand Richard Nzoko Fiemapong, Philéne Corine Aude
Um Nyobe, Charles Félix Bilong Bilong

Data type: plant species richness

Explanation note: Effect of anthropogenic pressures on floristic
composition from the forests of three localities of southern
Cameroon.

Copyright notice: This dataset is made available under the Open
Database License (http://opendatacommons.org/licenses/
odbl/1.0/). The Open Database License (ODbL) is a license
agreement intended to allow users to freely share, modify,
and use this Dataset while maintaining this same freedom
for others, provided that the original source and author(s)
are credited.

Link: https://doi.org/10.3897/jor.29.33373.suppl1

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)

34 C. OUMAROU NGOUTE, S. KEKEUNOU, M. LECOQ, A.R. NZOKO FIEMAPONG, P.C.A. UM NYOBE AND C.E BILONG BILONG
Appendix 1

Species composition and abundance of the grasshopper species in different seasons (Srs: Short rainy season; Sds: Short dry season; Ltrs:
Long rainy season; Lds: Long dry season).

Taxon Srs (n = 2) Sds (n = 2) Lrs (n = 1) Lds (n = 1) Total
Caelifera
Acridoidea
Acridomorpha
Acrididae
Acridinae
Holopercna gerstaeckeri (Bolivar, 1980) 9 13 3 1 26
Catantopinae
Apoboleus degener Karsch, 1891 0 0 2 0 2
Gemeneta opilionoides (Bolivar, 1905) 0 0 2 0 2
Gemeneta terrea Karsch, 1892 1 1 1 0) 3
Mazaea granulosa Stal, 1876 93 71 89 104 357
Pteropera carnapi Ramme, 1929 0 4 1 0) 5
Serpusia opacula Karsch, 1891 8 15 8 7 38
Coptacrinae
Cyphocerastis hopei Brunner, 1920 1 0 0 0 1
Oxyinae
Digentia fasciata Ramme, 1929 5 1 3 2 11
Pterotiltus apicalis Bolivar, 1905 6 4 0 5 15
Pyrgomorphidae
Pyrgomorphinae
Parapetasia femorata Bolivar, 1884 3 1 0) 0) 4
Taphronota ferruginea (Fabricius, 1791) 0 1 0 0 1
Site
Ongot 39 47 34 47 167
Zamakoe 58 49 64 55 226
Ngutadjap 29 15 11 17 72
Total 126 111 109 119 465

n indicates the number of seasons sampled.

JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(1)