Ecol Res
DOI 10.1007/s11284-017-1499-3
O R I GI N A L A R T IC L E
Julien Kouadio N’Dri • Fabrice Ange Seka •
Pacôme Konan Pokou • Rodolphe Arnaud Guy N’Da •
Jan Lagerlöf
Abundance and diversity of soil mite (Acari) communities
after conversion of tropical secondary forest into rubber
plantations in Grand-Lahou, Côte d’Ivoire
Received: 21 March 2017 / Accepted: 25 August 2017
 The Ecological Society of Japan 2017
Abstract The objective of this investigation was to
understand the modification of mite communities and
soil physico-chemical parameters after conversion of
secondary forests into rubber plantations and how these
change with the aging of the plantations. The sampling
was performed in a humid period and samples were ta-
ken from three of each age of secondary forests, 7-year-
old rubber plantations, 12-year-old rubber plantations
and 25-year-old rubber plantations. We hypothesized
that the stress imposed on mite communities during site
preparation and planting would be compensated for by
the reduction of the soil degradation index as the rubber
plantations age. Across the 12 sampling areas, 120 soil
cores were taken at 10 cm soil depth over a 40 m tran-
sect. Soil physico-chemical parameters were character-
ized and soil mites were extracted with a modified
Berlese-Tullgren funnel over the course of 10 days. The
results showed that conversion of secondary forests into
rubber plantations leads to a modification in the density
of mites ( 60 and +1%), species richness ( 48 and
15%), water content ( 62 and 31%), soil organic
carbon ( 67 and 51%) and total nitrogen ( 64 and
52%) respectively after about 7 and 25 years of con-
version. The investigation pointed out an improvement
Electronic supplementary material The online version of this article
(doi:10.1007/s11284-017-1499-3) contains supplementary material,
which is available to authorized users.
J. K. N’Dri (&) Æ F. A. Seka Æ P. K. Pokou Æ R. A. G. N’Da
Unité de Formation et de Recherche (UFR) des Sciences de la
Nature, Université Nangui Abrogoua, 02 BP 801, Abidjan 02, Côte
d’Ivoire
E-mail: ndri_jk@yahoo.fr
Tel.: +225 07 49 38 08
J. K. N’Dri
Centre de Recherche en Ecologie, 08 BP 109, Abidjan 08, Côte
d’Ivoire
J. Lagerlöf
Swedish University of Agricultural Sciences (SLU), Dept. of
Ecology, P. O. Box 7044, 750 07 Uppsala, Sweden
in soil ecological quality with the aging of rubber plan-
tations over time and this was characterized by an in-
crease in the density of mites (+150%), species richness
(+63%), water content (+84%), organic carbon
(+50%) and total nitrogen (+33%) in the 25-year-old
plantations compared to the 7-year-old plantations.
Keywords Density of mites and species richness Æ Soil
physico-chemical parameters Æ Soil degradation index Æ
Rubber plantations Æ Secondary forest
Introduction
The practice of irrational agriculture in the tropics by a
growing population has contributed to significant
reductions in natural forests. Global data from 2000 to
2010 indicate a destruction of 13 million hectares of
forests annually in the world (FAO 2012). The annual
rate of deforestation in Africa is about twice that ob-
served in the rest of the world (FAO 2001). The forests
of Côte d’Ivoire have not escaped deforestation caused
by various agricultural practices. The main perennial
crops observed in several localities are cocoa (Theo-
broma cacao), coffee (Coffea arabica), rubber (Hevea
brasiliensis), and oil palm (Elaeis guineensis) (Ruf 2000).
Introduced in Côte d’Ivoire in the 1950s, rubber culti-
vation has become a major perennial crop in the
southern part of the country (Ruf 2009). Over the last
two decades, rubber plantations have boomed, particu-
larly in the 2000s when the purchase price of rubber
increased dramatically. This boom was marked by an
increasing number of farmers, from 8, 000 in 2000 to
31,192 in 2007 (Brindoumi 2015), and cultivated area,
which increased from 200,000 ha in 2008 (Ruf 2009) to
more than 300,000 ha in 2012 (CNRA 2013). Thus
rubber production is considered the main cause of
deforestation (Allen et al. 2015).
 Several studies have shown that forest conversion conditions as in secondary forest. The restoration of soil
into agrosystems and continual agricultural manage- mite communities would indicate that rubber planta-
ment (clearings, weeding, inputs of pesticides and fer- tions promote a stable soil environment.
tilizers, etc.) can affect the communities of soil
mesofauna and microfauna either directly (Lindo and
Visser 2004; Minor and Cianciolo 2007; Maribie et al. Methods
2011; Zhao et al. 2013; Sarkar et al. 2014; Krashevska
et al. 2015, 2016; N’Dri et al. 2017) or indirectly (Ha- Study site
segawa et al. 2012). In general the impact of conversion
is characterized by a drop in mite abundance and species The investigation was conducted in the department of
richness (Maribie et al. 2011; Sarkar et al. 2014; N’Dri Grand-Lahou (513¢N; 503¢W) situated in southern
et al. 2017) and a loss of soil carbon and nitrogen con- Côte d’Ivoire about 140 km of Abidjan. The climate of
tent (Guillaume et al. 2015, 2016). As a consequence, a the area is an equatorial type with four seasons—a long
decrease in soil fertility is observed (Bedano et al. 2006). dry season from December to March, a long wet season
According to Minor and Cianciolo (2007), land use type from April to July, a short dry season from August to
is a significant factor influencing both small-scale (within September, and a short wet season from October to
individual soil cores) and site-scale diversity of Orib- November (Ettian et al. 2009). Annual rainfall during
atida. Most studies from forest conversion into the study year (2013) was about 1,085 mm, and the
agrosystems and agricultural land that involve mite average monthly temperature was about 27 C. The
community response have been devoted to teak, cocoa, department is crossed by the Bandama River. The veg-
and mixed crops fields (N’Dri et al. 2013, 2017) or to etation is of rainforest type, and various land uses such
coffee, maize, pine, cypress, and conifer plantations as secondary forests, rural domains, and fallow systems
(Maribie et al. 2011; Hasegawa et al. 2012). Nevertheless characterize the study site (Ettian et al. 2009).
some investigations based on soil mites have been per- The forest of Gobodiénou, located upstream of the
formed in rubber plantations in India, Brazil, and Bandama River, contains an important floristic diversity
Nigeria (Chakraborty and Bhattacharya 1993; Franklin and extends over an area of 59,800 ha (Fig. 1). It con-
et al. 2004; Oke et al. 2005).
In Côte d’Ivoire, no study has been devoted to the
impact of rubber plantations on soil mites, and on the
physico-chemical soil parameters that determine their
distribution. However, rubber plantations could serve as
an alternative solution to the loss of natural forest
ecosystems. One of the ecosystem services provided by
this agrosystem is its ability to store carbon (Gréggio
et al. 2008; Podong and Poolsiri 2012) in the different
compartments (the soil and the above- and below-
ground biomass). Soil mites play a key role in the eco-
logical functioning of soil. They are involved in litter
fragmentation, organic matter decomposition, and
minerals recycling (Gulvik 2007; Yang and Chen 2009),
and therefore are seen as good indicators of the eco-
logical quality of the soil (Zhao et al. 2013).
The objective of this investigation was to study the
modification of mite communities and soil physico-
chemical parameters after conversion of secondary for-
est into rubber plantations and how this changes with
the aging of the plantations. Specifically, the research
was based on (i) assessing the abundance and diversity
of mites in the different land use types (ii) measuring the
soil physico-chemical parameters, and (iii) evaluating the
response of mite communities, and the modification of
soil physico-chemical characteristics after secondary
forest conversion into rubber plantations. We hypothe-
sized that the stress imposed on mite communities dur-
ing site preparation and planting would be compensated
for by the reduction of the soil degradation index as the
rubber plantations age. We expected that over time the
density of mites, their diversity, and their community
structure in rubber plantations would reach similar Fig. 1 Map of the study area showing sites and sampling points
tains trees more than 25 m high, herbaceous plants with
several lianas and various palm species. Diospyros spp
and Mapania spp. are among the dominant plants. This
forest is a protected area. Despite this, it is highly
fragmented due to many agricultural activities (planta-
tions of rubber, oil palm and cocoa) carried out there.
Farmers suggested that most of the area we studied was
secondary forest since 1980 (33 years before the field
works) and was converted to rubber plantations in 1988.
A portion of the previous vegetation of rubber planta-
tions less than 10 years old was used for cocoa, coffee or
oil palm plantations in the former rotation. During the
management of plantations, fertilizers and pesticides are
applied once or twice per year during the first 4 years of
planting. Litter is abundant, whereas undergrowth and
herbaceous stratum are absent in rubber plantations.
Soils are ferralitic type (Perraud 1971).
Sampling design
Downstream of the Bandama River and around the
villages of Tieviessou, Betesso and Ahounianssou, nine
rubber plantations (1–2 ha) of different ages (7, 12 and
25 years after planting) were selected for sampling.
Three plantations per age class were selected and con-
sidered as replications. 7 years marks the beginning of
the latex harvest. At 12 years the plantations reach their
maximum production level, whereas at 25 years latex
production begins to decrease. After the productive
period (35–40 years), rubber trees are uprooted and a
new crop is planted after a few years of lying fallow.
Upstream of the Bandama River, three islets of sec-
ondary forest were selected and considered as control
plots. The ‘‘chronosequence’’ approach was used during
the fieldworks, and this allowed for the assessment of the
impact of rubber plantations’ spatio-temporal dynamics
on the variables being studied (Walker et al. 2010).
For each of the 12 sampling stands, soil cores were
taken during the wet period (April–June) with a steel
corer (˘ 5 cm) at 0–10 cm soil depth (Bedano and Ruf
2007) following a 40 m transect. Five sampling points
were allocated along each transect with a 10 m interval
between two consecutive points. At each sampling point,
two adjacent soil cores including litter thickness were
taken. This gave a total of 10 soil cores per transect. In
total, 120 soil cores were taken across the 12 sampling
stands and brought to the laboratory in plastic packets
for mite extraction. Soils for chemical analysis were
sampled according to the Tropical Soil Biology and
Fertility method recommended by Anderson and In-
gram (1993). Composite soil samples were obtained
from five cores taken in quadrats (50 cm · 50 cm · 10
cm) and at 50 cm from the sampling point. On each
sampling stand and following the same transect, three
composite soil samples were taken with 20 m intervals
between two consecutive points. Along the transect,
three soil cores (non-composite samples) were taken
with 20 m intervals between two consecutive points
using the cylinder method (Assié et al. 2008) for physical
measurements. A total of 36 composite soil samples and
36 non-composite soil samples were taken for the phy-
sico-chemical measurements.
Extraction and mite identification
Soil mites were extracted for 10 days with a Berlese-Tull-
gren apparatus (Ducarme et al. 2004; Bedano and Ruf
2007) and stored in a solution of 70% alcohol. Specimens
were mounted in permanent (using Hoyer’s medium) and
semi permanent (using lactic acid) slides and observed with
a digital camera VC.5000 mounted on a NOVEX light
microscope. In the absence of African keys, adult mites
were identified according to major group (Actinedida,
Gamasida, Oribatida and Acaridida), family, genus, and
morphospecies levels by using keys and illustrations pro-
vided in Balogh and Balogh (1992), Krantz (1978), and
Walter et al. (2013). Major groups (Oribatida, Acaridida)
were defined in the traditional sense used in soil zoology
and available in Krantz (1978) and Dindal (1990), not in
the more recent sense found in Krantz and Walter (2009).
SEM pictures were made at the Royal Museum for Cen-
tral Africa, Belgium.
Land use type analysis
Impact of rubber plantations on soil biological quality
The impact of human interference (rubber production)
on soil biological characteristics was estimated by using
an index of change V (Wardle 1995) for each organism
group. This index compares the relative increase or de-
crease in organism abundance between rubber planta-
tions and secondary forest. The index was calculated by
the formula
2MR
V= 1
M R + MF
where MR and MF were the abundance observed
respectively in the rubber plantations and the secondary
forest. The index V ranges from 1 when organisms
occur only in secondary forest to +1 when organisms
occur only in rubber plantations with 0 representing
equal abundance in both secondary forests and rubber
plantations. The magnitude of response to human
interference was expressed by the following categories:
extreme inhibition by rubber plantations V < 0.67;
moderate inhibition by rubber plantations
0.67 < V < 0.33;
mild inhibition by rubber plantations 0.33 < V < 0;
mild stimulation by rubber planta-
tions 0 < V < 0.33;
moderate stimulation by rubber plantations
0.33 < V < 0.67;
extreme stimulation by rubber plantations V > 0.67.
Soil degradation index
The soil degradation index—SDI was expressed as the
inverse of the number of years since the last clearing and
tillage operations (Parisi et al. 2005). This index was
estimated by using the age of the rubber plantations.
The first agrosystems were established in the forest of
Gobodienou in 1980, which was 33 years before the
sampling. Thus the soil degradation index values were:
0.03, 0.04, 0.08, and 0.14 respectively for secondary
forest, and 25-, 12- and 7-year-old rubber plantations.
Data analysis
Soil mite abundance was expressed as the mean
number of individuals per square meter. The mean
species richness, the Margalef diversity index (Gamito
2010), cumulated species richness (Colwell et al. 2004),
the Shannon index (Shannon and Wiener 1962),
evenness (Pielou 1969), the Berger Parker index
(Caruso et al. 2007), and the Sorensen dissimilarity
index expressed in percentages (Chao et al. 2005) were
used to assess the soil mite diversity. The community
structure was characterized by using dominant, ubiq-
uitous, and specialist species (Badejo and Ola-Adams
2000; N’Dri et al. 2016). Soil bulk density was esti-
mated using the cylinder method (Assié et al. 2008).
Soil water content was determined after drying at
105 C for 48 h. Soil pH-H2O was determined by
means of a glass electrode in 1:2.5 soil:water (Tondoh
et al. 2015). The Walkley and Black (1934) method
was used to determine soil organic carbon, namely a
wet oxidation of organic carbon in an acid dichromate
solution, followed by back titration of the remaining
dichromate with ferrous ammonium sulfate. Total
nitrogen was determined using the Kjeldahl method,
and the C/N ratio was also estimated.
Because the data did not follow a normal distribu-
tion, a Kruskal–Wallis test was used to compare the
density of mites in the different land use types. A one-
way ANOVA associated with the post hoc Fisher’s test
was performed to examine the effects of land use types
on the soil physico-chemical factors and mite diversity
parameters. The relationships between soil physico-
chemical and biological parameters along the
chronosequence were investigated with Pearson’s corre-
lation coefficient. The same test was used to determine
the interaction between the soil degradation index and
the overall measured parameters. All tests were carried
out using the software Statistica 7.1 (StatSoft Inc.,
Tulsa, USA). The cumulated species richness was esti-
mated after 500 randomizations by using the software
EstimateS 7.5 (Colwell 2005). The first-order jackknife
non-parametric estimator discussed by Colwell and
Coddington (1994) was used to estimate the true species
richness. The sampling efficiency in the four land use
types was also evaluated.
Results
Soil physical characteristics
The bulk density was significantly lower in the secondary
forest compared to the rubber plantations but did not
differ between the three ages of rubber plantations (one-
way ANOVA; F = 2.102, P = 0.1441). Water content
varied significantly across the land use types (Table 1),
and increased significantly (one-way ANOVA;
F = 3.878, P = 0.0347) with the increasing age of
rubber plantations. The highest value of water content
was measured in secondary forest. The soil water con-
tent decreased by 62 and 31%, respectively, in 7-, and
25-year-old rubber plantations compared to the sec-
ondary forest.
Soil chemical parameters
The soils in the rubber plantations and the secondary
forests were acidic. Mean values of soil organic carbon,
total nitrogen, C/N ratio, and pH were significantly
different across the land use types (Table 2). Apart from
pH, the highest values of chemical parameters were
measured in secondary forest. The amount of soil or-
ganic carbon (one-way ANOVA; F = 3.862,
P = 0.0351), total nitrogen (one-way ANOVA;
F = 4.028, P = 0.0310), C/N ratio (one-way ANOVA;
F = 6.510, P = 0.0055), and pH (one-way ANOVA;
F = 35.938, P = 0.0001) varied significantly with the
increasing age of rubber plantations. The soil organic
carbon decreased by 67 and 51% whereas total nitrogen
decreased by 64 and 52% respectively in 7-, and 25-year-
old rubber plantations compared to the secondary for-
est.
Density of soil mites
The mean density of soil mites changed significantly
(Kruskal–Wallis test, P = 0.0009) across the land use
types (Fig. 2), and increased significantly (Kruskal–
Table 1 Soil physical characteristics measured in the land use types
Bulk density (g cm 3) Water content (%)
SF 0.95 ± 0.06b 29.12 ± 3.95c
R7 1.27 ± 0.03a 11.02 ± 1.13a
R12 1.18 ± 0.02a 18.86 ± 2.14ab
R25 1.23 ± 0.04a 20.23 ± 3.64b
P value 0.0001*** 0.0017**
SF Secondary forest, R7 7-year-old rubber plantations, R12
12-year-old rubber plantations, R25 25-year-old rubber planta-
tions. N = 36, One-way ANOVA test
** P < 0.01, *** P < 0.001; means followed by the same super-
script lowercase letter within column are not significantly different
at the 0.05 level (Tukey’s multiple-comparison test)
Table 2 Chemical variables measured in the land use types
1 1
SOC (g kg soil) TN (g kg soil) C/N pH-H2O

SF 22.9 ± 3.31b 2.00 ± 0.29b 11.46 ± 0.30b 4.48 ± 0.09a

R7 7.5 ± 0.47a 0.72 ± 0.04a 10.35 ± 0.18a 5.87 ± 0.14c
R12 11.3 ± 1.41a 1.05 ± 0.11a 10.55 ± 0.21a 4.67 ± 0.06ab
R25 11.22 ± 1.20a 0.96 ± 0.08a 11.45 ± 0.27b 4.81 ± 0.12b
P value 0.0001*** 0.0001*** 0.0037** 0.0001***

SF Secondary forest, R7 7-year-old rubber plantations, R12 12-year-old rubber plantations, R25 25-year-old rubber plantations, SOC soil
organic carbon, TN total nitrogen, C/N carbon nitrogen ratio, pH-H2O potential of hydrogen–water. N = 36, One-way ANOVA test
** P < 0.01, *** P < 0.001; means followed by the same superscript lowercase letter within column are not significantly different at the
0.05 level (Tukey’s multiple-comparison test)

Major taxa groups

The soil mite communities consisted of four major taxa

Fig. 2 Soil mite density (Mean and SE) observed along the
chronosequence. SF Secondary forest, R7 7-year-old rubber
plantations, R12 12-year-old rubber plantations, R25 25-year-old
rubber plantations. N = 120, Kruskal–Wallis test, P < 0.001.
Means followed by the same lowercase letter are not significantly
different at the 0.05 level (post hoc multiple comparison, bilateral
test)
(Oribatida, Gamasida, Acaridida, and Actinedida).
Whatever the land use type, the Oribatida and Gama-
sida represented the dominant groups with an abun-
dance respectively exceeding 60 and 21%. The
Actinedida (2%) and Acaridida (1%) were minor groups
with highest relative abundance recorded in the 7-year-
old rubber plantations. A total of 14, 7, 14, and 16
species of Gamasida and 45, 22, 28, and 35 species of
Oribatida were observed, respectively, in secondary
forest and in 7-, 12-, and 25-year-old rubber plantations.
Thus the numbers of species of Gamasida and Oribatida
increased with the aging of rubber plantations.
Diversity of soil mites

The species richness of mites was composed of 100

Wallis test, P = 0.0033) with the increasing age of the
rubber plantations. The highest density
(7,429 ± 1,144 ind m 2) was observed in the 25-year-
old rubber plantations and was similar to the density
recorded in secondary forest (7,376 ± 874 ind m 2).
The lowest density was observed in the 7-year-old rub-
ber plantations (2,972 ± 413 ind m 2). The density of
mites decreased by 60% and increased by 1%, respec-
tively, in 7-, and 25-year-old rubber plantations com-
pared to the secondary forest.
morphospecies. A total of 61, 32, 44, and 52 species were
recorded, respectively, in secondary forest and in 7-, 12-,
and 25-year-old rubber plantations. Except for the
Berger Parker index, soil mite diversity parameters
varied significantly across the land use types (Table 3).
The mean species richness (one-way ANOVA;
F = 6.136, P = 0.0032), Shannon index (one-way
ANOVA; F = 4.186, P = 0.0183), Margalef diversity
index (one-way ANOVA; F = 6.175, P = 0.0031), and
the total species richness (one-way ANOVA;
F = 13.983, P = 0.0001) increased significantly with

Table 3 Soil mite diversity across the land use types

Diversity parameter Land use type

SF R7 R12 R25 P value

Mean species richness 5.7 ± 0.56c 2.56 ± 0.36a 4.1 ± 0.57ab 5.26 ± 0.66bc 0.0004***
Margalef diversity index 42.51 ± 5.41b 14.16 ± 2.95a 28.42 ± 5.34ab 39.34 ± 6.33b 0.0006***
Shannon–Wiener Index H¢ 1.48 ± 0.11c 0.74 ± 0.11a 1.05 ± 0.14ab 1.28 ± 0.13bc 0.0006***
Evenness J 0.87 ± 0.04b 0.64 ± 0.08a 0.67 ± 0.07a 0.81 ± 0.06ab 0.048*
Sobs cumulated 61 ± 2.90c 32 ± 1.54a 44 ± 2.05b 52 ± 2.41bc 0.0001***
Berger Parker index 0.34 ± 0.03a 0.48 ± 0.06a 0.46 ± 0.05a 0.42 ± 0.04a 0.455 ns

SF Secondary forest, R7 7-year-old rubber plantations, R12 12-year-old rubber plantations, R25 25-year-old rubber plantations.
N = 120, One-way ANOVA test
* P < 0.05, *** P < 0.001; means followed by the same superscript lowercase letter within rows are not significantly different at the 0.05
level (Tukey’s multiple-comparison test)
Table 4 Impact of rubber plantations on major groups of mites based on the data listed in Table S1

Index V Abundance Species richness

R7 R12 R25 R7 R12 R25

Actinedida +0.000 0.333 1.000 +0.000 +0.000 0.333

Gamasida 0.664 0.311 0.145 0.333 +0.000 +0.067
Oribatida 0.354 0.030 +0.087 0.343 0.233 0.125
Acaridida +1.000 +0.000 +0.000 +1.000 +0.000 +0.000
Total mites 0.451 0.126 +0.004 0.312 0.162 0.080

Index V ranges from 1 to +1 and is increasingly negative or positive as the group under consideration is increasingly decreased or
increased in rubber plantations compared to secondary forest
R7 7-year-old rubber plantations, R12 12-year-old rubber plantations, R25 25-year-old rubber plantations

the increasing age of the rubber plantations. The Berger

Parker index (one-way ANOVA; F = 0.591,
P = 0.5829) and evenness (one-way ANOVA;
F = 1.572, P = 0.2132) did not vary significantly with
the increasing age of rubber plantations. The highest
mean values of the diversity parameters were mainly
recorded in secondary forest, and the lowest values were
primarily recorded in the 7-year-old rubber plantations.
The species richness decreased by 48 and 15%, respec-
tively, in 7-, and 25-year-old rubber plantations com-
pared to the secondary forest. The total number of
observed species corresponded to approximately 69, 63,
69, and 72% of the value predicted by the first-order
jackknife estimator in secondary forest and in 7-, 12-,
and 25-year-old rubber plantations, respectively.

Response of soil mite density and species richness

to the establishment of rubber plantations
The mite community changed along with the age of the
rubber plantations (Table 4). The abundance of Gama-
sida, Oribatida, and total mites were negatively modified
in 7-year-old rubber plantations compared to the sec-
ondary forest. The reverse trend was observed in
25-year-old rubber plantations, where the abundance of
Oribatida and total mites were positively changed
compared to the secondary forest. In all ages of rubber
plantations, the species richness of Oribatida and total
mites were reduced compared to the secondary forest.
The species richness of Actinedida and Gamasida were
positively and negatively impacted, respectively, in
7-year-old rubber plantations. The species richness of
Actinedida and Gamasida were negatively and positively
affected, respectively, in the 25-year-old rubber planta-
tions.
Community structure
Overall, 20 dominant species composed of 14 Oribatida
and 6 Gamasida were collected (Mycobatidae sp.1,
Damaeidae sp.1, Scheloribatidae sp.1, Galumna sp.1,
Galumna sp.2, Javacarus sp.1, Eviphididae sp.1, Rho-
dacaridae sp.1, Mycobatidae sp.2, Acaronychus sp.2,
Fig. 3 Hierarchical classification of land-use types based on the
composition of mites species. UPGMA method (Unweighted Pair-
Group Method using Arithmetic averages) was used. SF Secondary
forest, R7 7-year-old rubber plantations, R12 12-year-old rubber
plantations, R25 25-year-old rubber plantations
Haplozetidae sp.1, Scheloribatidae sp.2, Afrotrachytes
sp.1, Afrotrachytes sp.3, Oppia sp.1, Phthiracarus sp.1,
Epilohmannia sp.1, Trachyuropodidae sp.1,
Trachyuropodidae sp.2, and Nothrus sp.2). The relative
abundance (proportion) of all dominant species put to-
gether varied across the land use types and decreased
with the increasing age of the rubber plantations. A total
of 6, 12, 11, and 7 species were dominants and repre-
sented 10, 38, 25, and 13% of total species observed,
respectively, in secondary forest and in 7-, 12-, and
25-year-old rubber plantations (Table S1). A total of 50
specialist and 10 ubiquitous species were recorded at the
landscape scale. Only two species, Galumna sp.1 and
Galumna sp.2 were considered both dominant and
ubiquitous. Values of the Sorensen dissimilarity index
calculated between the land use types were different. The
highest distance (66%) was obtained between 7-year-old
rubber plantations and secondary forest whereas the
lowest distance (35%) was estimated between 12- and
25-year-old rubber plantations (Fig. 3).
Relationships between mite community characteristics
and soil physico-chemical parameters
The different interactions showed that the density and
species richness of mites were not significantly influenced
by the soil physico-chemical parameters in 7-year-old
rubber plantations. However in 12-year-old rubber
plantations, soil organic carbon, total nitrogen, and pH
were significantly and negatively correlated with the
density of mites (Table 5). The density of mites was
significantly and positively correlated with bulk density
whereas the species richness of mites was significantly
and negatively correlated with water content, respec-
tively, in 25-year-old rubber plantations. Soil organic
carbon, total nitrogen, and pH were significantly and
positively correlated with species richness in secondary
forest.
Interactions between the soil degradation index
and edaphic physico-chemical and biological character-
istics
The soil degradation index decreased with the increasing
age of rubber plantations and impacted the soil physico-
chemical and biological characteristics. The Pearson
correlation showed that the soil degradation index was
negatively correlated with the density of mites
(R = 0.741, P = 0.022), species richness
(R = 0.746, P = 0.021), water content (R = 0.913,
P = 0.001), and soil organic carbon (R = 0.734,
P = 0.024), and positively correlated with the pH
(R = 0.846, P = 0.004) across the chronosequence.
Discussion
Abundance and species richness of soil mites
across the chronosequence
The study of soil fauna diversity along a chronosequence
is described in several research works (Gilot et al. 1995;
Walker et al. 2010; Hasegawa et al. 2012; Tondoh et al.
2015). Soil microarthropod communities are governed
by the age of the agrosystem and the time that has
elapsed since habitat modification (Lindo and Visser
2004; Maribie et al. 2011; N’Dri et al. 2013, 2017; Sarkar
et al. 2014). Our results showed a significant increase in
mite abundance and species richness with increasing age
of rubber plantations, and this trend might be explained

Table 5 Pearson correlation between mite characteristics and soil physico-chemical parameters according to the land use types

Density Richness

R P R P

7-year-old rubber plantations

Bulk density 0.039 0.975 0.095 0.939
Water content 0.769 0.441 0.848 0.356
Soil organic carbon 0.241 0.845 0.369 0.760
Total nitrogen 0.234 0.850 0.362 0.764
pH-H2O 0.235 0.849 0.363 0.763
12-year-old rubber plantations
Bulk density 0.735 0.475 0.497 0.669
Water content 0.195 0.875 0.515 0.656
Soil organic carbon 0.999 0.027* 0.481 0.681
Total nitrogen 0.999 0.032* 0.489 0.674
pH-H2O 0.999 0.034* 0.499 0.668
25-year-old rubber plantations
Bulk density 0.999 0.027* 0.917 0.261
Water content 0.895 0.295 0.998 0.007**
Soil organic carbon 0.983 0.116 0.805 0.404
Total nitrogen 0.990 0.091 0.828 0.379
pH-H2O 0.994 0.073 0.844 0.361
7-25-year-old rubber plantations
Bulk density 0.184 0.636 0.145 0.712
Water content 0.694 0.038* 0.896 0.001**
Soil organic carbon 0.392 0.296 0.118 0.764
Total nitrogen 0.346 0.361 0.104 0.784
pH-H2O 0.680 0.044* 0.348 0.359
Secondary forest
Bulk density 0.967 0.165 0.973 0.148
Water content 0.935 0.231 0.992 0.082
Soil organic carbon 0.862 0.338 0.999 0.025*
Total nitrogen 0.864 0.336 0.999 0.022*
pH-H2O 0.879 0.316 0.998 0.003**

* P < 0.05, ** P < 0.01

by the high availability of trophic resources due to an
increase in litter biomass. The investigation made by
Yéo (2017) on the same plantations pointed out an in-
crease of litterfall production with the increasing age of
rubber plantations (7-year-old rubber plantations:
3.88 ± 0.14 t ha 1 y 1, 12-year-old rubber plantations:
3.86 ± 0.71 t ha 1 y 1, 25-year-old rubber plantations:
5.11 ± 0.60 t ha 1 y 1). The research conducted by
Chaudhuri et al. (2013) in India revealed an increase of
litterfall production (3 years: 2.8 g m 2, 10 years:
200–700 g m 2, 14 years: 750–1400 g m 2, 25 years:
800–1500 g m 2), and a decreasing polyphenol (3 years:
90 lg mg 1, 10 years: 300 lg mg 1, 14 years: 30 lg
mg 1, 25 years: 45 lg mg 1 dry matter) and lignin
(3 years: 1.4 g, 10 years: 2.7 g, 14 years: 0.6 g, 25 years:
1.2 g per 100 g dry matter) content in leaf litters with
increasing age of rubber plantations. The increase of
litter input likely promotes the stability of the soil
microclimate for the development of soil mites. The in-
crease of water content in soil allows for rapid decom-
position of litter (Gréggio et al. 2008; Yang and Chen
2009; Podong and Poolsiri, 2012) and greater restoration
of nutrients in the soil (Podong and Poolsiri, 2012). In
contrast to our observation, Hasegawa et al. (2012)
noticed in the chronosequence of conifer plantations
that forest age was not significantly correlated with
density or species richness of the major mite groups.
The large number of mite species recorded in sec-
ondary forest soils could be explained by the hetero-
geneity of sources of organic residues (food web) with
vegetation consisting of several tree species (Hansen and
Coleman 1998). The complexity of ecological niches and
refuges enhance the stability of soil organism commu-
nities and therefore promote the co-existence of many
mite species (Schneider et al. 2004). Hasegawa et al.
(2012) showed that the vegetation structure in secondary
forest influences both the abundance and species rich-
ness of major groups of mites, and densities of Oribatida
were significantly correlated in broad-leaved sites with
forest age, twig litter weight, maximum DBH of tall
trees, and the species richness of tall trees, whereas
species richness of Mesostigmata was significantly cor-
related with water content in leaf litter, forest floor leaf
litter weight, soil respiration, and species richness of
forest floor plants. The total species richness of mites
observed in our study is superior to that recorded by
Oke et al. (2005) in rubber plantations (23 species). The
mean species richness of Oribatida estimated by
Franklin et al. (2004) in secondary forest and on aban-
doned rubber plantations was similar to ours. Indeed,
those authors recorded 4.8, 5.7, and 6.7 species of Ori-
batida, respectively, in secondary forest and polyculture
systems 1 and 2 based on four tree species (Hevea spp.,
Schizolobium amazonicum, Swietenia macrophylla, and
Carapa guianensis). Chakraborty and Bhattacharya
(1993) observed 37 species of Oribatida in rubber plan-
tations, which was higher than the 22 species of Orib-
atida they identified in 7-year-old rubber plantations.
The abundance, diversity, and community structure
of mites have been shown to be unaffected whether soil
cores are taken from plots under shade or exposed di-
rectly to sunlight (Badejo and Akinwole 2006; Schneider
et al. 2007). However, long-term exposure of 7-year-old
rubber plantations to sunlight increases soil drought and
consequently limits litter decomposition processes and
the biomass and diversity of fungi considered as a food
source for many Oribatida (Koukol et al. 2009). Clear-
ings, weeding, and inputs of pesticides and fertilizers in
rubber plantations could also eliminate some species,
specifically those with a life cycle longer than 1 year
(Behan-Pelletier 1999; Lindo and Visser 2004; Cao et al.
2011).
The investigation made by Maribie et al. (2011) re-
vealed a higher abundance and family richness of soil
mites in artificial forests (cypress forest, abundance: 607
individuals, richness: 16.8 families; pine forest, abun-
dance: 436.2 individuals, richness: 15.8 families) com-
pared to natural forest (abundance: 244 individuals,
richness: 12.3 families) and agrosystems (coffee, abun-
dance: 72.3 individuals, richness: 6.5 families). The
works performed by Sarkar et al. (2014) showed a lower
density of soil mites in brick field (1,612 ind m 2)
compared to agricultural field (4,785 ind m 2) and for-
est area (5,057 ind m 2). The same trend was observed
by N’Dri et al. (2017) in Cote d’Ivoire where the density
and mean species richness of soil mites decreased sig-
nificantly from primary forest to mixed crop fields
(primary forest, density: 2,264 ind m 2, richness: 9.6
species; secondary forest, density: 1,468 ind m 2, rich-
ness: 7 species; fallows, density: 1,264 ind m 2, richness:
5.6 species; multi-species planted trees, density:
1,020 ind m 2, richness: 5.6 species; 4-year-old teak
plantations, density: 979 ind m 2, richness: 5.4 species;
10-year-old teak plantations, density: 959 ind m 2,
richness: 4.8 species; cocoa plantations, density:
673 ind m 2, richness: 4 species; and mixed crops fields,
density: 204 ind m 2, richness: 1.4 species). Krashevska
et al. (2016) found that the species richness, density, and
biomass of living testate amoebae (Protist) were all
lower in rubber plantations compared to rainforest with
an impact that was more pronounced in litter than in
soil at about 13–29 years after conversion. In the litter of
rubber plantations, the concentrations of certain phos-
pholipid-derived fatty acids, including the gram-negative
bacteria marker cy17:0 and the gram-positive bacteria
marker i17:0, were reduced compared to rainforest
(Krashevska et al. 2015). The same variation was ob-
served with basal respiration and gram-negative bacteria
in soil (Krashevska et al. 2015).
Our results suggest that the sample size of 30 cores
per treatment was adequate to capture 63–72% of mite
assemblages in all examined land use types (Fig. 4).
However, the strong unsaturated relationships between
sample size and species richness is explained by two
fundamental factors. First, soil mites in rubber planta-
tions are presented in aggregate form (Chakraborty and
Bhattacharya 1993); second, Oribatid mites have low

Fig. 4 Sample-based rarefaction curves of species richness from the
land use types. The 95% confidence intervals are not shown. SF
Secondary forest, R7 7-year-old rubber plantations, R12 12-year-
old rubber plantations, R25 25-year-old rubber plantations
mobility (Berthet 1964) and this limits the ability to
capture rare species. These trends are similar to those
observed by Minor and Cianciolo (2007).
Rubber plantations and soil ecological quality
Several studies based on cocoa (Dawoe et al. 2014;
Tondoh et al. 2015; N’Dri et al. 2017), teak (N’Dri et al.
2013, 2017) and rubber (Gilot et al. 1995; Oke et al.
2005; Chaudhuri and Nath 2011; Chaudhuri et al. 2013;
Guillaume et al. 2015, 2016; Krashevska et al. 2015,
2016) have highlighted the decrease in soil physico-
chemical and biological parameters after conversion of
forests into agrosystems. This observation is in accord
with our results. In a study by Guillaume et al. (2015),
the carbon content in the Ah horizon of the soil in
rubber plantations was strongly reduced to 62% at
15 years after conversion. Soil organisms and especially
mites are impacted by such changes (Lindo and Visser
2004; N’Dri et al. 2013, 2017). With time, the abundance
of Oribatida and total mites, the species richness of
Gamasida reach similar conditions as in secondary for-
est whereas the species richness of Oribatida and total
species richness were reduced. The species richness of
Oribatida increases in stable ecosystems containing an
appropriate amount of organic matter (Behan-Pelletier
1999; Bedano et al. 2006). The decrease in the soil
degradation index following the aging of rubber plan-
tations would favor an improvement of soil ecological
quality along the chronosequence. The increase in soil
water content consecutive to an increase in litterfall
production following the aging of rubber plantations
(Chaudhuri et al. 2013, Yéo 2017) would enhance the
decomposition process (Gréggio et al. 2008; Yang and
Chen 2009, Yéo 2017). Annual litter mass decomposi-
tion was significantly different (one-way ANOVA,
F = 7.54; P = 0.00287) in the plantations and repre-
sented 3.78 ± 0.11 t ha 1 y 1, 2.73 ± 0.31 t ha 1 y 1,
and 4.21 ± 0.34 t ha 1 y 1, respectively, in 7-, 12-, and
25-year-old rubber plantations (Yéo 2017). Conse-
quently, the average carbon input (2.32 ± 0.18 t ha 1
y 1) and average nitrogen input (0.10 ± 0.008 t ha 1
y 1) in soil were highly concentrated in 25-year-old
rubber plantations (Yéo 2017).
This study points out the improvement in soil eco-
logical quality with the aging of rubber plantations,
which was characterized by an increase in the density of
mites (+150), species richness of mites (+63%), water
content (+84%), organic carbon (+50%), and total
nitrogen (+33%) in the 25-year-old plantations com-
pared to the 7-year-old plantations. It would be impor-
tant to extend the study to the oldest village plantations
(40–50 years old) in order to better appreciate the
implication of this agrosystem in improving soil quality.
For better management of plantations, it would be
interesting to see what effect there is from associated
cover crops and food crops during the first 3 years of
planting. This would enhance the quality of soil organic
matter and would ensure food security for the local
population.
Acknowledgements The authors would like to thank the farmers
and SODEFOR staff for their involvement in identifying suit-
able plantations for this work. We thank the field workers for their
assistance during the sampling period. A Big thank you goes out to
Prof Jérôme E. TONDOH for its assistance on the project and its
advices during data analysis. We thank Dr. Martine Kah Touao
GAUZE, Director of the Ecological Research Center, for accom-
modating the project in his institute. Thanks to Drs. Martinez A.
GUÉI and Ettien F. EDOUKOU for technical assistance. This
study was supported by the International Foundation for Science/
Ref. D/5287-1.
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