Intra-plant distribution and population fluctuation of Raoiella indica Hirst (Acari:

Tenuipalpidae) on coconuts in the state of Falcón, Venezuela

§
C. Vásquez *
1,2 2 3 3 2
, J. Dlouhy , H. Castillo , B. Gómez & J. Lorbes
1
Departamento Entomologia e Acarologia, Escola Superior de Agricultura “Luiz de Queiroz”,
Universidade de São Paulo, Piracicaba, SP 13498-900, Brazil
2
Present address: Universidad Técnica de Ambato, Facultad de Ciencias Agropecuarias,
Provincia de Tungurahua, Ecuador
3
Universidad Experimental Francisco de Miranda, Departamento de Entomologia, Coro,
estado Falcón, Venezuela

Knowledge of the ecology of the red palm mite, Raoiella indica Hirst, is still fragmentary in
South America since it was only recently introduced from East Asia. Gathering intra-plant
distribution and population fluctuation data could contribute to developing more efficient
sampling methods for R. indica. Taking this into consideration, studies were carried out to
determine spatial temporal distribution of R. indica in three commercial plantations in the
state of Falcón, in Venezuela. Higher mite numbers were registered on middle and lower leaf
portions collected from middle or lower plant strata, suggesting that sampling should be
allocated to these canopy areas on coconut trees. The R. indica population showed to be
negatively associated with rainfall. In 2010, population levels of R. indica were higher than in
2011 (maximum 7 mites/cm2); this was probably a function of lower rainfall values for that
year. In 2011, the highest population levels were reached between April and July, coinciding
with the lowest rainfall values over that year. Relating to this population spike, the
Amblyseius largoensis population showed a numerical response in relation to R. indica
density; this predator species does not appear to be exerting bio-control on the mite pest
population.
Key words: red palm mite, ecological studies, mite population, sampling methods.

INTRODUCTION

Raoiella indica Hirst (Acari: Tenuipalpidae) was
originally described from specimens collected on
the coconut leaves from Coimbature, in southern
India (Hirst 1924). Since then it has been consid-
ered a pest of date palms (Zaher et al. 1969), areca
palms (Saradamma 1972) and coconuts (Carrillo
et al. 2012).
Great emphasis has been given to the study of
the ecology of R. indica in Venezuela, Colombia
and northern Brazil (Carrillo et al. 2012) since its
introduction into the Caribbean (Flechtmann &
Etienne 2004; Rodrigues et al. 2007) and some
South American countries (Vásquez et al. 2008;
Carrillo et al. 2011; Návia et al. 2011). Amaro & de
Morais (2013) stated the potential economic and
social impacts of R. indica in Latin American coun-
tries, because the suitable habitat regions overlap
with agricultural areas for R. indica host plants
such as coconuts and bananas. In Mexico, Otero-
*Author for correspondence. E-mail: ca.vasquez@uta.edu.ec
Colina et al. (2016) observed that the coconut
cultivars Pacific Tall and Malayan Dwarf hosted
higher R. indica infestations than other arecaceous
and Musa species, however naturally decreased
with heavy rainfalls starting in June. Moreover,
because R. indica has the potential to inflict both
ecological and economic damage to plants native
to this region, management strategies have been
implemented to minimise the economic impact of
this pest.
The decision on which specific management
practices to apply are usually based on assess-
ments of pest population densities. Assessment of
pest population densities requires effective and
reliable sampling procedures (Fettig et al. 2005). A
basic knowledge of pest dynamics and population
ecology concerning densities in different parts
of the plant canopy (Negloh et al. 2010) are also
important to determine management strategies.

Received 18 January 2018. Accepted 10 August 2018

ISSN 1021-3589 [Print]; 2224-8854 [Online] African Entomology 27(1): 49–57 (2019)
DOI: https://doi.org/10.4001/003.027.0049 ©Entomological Society of Southern Africa
50 African Entomology Vol. 27, No. 1, 2019
These elements help understanding which mech-
anisms determine population size, and thus
enable prediction of future population dynamics
(Benton & Beckerman 2005). Spatio-temporal
distribution of phytophagous mites could be influ-
enced by several factors inherent to the host plant,
including competition with other phytophagous
pests, regulatory effects of natural enemies, and/or
environmental conditions (Sabelis 1985).
Except for the work of Nagesha Chandra &
ChannaBasavana (1984) on areca palm, and Taylor
et al. (2012) and Flores-Galano et al. (2017) on coco-
nut palm, there is little information about the fac-
tors influencing R. indica population fluctuations.
In addition to the work mentioned above, a single
paper was published dealing with the distribution
of R. indica within the coconut palm in Puerto Rico,
and Trinidad and Tobago (Roda et al. 2012). The
present study aims at evaluating the intra-plant
distribution and population dynamics of R. indica
in Venezuela.
MATERIAL AND METHODS
Intra-plant distribution
Intra-plant distribution of R. indica was evalu-
ated in three coconut plots (Chichiriviche, state of
Falcón, country of Venezuela) as follows: Plot 1
(P1), comprising 20-year-old plants on 3.4 ha,
about 1 km from Playa del Norte; Plot (P2), com-
prising of five-year-old plants on 8.0 ha, about
2.5 km from P1; and Plot 3 (P3), with five-year-old
plants on 2.2 ha, 1.4 km from P1 and 1.7 km from
P2. All plants were of the Caribbean Giant variety,
which predominates in Falcón state.
The regional climate of the study area has a
unimodal rain season, a mean annual precipita-
tion of 1100 mm, a mean annual temperature of
26.6 °C, and a mean evaporation of 1912 mm.
According to Holdridge (1979), this region is classi-
fied as Tropical Dry Forest, and contains mainly
xerophytic plants.
Pinnae samples were taken every 12–15 days,
between April 2010 and August 2011; the samples
were taken from 10 randomly selected coconut
palms from the central area of each plot. Each tree
was divided into three foliage strata, namely top,
median and basal canopy. At each sampling date,
two pinnae were selected from each of the top,
median and basal canopies. Pinnae from each leaf
section and palm stratum were then put into
plastic bags. The bags were placed in an icebox for
transport to the laboratory where the pinnae were
examined under a stereomicroscope to estimate
the R. indica density, counting all the post-
embryonic stages (within five regularly distrib-
uted rectangles of 10 cm2) along the lower pinnae
surface.
Mite density data were log-transformed [y =
log(mite density + 1] before the analyses. Data were
submitted to a nested analysis to evaluate varia-
tion between leaf sections and canopy strata using
SAS version 9.1. Mean R. indica densities in each
leaf section or canopy stratum were compared
using Tukey’s test (P < 0.05).
Population fluctuation
Two evaluations were conducted. The first used
data obtained in the study of intra-plant distribu-
tion (using the method previously described). For
the first evaluation, the data within each plot were
pooled, so as to have a single density for each
sampling date; samples collected from different
leaf sections and canopy strata were analysed
together. The second evaluation was conducted
between December 2010 and November 2011, in
the same plots. Samples were collected every
12–15 days, using the same procedure described
earlier, except that only five pinnae were collected
from each plant, and only from median stratum
leaves. In this study, the number of predators on
each leaflet were also evaluated as well as the
density of the scale insect (Aspidiotus destructor
Signoret (Hemiptera: Diaspididae)). Rainfall and
temperature data were obtained from a weather
station (serial number 0396) located in Tocuyo de
la Costa, state of Falcón, about 5 km from the coco-
nut plots where the study was carried out.
RESULTS AND DISCUSSION
Intra-plant distribution
Nested analysis accounted for 57 % of total varia-
tion (R2 = 0.57; F27,2699 = 128.99; P < 0.0001). Except
for triple interactions, variations of all factors and
their two-by-two combinations were statistically
significant. Significance levels determined by the
interactions indicate that the variation pattern of
R. indica densities in relation to one factor varies
according to the other factors. The P-value for the
interaction of coconut plantation × sampling
date × leaf section was near the threshold level of
significance (Table 1).
When strata and leaf section were considered
 Vásquez et al.: Raoiella indica (Acari) on coconuts in Falcón, Venezuela 51

Table 1. Nested analysis results relative to Raoiella indica densities were evaluated on three different coconut planta-
tions, on different sampling dates, (evaluations were made at 12–14 day intervals, in April and August 2010), and
include canopy strata (basal, middle, and apical third) and leaf portions (basal, middle and apical third). The three
plantations are found in the city of Chichiriviche, state of Falcón, Venezuela.

Source d.f. Type III error Mean square F P>F

Date 9 29367.70 3263.08 124.66 <0.0001

Stratum 2 3586.35 1793.18 68.51 <0.0001
Locality 2 3956.61 1978.31 75.58 <0.0001
Leaf section 2 22663.80 11331.90 432.93 <0.0001
Date × stratum 18 3015.90 167.55 6.40 <0.0001
Locality × date 18 3219.47 178.86 6.83 <0.0001
Date × leaf section 18 273.98 68.50 7.89 <0.0001
Locality × stratum 4 1163.00 290.75 2.62 0.0338
Stratum × leaf section 4 1163.00 290.75 11.11 <0.0001
Locality × leaf section 4 284.62 71.15 2.72 0.0285
Locality × date × stratum 36 1201.48 33.37 1.28 0.1296
Date × stratum × leaf section 36 1318.69 36.63 1.40 0.0611
Locality × date × leaf section 36 925.34 25.70 0.98 0.5002
Locality × stratum × leaf section 8 158.47 19.81 0.76 0.6412

together, average R. indica densities were signifi-
cantly higher in plots 2 and 3 (13.4 mites/cm2
and 14.0 mites/cm2 respectively) than in plot 1
(10.1 mites/cm2); no significant differences were
detected between plots 2 and 3 (Fig. 1). When the
plots and strata were considered together, average
R. indica densities were significantly higher on
basal pinnae of (11.8, 14.7 and 16.1 mites/cm2 in P1,
P2 and P3) and median portion pinnae (10.7, 13.5
and 14.9 mites/cm2 in P1, P2 and P3) than of the
apical section pinnae (7.9, 12.1 and 11.1 mites/cm2
in P1, P2 and P3); and again, no significant differ-
ences were detected between plots 2 and 3 (Fig. 2).
When plant stratum and leaf section were consid-
ered together, average R. indica densities were
significantly higher on leaf portion from basal and
median strata; again, no significant difference was
detected between plots 2 and 3 (Fig. 3).
The interactions observed between varying
distributions of R. indica on diverse canopy strata
and sampling dates seem to be related to progres-
sion of the R. indica population. From mid-
December to early May, the three plots recorded
the lowest population densities of R. indica; in the

Fig. 1. Mean number of Raoiella indica /cm2 in coconut plantations in Chichiriviche, Falcón state, Venezuela; from
April to August 2010.
52 African Entomology Vol. 27, No. 1, 2019

Fig. 2. Mean number of Raoiella indica /cm2 on basal, middle and apical strata of coconut trees in state of Falcón,
Venezuela; from April to August 2010.

apical stratum, densities ranged from 2.7 to
7.7 mites/cm2, and were not significantly different
from densities in the median or basal strata where
R. indica densities ranged from 5.9 to 9.2 mites/cm2.
However, when R. indica reached higher popula-
tion levels in mid-June, increased mite numbers in
the apical stratum were observed, ranging from
12.5 to 18.8 mites/cm2. Recent studies on R. indica
and intra-plant distribution of ‘Giant of Jamaica’
and ‘Malaysian Yellow Dwarf’ coconut cultivars
from Trinidad and Tobago, and Puerto Rico,
showed that higher R. indica densities were con-
centrated on the leaves of the median stratum
(Roda et al. 2012). Our results are similar to those
of Roda et al. (2012), since as R. indica population
levels increased, subsequent mite migration to

Fig. 3. Mean number of Raoiella indica /cm2 on basal, middle and apical leaf thirds taken from basal, middle and apical
coconut strata in the state of Falcón, Venezuela.
 Vásquez et al.: Raoiella indica (Acari) on coconuts in Falcón, Venezuela
the upper stratum was observed. Different re-
sults were reported for Areca catechu L. in India;
in this plant, when densities were low, pests
tended to concentrate on basal and middle strata.
When densities became higher, mites tended to
concentrate in the upper strata (YadavBabu &
Manjunatha 2007).
In general, changes in population levels of
phytophagous mites are associated with changes
in prevailing predation rates or cultivar (Duso &
Vettorazzo 1999), or according to prevailing
microenvironment (Perring et al. 1986) or leaf
metabolite concentration (Perring et al. 1983;
Karban & Myers 1989). Taylor et al. (2012) showed
that densities of R. indica depend on the combined
interaction between site temperature, site humid-
ity and phytoseiid densities on coconut. Leaf
quality influences herbivore abundance and
performance on plants (Barber & Marquis 2011).
In coconuts, it has been shown that nitrogen con-
centrations can vary with leaf age, with nitrogen
being lower in immature and senescent leaves;
lower nitrogen in leaves is also associated with a
decreasing rate of photosynthesis (Jakasekara et al.
1996), which could explain R. indica’s preference
for pinnae from median and basal strata. Addi-
tionally, higher number of eggs and motile forms
was observed on coconut plants irrigated with
100 % of its water requirement, suggesting possi-
ble biochemical changes in leaves due to the
suppression of irrigation could explain changes
in R. indica population dynamics (Villasmil et al.
2014).
Based on our results, it can be concluded that
R. indica densities vary between coconut fields
even when the fields are closely located to one
another. Our results suggest that older plants are
apparently less favourable to R. indica than youn-
ger plants; but the actual cause for this difference
could not be established. It could be due to outside
factors, edaphic characteristics, differences in
cultivation systems, nutrition, or endogenous
characteristics of the plants. In addition, based on
our results and on the valuable contributions of
Roda et al. (2012), it can be concluded that given
the variable R. indica densities of plant canopies, if
the highest levels are to be measured, samplings
should be taken from the median or basal sections
of leaves of the median or basal strata of coconut
canopies.
Interestingly, Roda et al. (2012) noted that higher
variance contribution (»50 %) was produced
53
when pinnae were considered, followed by
variance relative to leaf and leaflet position on
leaves. Accordingly, based on statistical argu-
ments, these authors concluded that increasing
the number of trees (>5 trees/sample unit) would
have a greater impact on the total variance. How-
ever, taking into account the cost to sample each
tree, this recommendation should only be consid-
ered in cases where high accuracy results are
required.
Population fluctuation
In the first evaluation, densities in P2 and P3
were always very similar, whereas densities in P1
were consistently lower than in P2 and P3 plots
(except for on 6 June). This difference could be
related to the age of the plants, which were much
older in P1; however, other environmental factors
should not be disregarded. One factor that might
account for differences between plots is that the
plots belonged to different owners who could
have adopted different cultivation practices not
accounted for in this study.
In this study, R. indica densities ranged between
5 and 25 mites/cm2 (Fig. 4). Marked reductions in
population densities were observed in three of the
four cases where rainfall was at least 47.5 mm
during the preceding period. A reduction in popu-
lation density was not observed on 29 August,
despite the occurrence of about 20.2–20.6 mm of
rainfall during the preceding period. This is proba-
bly because the deleterious effect of the rain on
R. indica was superseded by other changing envi-
ronmental factors (this period had the highest
average temperature for the time in which the
studies were conducted (average of 29.4 °C).
In the second study, differences in densities
between plots were small. The difference between
the second study and the former study could be
related to the much lower R. indica densities in the
second study, reducing the chances for differences
in densities to be expressed.
Corresponding with what was observed in the
first study, a major increase in rainfall during mid-
and late-March was related to major deflections of
the rising R. indica population in the three plots
from the second study (Fig. 5). Reduction of rain-
fall soon after March corresponded to a new
increases in R. indica populations, up to a maxi-
mum of 5.8 (P2) – 9.0 (P3); these densities were
very close to the lowest density observed in the
first study. The much lower R. indica population
54 African Entomology Vol. 27, No. 1, 2019

Fig. 4. Population fluctuation of Raoiella indica in three commercial coconut plantations (April–August 2010) in Falcón
state, Venezuela.

levels observed in the second study were probably
in part determined by more regular and higher
rainfall, as well as the lower temperature during
this period; during most of the second study, the
low rainfall levels matched the highest rainfall
levels in the first study. The progressive increase in
rainfall and the reduction of temperatures toward
the end of the second study corresponded to a
reduction of R. indica densities. Gondim et al.
(2012) observed that the highest density of R. in-
dica in March 2010 (in Roraima) was 1.5 mites/cm2
of coconut leaflet, whereas the second highest
density (0.35 mites/cm2) was observed in February
2011, coinciding with and/or preceded by periods
of high temperatures and low rainfall.
A significant negative correlation was observed
between rainfall and R. indica density (r = –0.3673,
P = 0.001). Conversely, a significant positive corre-
lation was observed between temperature and
R. indica density (r = 0.202, P = 0.005). However,
the correlational coefficients between rainfall and
R. indica density, as well as between temperature
and R. indica density, were very low, respectively;
this was probably because of interactions between
the different abiotic and biotic factors that could
influence R. indica densities. Taylor et al. (2012)
observed higher mite densities under high tem-
peratures and low rainfall environmental condi-
tions. Raoiella indica population densities showed
to be negatively associated with rainfall and
 Vásquez et al.: Raoiella indica (Acari) on coconuts in Falcón, Venezuela 55

Fig. 5. Population fluctuation of Raoiella indica in three commercial coconut farms (Dec 2010–Nov 2011) in state of
Falcón, Venezuela.
56 African Entomology Vol. 27, No. 1, 2019
relative humidity in India (Nagesha Chandra &
ChannaBasavana 1984; Taylor et al. 2012). As
expected, high temperatures (ranging from 29 to
32 °C) positively influenced population increases
in this pest (Nagesha Chandra & Channa-
Basavana 1984; Sarkar & Somchoudhury 1989;
Flores-Galano et al. 2017).
Within the R. indica densities, apparently
allowed by the prevailing abiotic environmental
conditions, it seems that the predator Amblyseius
largoensis (Muma) (Phytoseiidae) could also have
had a significant effect on densities attained by
R. indica (Fig. 5). Population fluctuations of
R. indica and A. largoensis were shown to be
partially synchronised during the study (r = 0.646,
P = 0.000). Predation of R. indica by A. largoensis
was confirmed by the red colouration of its diges-
tive tract. The systematic increase in predator den-
sities alongside increases in pest densities showed
a partial relationship between these populations;
similarly, the predator population declined only
after the pest populations decreased.
Amblyseius largoensis was the most frequent
predatory mite found in association with R. indica
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