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Reproductive Parameters of Diastocera trifasciata
(Fabricius, 1775) (Coleoptera: Cerambycidae:
Lamiinae), Cashew Branches Girdler in Côte d’Ivoire,
Under Semi-Natural Conditions
Ettien Narcice AKESSE*
, San-Whouly Mauricette OUALI-N’GORAN*
*
Faculty of Biosciences/Laboratory of Zoology and Animal Biology, Center of Excellence on Climate Change, Biodiversity and Sustainable
Agriculture, Félix HOUPHOUET-BOIGNY University Côte d'Ivoire
DOI: 10.29322/IJSRP.8.9.2018.p8193
http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193
Abstract- In Côte d'Ivoire, the cashew nut still known as "brown gold" represents a hope for the populations of the producing zones
and contributes to the international influence of the country. However, it is facing attacks from multiple insects including Diastocera
trifasciata formerly Analeptes trifasciata. The lack of data on the reproductive parameters of the species motivated this study. The
objective of this study was to provide knowledge related on the reproductive parameters of Diastocera trifasciata. The study was
conducted under semi-natural conditions at a temperature of 26.83 ± 2.36 ° C and a relative humidity of 75.37 ± 15.09%. Forty-five
pairs of adults were reared in insectarium to study the biology of the species. Data collected were analyzed with Statistica version
7.1 software. The study revealed a pre-mating period of 123.33 ± 8.45 days (mean ± SE) and pre-oviposition period 1.83 ± 0.79
days. Oviposition period was 64.26 ± 8.88 days with lifetime fecundity of 174.3 ± 18.77 eggs and fertility rate 82 ± 9.38%. In
females, body length was 39.71 ± 2.74 mm and longevity was 196.97 ± 6.43 days. There is a positive correlation between longevity,
lifetime fecundity, female size and fertility. In males, body length was 42.23 ± 3.03 mm and longevity 184.56 ± 7.47 days. Sex ratio
was 0.73 in favor of females. Incubation period of eggs was 11.81 ± 1.22 days. Development duration are 198.2 ± 16.22 days for
the larval stage and 18.69 ± 0.63 days for pupal stage. With a total life cycle 211.81 ± 18.87 days, D. trifasciata is univoltine species.
The reproductive parameters of D. trifasciata were obtained. Knowledge of these parameters could be used to support the
development of effective control strategies.
Index Terms- Diastocera trifasciata, branch girdler, reproductive cycle, longevity, lifetime fecundity, fertility.
I. INTRODUCTION
iastocera trifasciata (Fabricius, 1775), formerly Analeptes trifasciata, is a wood-boring insect that attacks cashew trees
(Anacardium occidentale, Anacardiaceae) in several areas of West Africa [1, 2]. Its damage has been observed and described
for the first in 1964 in the forest station Kokondekro to Bamoro and many smallholdings around Bouake in central Côte
d’Ivoire [3]. Since the work of these authors, no other study has made a case of the biology and ecology of this species to determine
effective means of control. In 2014, D. trifasciata has been identified as one of the main four major insect pests of cashew in the
country [2]. This insect girdle and cuts a large number of branches and trunks causing up to 55% cashew nut yield loss [4]. It
represents a serious threat on the farmers' yields and the country's economy [5].
The control technique used until now by producers in Côte d’Ivoire is to capture manually adults during periods of
occurrence and killing them with machetes or grilling. This practice does not make reach all individuals and proves to be very
inefficient with regard to the spread of growing outbreaks. In addition, this practice is not apply in the duration and the periods of
application are ill-chosen.
Since the work investigated by [6] as well as those conducted by [3] which focused on the morphological description of
this pest and this damage, no recent study has made any reference to the biological parameters of this species. The knowledge of
biology and especially the control of reproduction is essential for the development of an effective control strategy.
The objective of this work is to know the reproductive parameters of D. trifasciata. Specifically, it is to determine the egg
incubation period, the duration of the development cycle, female lifetime fecundity, fertility, duration of sexual maturation of adult,
emergence rate, sex ratio and adult longevity.
D
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II. MATERIALS AND METHODS
A. Study site
The study was conducted in the locality of Brobo located 20 km from Bouake on the axis Bouaké-M'Bahiakro, in the center
of Côte d'Ivoire (latitude: 07°38.18N, longitude: 004°49.29W). The vegetation is herbaceous savannah, shrubby or wooded with
forest islets mainly in the lowlands [7]. The study period extended from October 2015 to September 2017 in semi-natural conditions.
B. Material
B1. Biological material
The plant material used was branches and leaves of cashew obtained from the orchards of Brobo. These organs serve both
as food for the individuals and as a female’s egg-laying support. The adult used come from the branches of cashew trees containing
eggs of D. trifasciata kept in rearing in the insectarium.
B2. Breeding device
Thirty wooden cages in the form of a parallelepiped of dimensions 80 cm x 60 cm x 60 cm were used for the rearing of adults.
The four (04) lateral sides are in a metal screen of fine mesh of 2 mm for ventilation of cages. The base and the roof are plywood.
One of the four lateral side serves as the door for cleaning droppings and dead individuals. The plywood of the roof is free to
facilitate the renewal of the branches inside the cage. The temperature inside the insectarium was 26.83 ± 2.36 ° C (mean ± SE)
ranged from 22.6 - 32 ° C. The relative humidity was 75.37 ± 15.09% (mean ± SE) (range 40.94 – 83). These data were recorded
by a digital thermohygrometer MEDISANA. The shots were taken with a NIKON B700 digital camera. The measure of every instar
of the insect were recorded with the help of a vernier slide caliper.
C. Study of reproductive parameters
C1. Adult pre-mating, mating and egg-laying
Forty-five pairs of adults (45 females and 45 males) newly emerged from the branches collected in different fields were
observed in fifteen (15) cages at the rate of three pairs per cage. The body length of each adult was measured from the top of the
head to the apex of the elytra using a vernier slide caliper. The date of emergence is noted. Individuals emerged from the same
branch and from the same orchard were never put in the same cages to avoid consanguinity [8]. In each cage, small fresh branches
of 10 to 15 mm in diameter and dried leaves of cashew trees are provided to insects as food. These branches are renewed every two
days until observation of the first mating. The date of the first mating is noted. Pre-mating period, which is time elapsed between
the adults’ date of emergence and that of the first mating of each pair was recorded.
For mating and egg-laying experiments, as soon as the first mating are observed, the pair is isolated in another cage. In each
cage, a branch of larger size whose circumference is between 15 and 25 cm is made available to them. This branch serves both food
for the pair and female support for laying. Daily, branches are removed from cages and inspected to note the presence or not of
laying. Oviposition pit is marked by the presence of gelatinous substance at the entrance of the holes [8, 9]. Egg counting is based
on the presence of this substance. In case of death of the male before the female, it is replaced by another and observations continue.
The dates and times of mating and laying are observed and noted until the death of the female. The date of female death is noted.
The Preoviposition period (elapsed time between first mating and first oviposition) the oviposition period which is elapsed
time between first and last oviposition were determined.
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The number of eggs deposited per female in the branches was recorded daily. The experiment continued until the death of the
female. Lifetime fecundity (total number of eggs laid by a female during its life) was determined.
After the laying, the oviposition pit are inspected from the outside every day until the observation of wood dust coming out
of these holes. Wood dust is powder from wood rejected by the larva during its feeding activity in the branch. As soon as detection
of this wood dust, the pit is then opened to observe first instar larvae. Number of hatched and unhatched eggs was counted and egg
fertility rate (percentage of the number of eggs hatched to the total number of eggs laid) was determined. The longevity (elapsed
time between date of emergence and the date of death) was determined.
C2. Emergence rate and sex ratio
The emergence rate was studied on 76 fresh branches of average length 149.31 ± 53.34 cm collected in different orchards.
These branches contain eggs newly laid. The number of eggs laid and laying date are recorded on each branch with permanent
marker. They are then brought back to an insectarium to be stored on tables. They are followed until the emergence of adults. The
number of emerged adults was noted. Emergence rate and sex ratio (number of males emerged on number of females emerged)
were calculated.
C3. Incubation period, larval and pupal development of D. trifasciata
During attacks, 100 freshly girdled branches containing eggs were collected from different orchard of Brobo. These selected
branches are those cut the day before the visits. They are recognized by the fresh state of the substance covering the oviposition pit.
They were divided into 50 lots of 02 branches. The eggs are observable by superficial opening of the oviposition pit with a knife.
On each branch, laying date and the number of notch containing the egg are recorded in the indelible marker. According to [10], the
time between egg-laying and hatching of Cerambycidae eggs in general ranges from 3 - 7 days but can be up to 25 days. A batch of
two (02) branches is dissected every three (03) days during the first month following the laying date to observe the evolution of the
eggs. The number of hatched eggs was noted at each observation and the date of hatching was also on each branch. Incubation
period (time elapsed between egg laying and hatching) has been recorded.
The determination of larval and pupal duration was made from the 50 lots of branches formed. After hatching, the branches
are dissected every seven (07) days for seven (07) months to see the evolution of the larvae until observation of pupae and adult
emergence. After dissection of the branches, the shape, the dimensions and the color of the observed stages are described. The
following parameters were recorded: duration of larval development (elapsed time between the date of hatching of the eggs of the
moulting pupal), duration of pupation (time that covers the process that allows the passage of larvae of last stage to the pupa),
duration of pupal development (elapsed time between the date of the pupal moulting and that of obtaining the adult), duration of
development cycle (elapsed time between the date of egg laying and obtaining of the adult stage).
D. Data Analysis
Data processing was done using Statistica software version 7.1 StatSoft [22]. A one-way analysis of variance (ANOVA)
followed by the Newman and Keuls test at the 5% threshold allowed the averages to be compared. Linear regression analyses
(Pearson’s correlation) was used to test the correlation between some reproductive variables in females (lifetime fecundity, fertility,
longevity, oviposition period, and female length) and in males (pre-mating period, longevity and body length).
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III. RESULTS
A. Pre-mating and pre-oviposition periods of adults
The individuals made the first mating between the 104th and 138th days after emergence, pre-mating period of 123.33 ±
8.45 days (mean ± SE). Mating takes place during the day. The first oviposition was observed between the 1st and 4th day after
mating, pre-oviposition period of 1.83 ± 0.79 days (mean ± SE). Pre-mating and pre-oviposition periods are not correlated with any
of the biological parameters studied in the female in this study (Table 1).
Table 1: Linear regression analyses (Pearson’s R correlation matrix) between the different parameters of the female of D. trifasciata
in semi-natural conditions (temperature 26.83 ± 2.36 °C and relative humidity 75.37 ± 15.09 %)
Parameters
Pre-mating
period (days)
Longevity
(days)
Oviposition
period
(days)
Pre-
oviposition
period (days)
Lifetime
fecundity
Body
length
(mm)
Pre-mating period
(days) 1
Longevity (days) 0.356 ns 1
Oviposition period
(days) -0.058 ns 0.315 ns 1
Pre-oviposition
period (days) -0.058 ns 0.114 ns -0.087 ns 1
Lifetime fecundity 0.291 ns 0.958‫٭‬ 0.265 ns 0.181 ns 1
Body length (mm) 0.270 ns 0.813‫٭‬ 0.155 ns 0.259 ns 0.875‫٭‬ 1
Fertility 0.076 ns 0.477‫٭٭‬ 0.608‫٭‬ -0.035 ns 0.518‫٭٭‬ 0.424‫٭٭٭‬
Significance of P-values: ns: non-significant: ‫٭‬P < 0.0001; ‫٭٭‬P < 0.01; ‫٭٭٭‬P < 0.05
B. Oviposition period, lifetime fecundity, fertility rate and longevity of adults
Egg-laying begin at dusk and end at dawn. Oviposition period was 64.26 ± 8.88 days (mean ± SE). Female lifetime fecundity
was 174.3 ± 18.77 eggs, ranged from 126 - 200 eggs. On 174.3 ± 18.77 eggs laid per female, 143.46 ± 22.92 hatched, fertility rate
was 82 ± 9.38% (mean ± SE). Males of D. trifasciata emerge firstly but a longevity was 184.56 ± 7.47 days (range 168 – 201). In
females, longevity was 196.97 ± 6.43 days (range 178 - 208) (Table 2). The linear correlation test shows that there is a positive
correlation between female lifetime fecundity, longevity and egg fertility (Figure 1) (Table 1).There are significant difference
between the longevity in males and females (F = 47.43, ddl = 1, P < 0.05).
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C. Correlation between adult length, fecundity and longevity
In female, body length was 39.71 ± 2.74 mm (means ± SE), ranged 33.10 - 43.50. The linear correlation test between female
length and lifetime fecundity shows that large females lay more eggs (R = 0.87, P < 0.05) (Figure 2). Female length is also positively
correlated with longevity (R = 0.81, P < 0.05) (Figure 3). There were mating between some females who lost their male partners
and newcomers. Females were therefore polyandrous and males were polygynous. In males, body length was 42.23 ± 3.03 mm
(range 31.49 - 46.20). Males are larger in size than females. Analysis of variance at the 5% threshold showed that there is a significant
difference between the body length in males and females (F = 11.38, ddl = 1, P < 0.05). The sexual dimorphism is marked by the
length of the antennae which is 59.72 ± 4.88 mm in the males against 47.98 ± 6.77 mm in the females (Table 2). Male length
correlated neither with longevity (P > 0.05, R2
= 0.002) nor with the pre-copulation period (P > 0.05, R2
= 0.001).
Figure 1: Correlation between female longevity and lifetime
fecundity
175 180 185 190 195 200 205 210
Longevity (days)
120
130
140
150
160
170
180
190
200
210
Lifetimefecundity
R2
= 0,91
y = 2,8x - 360,6
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Figure 2: Correlation between female body length and lifetime fecundity
32 34 36 38 40 42 44
Female body length (mm)
120
130
140
150
160
170
180
190
200
210
Lifetimefecundity
R2 = 0,76
y = 6,02x - 66,63
Figure 3: Correlation between longevity and female body length
170 175 180 185 190 195 200 205 210
Longevity (days)
28
30
32
34
36
38
40
42
44
Femalebodylength(mm)
R2
= 0,66
y = 0,34x - 28,45
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Table 2: Some morphological characteristics of adults of D. trifasciata
Sex Body length
(mm)
Antennae length
(mm) Longevity (days)
Males 42,23 ± 3,03
(31,49 - 46,20
59,72 ± 4,88
(50,8 - 69,95)
184,56 ± 7,47
(168 - 201)
Females 39, 71 ± 2,74
(33,10 - 43,50)
47,98 ± 6,77
(38,18 - 59,41)
196,97 ± 6,43
(178 - 208)
n = 45 in each sex; Mean ± SE (Standard Error of the mean); (minimum – maximum values)
D. Emergence rate and sex ratio of D. trifasciata
Out of a total of 2953 eggs monitored, only 247 adult individuals have emerged, an average emergence or rate of 8.36%. The
emerged adults are divided into 142 females and 105 males, a sex ratio of 0.73 in favor of females.
E. Egg of D. trifasciata and incubation period
The laying were recorded between 18h and 07h in the morning. The eggs are laid in notches made in the bark by the females
only. Each notch of eggs contains only one egg disposed parallel to the longitudinal axis of the branch or stem cut (Figure 4a-b).
The newly laid of D. trifasciata egg is white, oval and similar to a rice grains (Figure 4 b). Eggs length were 5.52 ± 0.23 mm (mean
± SE), ranging from 5.04 - 6.1 mm. The width of the eggs ranged 1.05 - 1.71 mm with a mean of 1.28 ± 0.11 mm. The incubation
period of the eggs was 11.81 ± 1.22 days (range 9 – 15).
(a) (b)
Figure 4: Laying site and fresh eggs of D. trifasciata
(a) Laying notch (red encircled); (b) egg (red encircled) still inlaid in the branch
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F. Larval and pupal development of D. trifasciata
The first instar larvae of D. trifasciata are translucent rosy white color. The body length of larvae (mean ± SE) was 6.39 ±
1.20 mm (range 4.80 - 8.70). The body width was 1.34 ± 0.16 mm (range 1.01 - 1.77). The length of late-instar larvae was 36.45 ±
4.56 mm (range 27.60 - 48.12). The width of late-instar larvae was 7.72 ± 1.2 mm (range 5.8 - 10.60). They are cream-white tinged
with yellow and it is located under the bark. The duration of larval development was 198.2 ± 16.22 days (range 169 - 226).
Before the end of their development, the larvae reduce their gallery in the wood to construct a pupal chambers. At the ends
of this chamber are piled woody debris undigested that look like sawdust. Pupation and pupal development occurs at the end of the
larval development in this pupal chamber. The length of the pupal chamber was 89.26 ± 13.01 mm (range 55.1 – 135). Its width
was 24.95 ± 3.36 mm (range 15.7 - 34.9). The pupation duration was (mean ± SE) 9.02 ± 2.13 days.
The pupa is white or black according on whether it is at beginning or end of its development. The head is inflected toward
the ventral side and carries the antennae which are elongated dorsally along the body before wrapping in a spiral at the end on the
legs. The wing cases are concealed under the folded legs on the ventral side. The length of the pupae was (mean ± SE) 38.64 ± 2.99
mm (range 32.5 - 46.9). The width was 10.66 ± 1.11 mm (range 8.1 - 14.7).
Pupal development occurred between days 16 and 19 after pupation, for an average duration of 18.69 ± 0.63 days. The young
imago who comes out, the wings and the whole body were soft. The head, legs and abdomen have a grey and black color at the
joints. The bands of the wings have a grey coloration. The average duration of pigmentation is 3.68 ± 1.03 days (mean ± SE).
G. Duration of the development cycle of D. trifasciata
The duration of the life cycle was 211.81 ± 18.87 days (range 167 – 240). D. trifasciata is therefore a species univoltine. The
adult, once the pigmentation is complete, emerges through a circular hole made with his mandibles in the bark covering pupal
chamber. The diameter of these exit hole was 17.63 ± 2.43 mm (mean ± SE) (range 11.4 - 28.1) (Figure 5).
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F
Pigmentation
3.68 ± 1.03 days
Preoviposition
1.83 ± 0.79 days
Premating
123.33 ± 8.45 days
Incubation period
11.81 ± 1.22 days
Larval development
198.2 ± 16.22 days
Pupation
9.02 ± 2.13 days
Pupal development
18.69 ± 0.63 days
♀ ♂
(b)
(h)(g)(i)
(j)
a
(f)(e)
(c) (d)
(a)
(k)
Figure 5. Life cycle of Diastocera trifasciata (Coleoptera: Cerambycidae: Lamiinae) under semi-natural
conditions (temperature 26.83 ± 2.36 ° C and relative humidity 75.37 ± 15.09%.
(a): egg (red encircled) observed after opening of a notch; (b): first instar larva (red arrowed); (c) : late instar
larva lateral view; (d): late instar larva ventral view; (e): pupa lateral view; (f): pupa ventral view; (g): young
imago (not emerged from branch) dorsal view; (h): young imago (not emerged from branch) ventral view;
(i): adults emerged; (j): mating on cashew branch; (k): female laying in a cashew branch.
Scale a: 0.79 cm
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IV.DISCUSSION
In this study, the results revealed that the pre-mating period was 123.33 ± 8.45 days. This period is very high compared to that
of Cerambycidae of the Lamiinae subfamily, whose adults feed from a few days to three weeks before becoming sexually mature [8,
12]. Among Cerambycidae Anoplophora glabripennis (Motschulsky), adult females become sexually mature about 9 to 17 days after
emergence. The pre-mating period of D. trifasciata is explained by the fact that Cerambycidae Lamiinae are synovigenic pattern that do
not mate immediately after emergence. Adults emerge with immature sexual organs. They must feed on young shoots and dry leaves
for a period ranging from several days to weeks before reaching sexual maturity [8, 13]. The duration of this period may also depend on
the phenology of the host plant Anacardium occidentale. Indeed, the mating period begins when the plant begins its push vegetative
pre-flowering and extends until flowering. There would be a synchronization between the reproductive period of D. trifasciata and that
of the host plant. According to [14], this phenomenon of synchronization of the cycle of insects and their host plant is frequent and
determines the quantity and the quality of the resources available for reproduction.
The pre-oviposition period obtained in this study (1.83 ± 0.79 days) is close to that obtained in Cerambycidae Cerambyx welensii
which is 2.8 ± 0.2 days, investigated by [15].
The oviposition period of D. trifasciata recorded was 64 ± 8.88 days. These observations are close to those of Cerambycidae
Lamiinae that have an oviposition period of 60 ± 7 days [14]. On the other hand, in the species of the subfamily Cerambycinae, the
oviposition period was 41 ± 17 days [15].
Regarding female’s lifetime fecundity (174.3 ± 18.77 eggs) in breeding condition is superior to the lifetime fecundity in
Cerambycidae Lamiinae which is 155 ± 30 eggs [14]. [16] showed a higher fecundity of 284 ± 27 eggs in the female Cerambycinae
Phoracantha recurva. The positive correlation between fecundity and female size shows that larger females lay more eggs than smaller
females. According to [17], female size is usually a good indicator of potential fecundity.
The phenomena of polyandry and polygyny were observed respectively in females and males of D. trifasciata as in Cerambyx
cerdo [18]. The egg fertility rate recorded in D. trifasciata was 82.37 ± 9.38%. This result is close to those of Cerambycidae such as
Anoplophora glabripennis which is 83 ± 2% [19], Cerambyx cerdo 78 ± 1% [17]. On the other hand, other authors have recorded a
fertility rate ranging from 90 to 98%, in Monochamus galloprovincialis and Phoracantha semipunctata [16, 20]. Rates of between 48-
66% were in Anoplophora glabripennis [21].
Adult males measure 42.23 ± 3.03 mm and live 184.56 ± 7.47 days while females measure 39.71 ± 2.74 mm and live 196.97 ±
6.43 days. This difference in longevity has been reported by various authors. In Cerambycidae, females generally live longer (54 ± 6
days) than males (42 ± 6 days) for a given species [15, 22]. Adults’ longevity of D. trifasciata could be influenced by the long period of
nutrition that precedes reproduction. According to [23], the adult phase of Cerambycidae is entirely devoted to reproduction. Female
longevity was positively correlated with lifetime fecundity, egg fertility. Similar results have been obtained in other Cerambycidae [15,
24]. Longevity can also be influenced by body size, although factors such as temperature, diet, and overall health of an individual may
play a larger role [25].
The emergence rate obtained in this study was 8.36%. These results are quite similar to those of [26] who recorded an emergence
rate of 12% in adults of Cerambycidae Monochamus carolinensis. These findings could be justified by competitions within the branches
at the time of larval life. Indeed, various studies have shown that competition for food resources in cases of high larval density increases
the mortality rate in Cerambycids and / or decreases the body weight of adults. The cannibalism in the larval stage may occur when
competition is intense [10, 27].
The sex ratio obtained in D. trifasciata was 0.73 in favor of females. But in Cerambycidae Monochamus galloprovincialis, the
sex ratio is in favor of males [8].
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The incubation period of D. trifasciata obtained was 11.81 ± 1.22 days. These results are similar to those of [10] who reported
that the time between egg-laying and hatching of eggs’ Cerambycidae varies from 3 to 7 days in general but can go up to 25 days.
In this study, the total duration of larval development of D. trifasciata (198.2 ± 16.22 days) is close to that obtained by [28] who
reported that the larval period of Cerambycidae Monochamus leuconotus was approximately 210 days. In Anoplophora versteegi and
Aeolesthes holosericea, larval development requires respectively 263.0 ± 22.64 days and 516.9 days [29, 30]. The duration of pupal
development of D. trifasciata was 18.69 ± 0.63 days. It is similar to that recorded in Anoplophora versteegi which is 25.76 ± 2.77 days
[29]. On the other hand, this duration is 145.16 ± 29.10 days in females of Plocaederus ferrugenius [31].
After pupal development, adult stay 3.68 ± 1.03 days in the pupal chamber before emerging from the exile hole. This time is
important because many physiological changes occur in these recently emerged adults of the pupal development, including the
pigmentation of the exoskeleton. This process may take several days before the young adult emerges from the wood through a circular
hole as mentioned by [32] as well as [8].
The complete lifecycle of D. trifasciata under rearing conditions was 211.81 ± 18.87 days. These results are close to those
obtained on D. trifasciata by [3]. Indeed, these authors observed emergence of adults by mid-June from laying at the end of November,
about 210 days after laying.
V.CONCLUSION
This study was motivated by the almost total absence of data on the biological parameters of D. trifasciata. The study of the
development cycle made it possible to determine for the first time the reproductive parameters of this species. The pre-mating and pre-
oviposition periods were 123.33 ± 8.45 days and 1.83 ± 0.79 days, respectively. Lifetime fecundity 174.3 ± 18.77 eggs was obtained
with a fertility rate 82 ± 9.38%. Longevity of D. trifasciata was 184.56 ± 7.47 days in males and 196.97 ± 6.43 days in females. The
emergence rate was 8.36% and the sex ratio has been in favor of females. The results revealed that D. trifasciata is a univoltine patterns
species with a total development cycle duration of 211.81 ± 18.87 days. The knowledge of the reproductive parameters of D. trifasciata
could help identify the appropriate methods and times of intervention in developing effective control strategies.
ACKNOWLEDGMENT
The authors thank Kouamé Francis Koffi, Mathieu Zogbe, Léon Kouamé and all the producers of the study area for their
availability and help with data collection throughout the year. They thank N'guadi Blaise Allou, CSIR-TWAS fellow (FR number:
3240280453) and CSIR-NEIST, Jorhat, India for agreeing to revise the English version of the manuscript. They are also grateful to the
institution such as cotton and cashew council, African center of excellence on Climate Change, biodiversity and sustainable agriculture,
led by Professor Koné for collaboration and financial support.
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AUTHORS
First Author – Ettien Narcice AKESSE (PhD student), Félix HOUPHOUET-BOIGNY University, Center of Excellence on Climate
Change, Biodiversity and Sustainable Agriculture, Côte d'Ivoire, narcice1985@gmail.com, 00225 09 91 14 45.
Second Author – San-Whouly Mauricette OUALI-N’GORAN (Dr.), Félix HOUPHOUET-BOIGNY University, Center of
Excellence on Climate Change, Biodiversity and Sustainable Agriculture, Côte d'Ivoire, ngoransw@yahoo.fr , 00225 07 40 57 05
Correspondence Author –Ettien Narcice AKESSE, PhD student, Félix HOUPHOUET-BOIGNY University, Center of Excellence
on Climate Change, Biodiversity and Sustainable Agriculture, Côte d'Ivoire, narcice1985@gmail.com, 00225 09 91 14 45.

Reproductive Parameters of Diastocera trifasciata (Fabricius, 1775) (Coleoptera: Cerambycidae: Lamiinae), Cashew Branches Girdler in Côte d’Ivoire, Under Semi-Natural Conditions

  • 1.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 686 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org Reproductive Parameters of Diastocera trifasciata (Fabricius, 1775) (Coleoptera: Cerambycidae: Lamiinae), Cashew Branches Girdler in Côte d’Ivoire, Under Semi-Natural Conditions Ettien Narcice AKESSE* , San-Whouly Mauricette OUALI-N’GORAN* * Faculty of Biosciences/Laboratory of Zoology and Animal Biology, Center of Excellence on Climate Change, Biodiversity and Sustainable Agriculture, Félix HOUPHOUET-BOIGNY University Côte d'Ivoire DOI: 10.29322/IJSRP.8.9.2018.p8193 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 Abstract- In Côte d'Ivoire, the cashew nut still known as "brown gold" represents a hope for the populations of the producing zones and contributes to the international influence of the country. However, it is facing attacks from multiple insects including Diastocera trifasciata formerly Analeptes trifasciata. The lack of data on the reproductive parameters of the species motivated this study. The objective of this study was to provide knowledge related on the reproductive parameters of Diastocera trifasciata. The study was conducted under semi-natural conditions at a temperature of 26.83 ± 2.36 ° C and a relative humidity of 75.37 ± 15.09%. Forty-five pairs of adults were reared in insectarium to study the biology of the species. Data collected were analyzed with Statistica version 7.1 software. The study revealed a pre-mating period of 123.33 ± 8.45 days (mean ± SE) and pre-oviposition period 1.83 ± 0.79 days. Oviposition period was 64.26 ± 8.88 days with lifetime fecundity of 174.3 ± 18.77 eggs and fertility rate 82 ± 9.38%. In females, body length was 39.71 ± 2.74 mm and longevity was 196.97 ± 6.43 days. There is a positive correlation between longevity, lifetime fecundity, female size and fertility. In males, body length was 42.23 ± 3.03 mm and longevity 184.56 ± 7.47 days. Sex ratio was 0.73 in favor of females. Incubation period of eggs was 11.81 ± 1.22 days. Development duration are 198.2 ± 16.22 days for the larval stage and 18.69 ± 0.63 days for pupal stage. With a total life cycle 211.81 ± 18.87 days, D. trifasciata is univoltine species. The reproductive parameters of D. trifasciata were obtained. Knowledge of these parameters could be used to support the development of effective control strategies. Index Terms- Diastocera trifasciata, branch girdler, reproductive cycle, longevity, lifetime fecundity, fertility. I. INTRODUCTION iastocera trifasciata (Fabricius, 1775), formerly Analeptes trifasciata, is a wood-boring insect that attacks cashew trees (Anacardium occidentale, Anacardiaceae) in several areas of West Africa [1, 2]. Its damage has been observed and described for the first in 1964 in the forest station Kokondekro to Bamoro and many smallholdings around Bouake in central Côte d’Ivoire [3]. Since the work of these authors, no other study has made a case of the biology and ecology of this species to determine effective means of control. In 2014, D. trifasciata has been identified as one of the main four major insect pests of cashew in the country [2]. This insect girdle and cuts a large number of branches and trunks causing up to 55% cashew nut yield loss [4]. It represents a serious threat on the farmers' yields and the country's economy [5]. The control technique used until now by producers in Côte d’Ivoire is to capture manually adults during periods of occurrence and killing them with machetes or grilling. This practice does not make reach all individuals and proves to be very inefficient with regard to the spread of growing outbreaks. In addition, this practice is not apply in the duration and the periods of application are ill-chosen. Since the work investigated by [6] as well as those conducted by [3] which focused on the morphological description of this pest and this damage, no recent study has made any reference to the biological parameters of this species. The knowledge of biology and especially the control of reproduction is essential for the development of an effective control strategy. The objective of this work is to know the reproductive parameters of D. trifasciata. Specifically, it is to determine the egg incubation period, the duration of the development cycle, female lifetime fecundity, fertility, duration of sexual maturation of adult, emergence rate, sex ratio and adult longevity. D
  • 2.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 687 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org II. MATERIALS AND METHODS A. Study site The study was conducted in the locality of Brobo located 20 km from Bouake on the axis Bouaké-M'Bahiakro, in the center of Côte d'Ivoire (latitude: 07°38.18N, longitude: 004°49.29W). The vegetation is herbaceous savannah, shrubby or wooded with forest islets mainly in the lowlands [7]. The study period extended from October 2015 to September 2017 in semi-natural conditions. B. Material B1. Biological material The plant material used was branches and leaves of cashew obtained from the orchards of Brobo. These organs serve both as food for the individuals and as a female’s egg-laying support. The adult used come from the branches of cashew trees containing eggs of D. trifasciata kept in rearing in the insectarium. B2. Breeding device Thirty wooden cages in the form of a parallelepiped of dimensions 80 cm x 60 cm x 60 cm were used for the rearing of adults. The four (04) lateral sides are in a metal screen of fine mesh of 2 mm for ventilation of cages. The base and the roof are plywood. One of the four lateral side serves as the door for cleaning droppings and dead individuals. The plywood of the roof is free to facilitate the renewal of the branches inside the cage. The temperature inside the insectarium was 26.83 ± 2.36 ° C (mean ± SE) ranged from 22.6 - 32 ° C. The relative humidity was 75.37 ± 15.09% (mean ± SE) (range 40.94 – 83). These data were recorded by a digital thermohygrometer MEDISANA. The shots were taken with a NIKON B700 digital camera. The measure of every instar of the insect were recorded with the help of a vernier slide caliper. C. Study of reproductive parameters C1. Adult pre-mating, mating and egg-laying Forty-five pairs of adults (45 females and 45 males) newly emerged from the branches collected in different fields were observed in fifteen (15) cages at the rate of three pairs per cage. The body length of each adult was measured from the top of the head to the apex of the elytra using a vernier slide caliper. The date of emergence is noted. Individuals emerged from the same branch and from the same orchard were never put in the same cages to avoid consanguinity [8]. In each cage, small fresh branches of 10 to 15 mm in diameter and dried leaves of cashew trees are provided to insects as food. These branches are renewed every two days until observation of the first mating. The date of the first mating is noted. Pre-mating period, which is time elapsed between the adults’ date of emergence and that of the first mating of each pair was recorded. For mating and egg-laying experiments, as soon as the first mating are observed, the pair is isolated in another cage. In each cage, a branch of larger size whose circumference is between 15 and 25 cm is made available to them. This branch serves both food for the pair and female support for laying. Daily, branches are removed from cages and inspected to note the presence or not of laying. Oviposition pit is marked by the presence of gelatinous substance at the entrance of the holes [8, 9]. Egg counting is based on the presence of this substance. In case of death of the male before the female, it is replaced by another and observations continue. The dates and times of mating and laying are observed and noted until the death of the female. The date of female death is noted. The Preoviposition period (elapsed time between first mating and first oviposition) the oviposition period which is elapsed time between first and last oviposition were determined.
  • 3.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 688 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org The number of eggs deposited per female in the branches was recorded daily. The experiment continued until the death of the female. Lifetime fecundity (total number of eggs laid by a female during its life) was determined. After the laying, the oviposition pit are inspected from the outside every day until the observation of wood dust coming out of these holes. Wood dust is powder from wood rejected by the larva during its feeding activity in the branch. As soon as detection of this wood dust, the pit is then opened to observe first instar larvae. Number of hatched and unhatched eggs was counted and egg fertility rate (percentage of the number of eggs hatched to the total number of eggs laid) was determined. The longevity (elapsed time between date of emergence and the date of death) was determined. C2. Emergence rate and sex ratio The emergence rate was studied on 76 fresh branches of average length 149.31 ± 53.34 cm collected in different orchards. These branches contain eggs newly laid. The number of eggs laid and laying date are recorded on each branch with permanent marker. They are then brought back to an insectarium to be stored on tables. They are followed until the emergence of adults. The number of emerged adults was noted. Emergence rate and sex ratio (number of males emerged on number of females emerged) were calculated. C3. Incubation period, larval and pupal development of D. trifasciata During attacks, 100 freshly girdled branches containing eggs were collected from different orchard of Brobo. These selected branches are those cut the day before the visits. They are recognized by the fresh state of the substance covering the oviposition pit. They were divided into 50 lots of 02 branches. The eggs are observable by superficial opening of the oviposition pit with a knife. On each branch, laying date and the number of notch containing the egg are recorded in the indelible marker. According to [10], the time between egg-laying and hatching of Cerambycidae eggs in general ranges from 3 - 7 days but can be up to 25 days. A batch of two (02) branches is dissected every three (03) days during the first month following the laying date to observe the evolution of the eggs. The number of hatched eggs was noted at each observation and the date of hatching was also on each branch. Incubation period (time elapsed between egg laying and hatching) has been recorded. The determination of larval and pupal duration was made from the 50 lots of branches formed. After hatching, the branches are dissected every seven (07) days for seven (07) months to see the evolution of the larvae until observation of pupae and adult emergence. After dissection of the branches, the shape, the dimensions and the color of the observed stages are described. The following parameters were recorded: duration of larval development (elapsed time between the date of hatching of the eggs of the moulting pupal), duration of pupation (time that covers the process that allows the passage of larvae of last stage to the pupa), duration of pupal development (elapsed time between the date of the pupal moulting and that of obtaining the adult), duration of development cycle (elapsed time between the date of egg laying and obtaining of the adult stage). D. Data Analysis Data processing was done using Statistica software version 7.1 StatSoft [22]. A one-way analysis of variance (ANOVA) followed by the Newman and Keuls test at the 5% threshold allowed the averages to be compared. Linear regression analyses (Pearson’s correlation) was used to test the correlation between some reproductive variables in females (lifetime fecundity, fertility, longevity, oviposition period, and female length) and in males (pre-mating period, longevity and body length).
  • 4.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 689 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org III. RESULTS A. Pre-mating and pre-oviposition periods of adults The individuals made the first mating between the 104th and 138th days after emergence, pre-mating period of 123.33 ± 8.45 days (mean ± SE). Mating takes place during the day. The first oviposition was observed between the 1st and 4th day after mating, pre-oviposition period of 1.83 ± 0.79 days (mean ± SE). Pre-mating and pre-oviposition periods are not correlated with any of the biological parameters studied in the female in this study (Table 1). Table 1: Linear regression analyses (Pearson’s R correlation matrix) between the different parameters of the female of D. trifasciata in semi-natural conditions (temperature 26.83 ± 2.36 °C and relative humidity 75.37 ± 15.09 %) Parameters Pre-mating period (days) Longevity (days) Oviposition period (days) Pre- oviposition period (days) Lifetime fecundity Body length (mm) Pre-mating period (days) 1 Longevity (days) 0.356 ns 1 Oviposition period (days) -0.058 ns 0.315 ns 1 Pre-oviposition period (days) -0.058 ns 0.114 ns -0.087 ns 1 Lifetime fecundity 0.291 ns 0.958‫٭‬ 0.265 ns 0.181 ns 1 Body length (mm) 0.270 ns 0.813‫٭‬ 0.155 ns 0.259 ns 0.875‫٭‬ 1 Fertility 0.076 ns 0.477‫٭٭‬ 0.608‫٭‬ -0.035 ns 0.518‫٭٭‬ 0.424‫٭٭٭‬ Significance of P-values: ns: non-significant: ‫٭‬P < 0.0001; ‫٭٭‬P < 0.01; ‫٭٭٭‬P < 0.05 B. Oviposition period, lifetime fecundity, fertility rate and longevity of adults Egg-laying begin at dusk and end at dawn. Oviposition period was 64.26 ± 8.88 days (mean ± SE). Female lifetime fecundity was 174.3 ± 18.77 eggs, ranged from 126 - 200 eggs. On 174.3 ± 18.77 eggs laid per female, 143.46 ± 22.92 hatched, fertility rate was 82 ± 9.38% (mean ± SE). Males of D. trifasciata emerge firstly but a longevity was 184.56 ± 7.47 days (range 168 – 201). In females, longevity was 196.97 ± 6.43 days (range 178 - 208) (Table 2). The linear correlation test shows that there is a positive correlation between female lifetime fecundity, longevity and egg fertility (Figure 1) (Table 1).There are significant difference between the longevity in males and females (F = 47.43, ddl = 1, P < 0.05).
  • 5.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 690 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org C. Correlation between adult length, fecundity and longevity In female, body length was 39.71 ± 2.74 mm (means ± SE), ranged 33.10 - 43.50. The linear correlation test between female length and lifetime fecundity shows that large females lay more eggs (R = 0.87, P < 0.05) (Figure 2). Female length is also positively correlated with longevity (R = 0.81, P < 0.05) (Figure 3). There were mating between some females who lost their male partners and newcomers. Females were therefore polyandrous and males were polygynous. In males, body length was 42.23 ± 3.03 mm (range 31.49 - 46.20). Males are larger in size than females. Analysis of variance at the 5% threshold showed that there is a significant difference between the body length in males and females (F = 11.38, ddl = 1, P < 0.05). The sexual dimorphism is marked by the length of the antennae which is 59.72 ± 4.88 mm in the males against 47.98 ± 6.77 mm in the females (Table 2). Male length correlated neither with longevity (P > 0.05, R2 = 0.002) nor with the pre-copulation period (P > 0.05, R2 = 0.001). Figure 1: Correlation between female longevity and lifetime fecundity 175 180 185 190 195 200 205 210 Longevity (days) 120 130 140 150 160 170 180 190 200 210 Lifetimefecundity R2 = 0,91 y = 2,8x - 360,6
  • 6.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 691 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org Figure 2: Correlation between female body length and lifetime fecundity 32 34 36 38 40 42 44 Female body length (mm) 120 130 140 150 160 170 180 190 200 210 Lifetimefecundity R2 = 0,76 y = 6,02x - 66,63 Figure 3: Correlation between longevity and female body length 170 175 180 185 190 195 200 205 210 Longevity (days) 28 30 32 34 36 38 40 42 44 Femalebodylength(mm) R2 = 0,66 y = 0,34x - 28,45
  • 7.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 692 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org Table 2: Some morphological characteristics of adults of D. trifasciata Sex Body length (mm) Antennae length (mm) Longevity (days) Males 42,23 ± 3,03 (31,49 - 46,20 59,72 ± 4,88 (50,8 - 69,95) 184,56 ± 7,47 (168 - 201) Females 39, 71 ± 2,74 (33,10 - 43,50) 47,98 ± 6,77 (38,18 - 59,41) 196,97 ± 6,43 (178 - 208) n = 45 in each sex; Mean ± SE (Standard Error of the mean); (minimum – maximum values) D. Emergence rate and sex ratio of D. trifasciata Out of a total of 2953 eggs monitored, only 247 adult individuals have emerged, an average emergence or rate of 8.36%. The emerged adults are divided into 142 females and 105 males, a sex ratio of 0.73 in favor of females. E. Egg of D. trifasciata and incubation period The laying were recorded between 18h and 07h in the morning. The eggs are laid in notches made in the bark by the females only. Each notch of eggs contains only one egg disposed parallel to the longitudinal axis of the branch or stem cut (Figure 4a-b). The newly laid of D. trifasciata egg is white, oval and similar to a rice grains (Figure 4 b). Eggs length were 5.52 ± 0.23 mm (mean ± SE), ranging from 5.04 - 6.1 mm. The width of the eggs ranged 1.05 - 1.71 mm with a mean of 1.28 ± 0.11 mm. The incubation period of the eggs was 11.81 ± 1.22 days (range 9 – 15). (a) (b) Figure 4: Laying site and fresh eggs of D. trifasciata (a) Laying notch (red encircled); (b) egg (red encircled) still inlaid in the branch
  • 8.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 693 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org F. Larval and pupal development of D. trifasciata The first instar larvae of D. trifasciata are translucent rosy white color. The body length of larvae (mean ± SE) was 6.39 ± 1.20 mm (range 4.80 - 8.70). The body width was 1.34 ± 0.16 mm (range 1.01 - 1.77). The length of late-instar larvae was 36.45 ± 4.56 mm (range 27.60 - 48.12). The width of late-instar larvae was 7.72 ± 1.2 mm (range 5.8 - 10.60). They are cream-white tinged with yellow and it is located under the bark. The duration of larval development was 198.2 ± 16.22 days (range 169 - 226). Before the end of their development, the larvae reduce their gallery in the wood to construct a pupal chambers. At the ends of this chamber are piled woody debris undigested that look like sawdust. Pupation and pupal development occurs at the end of the larval development in this pupal chamber. The length of the pupal chamber was 89.26 ± 13.01 mm (range 55.1 – 135). Its width was 24.95 ± 3.36 mm (range 15.7 - 34.9). The pupation duration was (mean ± SE) 9.02 ± 2.13 days. The pupa is white or black according on whether it is at beginning or end of its development. The head is inflected toward the ventral side and carries the antennae which are elongated dorsally along the body before wrapping in a spiral at the end on the legs. The wing cases are concealed under the folded legs on the ventral side. The length of the pupae was (mean ± SE) 38.64 ± 2.99 mm (range 32.5 - 46.9). The width was 10.66 ± 1.11 mm (range 8.1 - 14.7). Pupal development occurred between days 16 and 19 after pupation, for an average duration of 18.69 ± 0.63 days. The young imago who comes out, the wings and the whole body were soft. The head, legs and abdomen have a grey and black color at the joints. The bands of the wings have a grey coloration. The average duration of pigmentation is 3.68 ± 1.03 days (mean ± SE). G. Duration of the development cycle of D. trifasciata The duration of the life cycle was 211.81 ± 18.87 days (range 167 – 240). D. trifasciata is therefore a species univoltine. The adult, once the pigmentation is complete, emerges through a circular hole made with his mandibles in the bark covering pupal chamber. The diameter of these exit hole was 17.63 ± 2.43 mm (mean ± SE) (range 11.4 - 28.1) (Figure 5).
  • 9.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 694 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org F Pigmentation 3.68 ± 1.03 days Preoviposition 1.83 ± 0.79 days Premating 123.33 ± 8.45 days Incubation period 11.81 ± 1.22 days Larval development 198.2 ± 16.22 days Pupation 9.02 ± 2.13 days Pupal development 18.69 ± 0.63 days ♀ ♂ (b) (h)(g)(i) (j) a (f)(e) (c) (d) (a) (k) Figure 5. Life cycle of Diastocera trifasciata (Coleoptera: Cerambycidae: Lamiinae) under semi-natural conditions (temperature 26.83 ± 2.36 ° C and relative humidity 75.37 ± 15.09%. (a): egg (red encircled) observed after opening of a notch; (b): first instar larva (red arrowed); (c) : late instar larva lateral view; (d): late instar larva ventral view; (e): pupa lateral view; (f): pupa ventral view; (g): young imago (not emerged from branch) dorsal view; (h): young imago (not emerged from branch) ventral view; (i): adults emerged; (j): mating on cashew branch; (k): female laying in a cashew branch. Scale a: 0.79 cm
  • 10.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 695 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org IV.DISCUSSION In this study, the results revealed that the pre-mating period was 123.33 ± 8.45 days. This period is very high compared to that of Cerambycidae of the Lamiinae subfamily, whose adults feed from a few days to three weeks before becoming sexually mature [8, 12]. Among Cerambycidae Anoplophora glabripennis (Motschulsky), adult females become sexually mature about 9 to 17 days after emergence. The pre-mating period of D. trifasciata is explained by the fact that Cerambycidae Lamiinae are synovigenic pattern that do not mate immediately after emergence. Adults emerge with immature sexual organs. They must feed on young shoots and dry leaves for a period ranging from several days to weeks before reaching sexual maturity [8, 13]. The duration of this period may also depend on the phenology of the host plant Anacardium occidentale. Indeed, the mating period begins when the plant begins its push vegetative pre-flowering and extends until flowering. There would be a synchronization between the reproductive period of D. trifasciata and that of the host plant. According to [14], this phenomenon of synchronization of the cycle of insects and their host plant is frequent and determines the quantity and the quality of the resources available for reproduction. The pre-oviposition period obtained in this study (1.83 ± 0.79 days) is close to that obtained in Cerambycidae Cerambyx welensii which is 2.8 ± 0.2 days, investigated by [15]. The oviposition period of D. trifasciata recorded was 64 ± 8.88 days. These observations are close to those of Cerambycidae Lamiinae that have an oviposition period of 60 ± 7 days [14]. On the other hand, in the species of the subfamily Cerambycinae, the oviposition period was 41 ± 17 days [15]. Regarding female’s lifetime fecundity (174.3 ± 18.77 eggs) in breeding condition is superior to the lifetime fecundity in Cerambycidae Lamiinae which is 155 ± 30 eggs [14]. [16] showed a higher fecundity of 284 ± 27 eggs in the female Cerambycinae Phoracantha recurva. The positive correlation between fecundity and female size shows that larger females lay more eggs than smaller females. According to [17], female size is usually a good indicator of potential fecundity. The phenomena of polyandry and polygyny were observed respectively in females and males of D. trifasciata as in Cerambyx cerdo [18]. The egg fertility rate recorded in D. trifasciata was 82.37 ± 9.38%. This result is close to those of Cerambycidae such as Anoplophora glabripennis which is 83 ± 2% [19], Cerambyx cerdo 78 ± 1% [17]. On the other hand, other authors have recorded a fertility rate ranging from 90 to 98%, in Monochamus galloprovincialis and Phoracantha semipunctata [16, 20]. Rates of between 48- 66% were in Anoplophora glabripennis [21]. Adult males measure 42.23 ± 3.03 mm and live 184.56 ± 7.47 days while females measure 39.71 ± 2.74 mm and live 196.97 ± 6.43 days. This difference in longevity has been reported by various authors. In Cerambycidae, females generally live longer (54 ± 6 days) than males (42 ± 6 days) for a given species [15, 22]. Adults’ longevity of D. trifasciata could be influenced by the long period of nutrition that precedes reproduction. According to [23], the adult phase of Cerambycidae is entirely devoted to reproduction. Female longevity was positively correlated with lifetime fecundity, egg fertility. Similar results have been obtained in other Cerambycidae [15, 24]. Longevity can also be influenced by body size, although factors such as temperature, diet, and overall health of an individual may play a larger role [25]. The emergence rate obtained in this study was 8.36%. These results are quite similar to those of [26] who recorded an emergence rate of 12% in adults of Cerambycidae Monochamus carolinensis. These findings could be justified by competitions within the branches at the time of larval life. Indeed, various studies have shown that competition for food resources in cases of high larval density increases the mortality rate in Cerambycids and / or decreases the body weight of adults. The cannibalism in the larval stage may occur when competition is intense [10, 27]. The sex ratio obtained in D. trifasciata was 0.73 in favor of females. But in Cerambycidae Monochamus galloprovincialis, the sex ratio is in favor of males [8].
  • 11.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 696 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org The incubation period of D. trifasciata obtained was 11.81 ± 1.22 days. These results are similar to those of [10] who reported that the time between egg-laying and hatching of eggs’ Cerambycidae varies from 3 to 7 days in general but can go up to 25 days. In this study, the total duration of larval development of D. trifasciata (198.2 ± 16.22 days) is close to that obtained by [28] who reported that the larval period of Cerambycidae Monochamus leuconotus was approximately 210 days. In Anoplophora versteegi and Aeolesthes holosericea, larval development requires respectively 263.0 ± 22.64 days and 516.9 days [29, 30]. The duration of pupal development of D. trifasciata was 18.69 ± 0.63 days. It is similar to that recorded in Anoplophora versteegi which is 25.76 ± 2.77 days [29]. On the other hand, this duration is 145.16 ± 29.10 days in females of Plocaederus ferrugenius [31]. After pupal development, adult stay 3.68 ± 1.03 days in the pupal chamber before emerging from the exile hole. This time is important because many physiological changes occur in these recently emerged adults of the pupal development, including the pigmentation of the exoskeleton. This process may take several days before the young adult emerges from the wood through a circular hole as mentioned by [32] as well as [8]. The complete lifecycle of D. trifasciata under rearing conditions was 211.81 ± 18.87 days. These results are close to those obtained on D. trifasciata by [3]. Indeed, these authors observed emergence of adults by mid-June from laying at the end of November, about 210 days after laying. V.CONCLUSION This study was motivated by the almost total absence of data on the biological parameters of D. trifasciata. The study of the development cycle made it possible to determine for the first time the reproductive parameters of this species. The pre-mating and pre- oviposition periods were 123.33 ± 8.45 days and 1.83 ± 0.79 days, respectively. Lifetime fecundity 174.3 ± 18.77 eggs was obtained with a fertility rate 82 ± 9.38%. Longevity of D. trifasciata was 184.56 ± 7.47 days in males and 196.97 ± 6.43 days in females. The emergence rate was 8.36% and the sex ratio has been in favor of females. The results revealed that D. trifasciata is a univoltine patterns species with a total development cycle duration of 211.81 ± 18.87 days. The knowledge of the reproductive parameters of D. trifasciata could help identify the appropriate methods and times of intervention in developing effective control strategies. ACKNOWLEDGMENT The authors thank Kouamé Francis Koffi, Mathieu Zogbe, Léon Kouamé and all the producers of the study area for their availability and help with data collection throughout the year. They thank N'guadi Blaise Allou, CSIR-TWAS fellow (FR number: 3240280453) and CSIR-NEIST, Jorhat, India for agreeing to revise the English version of the manuscript. They are also grateful to the institution such as cotton and cashew council, African center of excellence on Climate Change, biodiversity and sustainable agriculture, led by Professor Koné for collaboration and financial support. REFERENCES [1] E. U. Asogwa, T.C.N. Ndubuaku, A. T. Hassan, “Distribution and damage characteristics of Analeptes trifasciata Fabricius 1775 (Coleoptera: Cerambycidae) on cashew (Anacardium occidentale Linnaeus 1753) in Nigeria.” Agriculture and Biology Journal of North America. 2011, 2(3): 421-431. [2] O. R. N’Dépo, M. Chérif, F. Johnson, K. F. JM. Kassi, A. C. N’Guessan, N. Silué, E. N. Akessé, D. Koné, O. M. N’Goran, “Inventaire des insectes ravageurs du verger anacardier dans les régions de Bounkani, Gontougo et Indénie-Djablun au Nord-Est en Côte d’Ivoire.” Afrique Sciences. 2017, 13: 333–343. [3] F. Brunck, J. P. Fabre, “Note sur Analeptes trifasciata Fabricius, Coléoptère cérambycide, grave ravageur d'Anacardium occidentale en Côte d'Ivoire.” Bois ET Forêt des Tropiques. 1970, 134: 15–19.
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    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 697 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org [4] Q. Wang, “Cerambycid Pests in Agricultural and Horticultural Crops,” In Cerambycidae of the World: Biology and Pest Management. 1st ed, Q. Wang, London: CRC Press. 2017, pp. 409-562. [5] E. N. Akessé, S. W. M. Ouali-N’goran, O. R. N’Dépo, T. Koné, D. Koné, “Prospective Study of the Insect Fauna Associated with Anacardium occidentale L. (Salpindales: Anacardiaceae) in Five Producing Areas of Côte d’Ivoire,” In Proceedings on the Third International Cashew Conference: Cashew for Health Wealth and Environment. Held at Serena Hotel, Dar Es Salaam, Tanzania. 2015, pp: 164–172. [6] H. Roberts, “Further observations on the biology of Analeptes trifasciata (P.) 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Keena, “Anoplophora glabripennis (Coleoptera: Cerambycidae) fecundity and longevity under laboratory conditions: comparison of populations from New York and Illinois on Acer saccharum.” Environmental Entomology. 2002, 31: 490–498. [20] P. M. Naves, E. M. De Sousa, J. A. Quartau, “Reproductive traits of Monochamus galloprovincialis (Coleoptera: Cerambycidae) under laboratory conditions.” Bulletin of entomological research. 2006, 96: 289–294. [21] M.T. Smith, J. Bancroft, J. Tropp, “Age-specific fecundity of Anoplophora glabripennis (Coleoptera: Cerambycidae) on three tree species infested in the United States.” Environmental Entomology. 2002, 31: 76–83. [22] R. A. Haack, M. A. Keena, D. Eyre, “Life History and Population Dynamics of Cerambycids,” in Cerambycidae of the World: Biology and Pest Management, 1st ed, Q. Wang, London: CRC Press, 2017, pp. 71-103. [23] A. Berkov, “Do Lecythidaceae Specialists (Coleoptera, Cerambycidae) Shun Fetid Tree Species?” Biotropica. 2000, 32: 440–451. [24] K. Togashi, J. Appleby, H. Oloumi-Sadeghi, R. B. Malek, “Age-specific survival rate and fecundity of adult Monochamus carolinensis (Coleoptera: Cerambycidae) under field conditions.” Applied Entomology and Zoology. 2009, 44: 249–256. [25] N. Sokolovska, L. Rowe, F. Johansson, “Fitness and body size in mature odonates.” Ecological Entomology. 2000, 25: 239–248. [26] S. Akbulut, W. T. Stamps, M. J. Linit, “Population dynamics of Monochamus carolinensis (Col., Cerambycidae) under laboratory conditions.” Journal of Applied Entomology. 2004, 128:17–21. [27] M. L. Richardson, R. F Mitchell, P. F. Reagel, L. M. Hanks, “Causes and consequences of cannibalism in noncarnivorous insects.” Annual Review of Entomology. 2010, 55: 39–53. [28] J. M. Gichuhi, “Rearing Method and Life History of Laboratory Bred African Coffee White Stem Borer, Monochamus leuconotus (Pascoe) (Coleoptera: Cerambycidae).” Ph. D Thesis, School Of Biological Sciences, University Of Nairobi, Kenya. 2016, 97 [29] K. N. S. Saikia, A. Thakur, A. Alemla, “Biology of Citrus Trunk Borer Anoplophora versteegi Rits. (Coleoptera: Cerambycidae) under Laboratory Conditions.” Indian Journal of Hill Farming. 2011, 24; 19-24. [30] R. Gupta, J. S. Tara, “First record on the biology of Aeolesthes holosericea Fabricius, 1787 (Coleoptera: Cerambycidae), an important pest on apple plantations (Malus domestica Borkh.) in India.” Munis Entomology & Zoology. 2013, 8: 243–251.
  • 13.
    International Journal ofScientific and Research Publications, Volume 8, Issue 9, September 2018 698 ISSN 2250-3153 http://dx.doi.org/10.29322/IJSRP.8.9.2018.p8193 www.ijsrp.org [31] P. T. Vasanthi, N. Raviprasad, “Biology and Morphometrics of Cashew Stem and Root Borers (CSRB) Plocaederus ferrugenius and Plocaederus obesus (Coleoptera: Cerambycidae) Reared on Cashew Bark.” International Journal of Science and Research. 2013, 3:1-7. [32] H. F. Paulino-Neto, J. Vasconcellos-Neto, S. M. Carmello-Guerreiro, “The biology of Oncideres humeralis Нorms (Coleoptera: Cerambycidae: Lamiinae) and new Cerambycidae-Melastomataceae host-plant associations.” Studies on Neotropical Fauna and Environment. 2006, 41: 227–233. AUTHORS First Author – Ettien Narcice AKESSE (PhD student), Félix HOUPHOUET-BOIGNY University, Center of Excellence on Climate Change, Biodiversity and Sustainable Agriculture, Côte d'Ivoire, [email protected], 00225 09 91 14 45. Second Author – San-Whouly Mauricette OUALI-N’GORAN (Dr.), Félix HOUPHOUET-BOIGNY University, Center of Excellence on Climate Change, Biodiversity and Sustainable Agriculture, Côte d'Ivoire, [email protected] , 00225 07 40 57 05 Correspondence Author –Ettien Narcice AKESSE, PhD student, Félix HOUPHOUET-BOIGNY University, Center of Excellence on Climate Change, Biodiversity and Sustainable Agriculture, Côte d'Ivoire, [email protected], 00225 09 91 14 45.