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J. Bio. Env. Sci. 2019
15 | Ningatoloum et al.
RESEARCH PAPER OPEN ACCESS
Pollination efficiency of Chalicodoma cincta (Fabricius)
(Hymenoptera: Megachilidae) on Cajanus cajan (L.) Millsp.
(Fabaceae) flowers at Doyaba (Sarh, Chad)
Clautin Ningatoloum*1,2
, Guiguindibaye Madjimbe3
, Sidonie Fameni Tope4
,
Fernand-Nestor Tchuenguem Fohouo1
1
Laboratory of Applied Zoology, Department of Biological Sciences, Faculty of Science,
University of Ngaoundéré, Ngaoundéré, Cameroon
2
Department of Biological Sciences, University Adam Barka of Abéché, Abéché, Chad
3Department of Environmental Sciences, University Institute of Sciences Agronomic and the Environment,
Sarh, Chad
4
Laboratory of Zoology, Department of Biological Sciences, Faculty of Science,
University of Maroua, Maroua, Cameroon
Article published January 26, 2019
Key words: Cajanus cajan, Chalicodoma cincta, Flowers, Yield, Pollination.
Abstract
To evaluate Chalicodoma cincta impact on pod and seed yields of Cajanus cajan, its foraging and pollinating
activities were studied in Doyaba, during the rainy season of 2015 and 2016. Each year, treatments included
flowers accessible to all visitors, bagged flowers to avoid insect visitors, bagged flowers using gauze bags destined
to be visited exclusively by C. cincta and bagged flowers destined to opening and closing without the visit of
insects or any other organism. For each year of study, observations were made on 1028 ± 90 flowers per
treatment. Chalicodoma cincta daily rhythm of activity, its foraging behaviour on flowers and its pollination
efficiency were evaluated. On flowers, individual bees intensely harvested exclusively nectar. The fruiting rate, the
number of seeds per pod and the percentage of normal seeds of unprotected flowers were significantly higher
than those of flowers protected from insects. Through its pollination efficiency, C. cincta provoked a significant
increment of the fruiting rate by 21.40% and 7.55%, the number of seeds per pod by 16.69% and 14.96% and the
percentage of normal seeds by 32.95% in 2015 and 36.30% in 2016 respectively. The Conservation of C. cincta
nests close to C. cajan fields is recommended to improve pod and seed productions in the region.
*Corresponding Author: Clautin Ningatoloum  gclautin@gmail.com
Journal of Biodiversity and Environmental Sciences (JBES)
ISSN: 2220-6663 (Print) 2222-3045 (Online)
Vol. 14, No. 1, p. 15-25, 2019
http://www.innspub.net
J. Bio. Env. Sci. 2019
16 | Ningatoloum et al.
Introduction
Several plant species depend on insect pollinators for
their reproduction; in agro ecosystems, these
pollinators have a great ecological and economic
importance because they influence positively the plant
production (McGregor, 1976; Philippe, 1991;
Tchuenguem, 2005). Cajanus cajan is an ideal pulse
crop of rainfed tropics and subtropics (Saxena et al.,
2002). Its grows up right to 4 m (Niyonkuru, 2002).
The leaves are generally trifoliate; flower is pink, but
can vary from white to red (ICRISAT, 1981) and
produces nectar and pollen which attract insects
(Grewal et al., 1990; Saxena et al., 1990; Reddy et al.,
2004; Sarah et al., 2010).
Indian is the largest producers of pigeon pea in the
world (Kimani, 2000). The fruit is a pod containing
four raw of seven seeds (Pando et al., 2011b). Seeds
contain 21 to 30% proteins important for human’s
diets (Sharma and Green, 1980; Gupta et al., 2001;
Saxena et al., 2002). Cajanus cajan flowers were
reported to produce fewer seeds per pod in the
absence of insect pollinators in Great Britain
(Kendall and Smith, 1976) and in Cameroon (Pando
et al., 2011b).
The research conducted in the United Stated of
America (Grewal et al., 1990; Ibarra-Perez et al.,
1999) and in Cameroon (Pando, 2012; Mazi, 2015)
has revealed that bees of the genus Chalicodoma and
Megachile visits C. cajan flowers and collect nectar
and pollen. In Chad, the quantity of Ca. cajan
available to consumers is very low, the demand for
pigeon pea seeds is high, and its pod and seed yields
are weak because notably of the insufficiency of
knowledge on its relationships with anthophilous
insects in general and C. cincta in particular.
Therefore, it is important to investigate how the
production of this plant could be increased in Chad.
Prior to these studies, no previous research has
been reported on the interactions between Ca. cajan
and insects in Chad. The main objective of this work
is to contribute to the understanding of the
relationships between C. cajan and C. cincta, for
their optimal management in this country. Specific
objectives are to: (a) study the activity of this
Megachilidae on C. cajan flowers, (b) evaluate the
impact of flowering insects including Ch. cincta on
pollination and fruit and seed yields of this
Fabaceae, (c) estimate the pollination efficiency of C.
cincta on this plant species.
Materials and methods
Site and biological materials
The studies were conducted twice, June to November
in 2015 and 2016 in a field located at Doyaba, Moyen-
Chari Region of Chad. This Region belongs to the
Sudanian agro-ecological zone (Cabot and Bouquet,
1972). The climate is tropical type with two seasons: a
dry season (November to April) and a rainy season
(May to October).
The mean annual temperature is 28°C; the mean
rainfall is 1100 mm (Madjimbé, 2013). The
experimental plot was an area of 437 m2 (Latitude:
09° 04.875’ N, Longitude: 018° 25.721’ E, Altitude
363.3 m.a.s.l.). The animal material was represented
by insects naturally present in the environment.
The vegetation was represented by wild and cultivated
species. The plant material was represented by the
seeds of C. cajan variety ICPL 9145, provided by the
Agronomic Institute of the University of Sarh.
Sowing and weeding
On June 14th, 2015 and 2016 the experimental plot
was divided into eight subplots (8*4.5 m2). Seeds
were sown on 16 lines per subplot; each line had nine
holes and each hole received three seeds.
The spacing was 50 cm between rows and 50 cm on
rows. Each hole was 5 cm depth. Two weeks after
germination, the plants were thinned and only two
were left per hole (Pando et al., 2011b). Weeding was
performed manually as necessary to maintain
subplots weeds-free (Tchuenguem et al, 2009a;
Pando et al., 2013).
J. Bio. Env. Sci. 2019
17 | Ningatoloum et al.
Determining the reproduction mode
In October 12th, 2015, 2282 flowers of Ca. cajan at the
bud stage were labeled, among which 1136 were
allowed for treatment 1 (open pollination) and 1146
for treatment 2 (bagged with gauze bags to prevent
visitors) (Tchuenguem, 2005; Delaplane et al., 2013)
(Fig. 1). In October 11th, 2016, 2039 flowers at the bud
stage were labeled, among which 992 were provided
for treatment 3 (open pollination) and 1047 flowers
for treatment 4 (bagged with gauze bags). For each
year, ten days after the wilting of the last flower, the
number of pods formed in each treatment was
counted. For each treatment, the fruiting index (Ifr)
was then calculated using the following formula: Ifr
= (F1/F2), where F1 is the number of pods formed and
F2 the number of flowers initially labeled
(Tchuenguem et al., 2001). For each year, the out
crossing rate (TC) was calculated using the formula:
TC = {[(Ifra – Ifrb /fra]*100}, where Ifra and Ifrb are
the mean fruiting indexes in treatment a (flowers in
open pollination) and treatment b (bagged flowers)
respectively (Demarly, 1977). The rate of self-
pollination in the broad sense (TA) was calculated
using the formula: TA = (100 - TC) (Tchuenguem,
2005; Népidé and Tchuenguem, 2016).
Study of the activity of Chalicodoma cincta on
Cajanus cajan flowers
Observations were conducted on individually opened
flowers of treatments 1 and 3, each day, from October
13th to November 03th 2015 and from October 12th to
November 04th 2016, at one hour interval, from 6 am
to 15 pm (6-7 am, 8-9 am, 10-11 am, 12-13 pm and 14-
15 pm). In a slow walk along all labeled flowers of
treatments 1 and 3, the identity of each insect that
visited Ca. cajan flowers was noted. All insects
encountered on flowers were recorded and the
cumulated results expressed in number of visits to
determine the relative frequency of C. cincta in the
anthophilous entomofauna of C. cajan (Pando et al.,
2011b). Direct observations of the foraging activity
of this bee on flowers were made. The floral
products (nectar and/or pollen) collected by Ch.
cincta were recorded during the same dates and
time slots as that of the insect counts (Pando et al.,
2011b). The duration of visits per flower was recorded
(using stopwatch) according to five time frames (7-8
am, 9-10 am, 11-12 am, 13-14 pm and 15-16 pm). The
foraging speed (number of flowers visited by
individual bee per minute (Jacob-Remacle, 1989) was
recorded during the same date and daily periods as
the registration of the duration of visits. Abundances
(highest number of individuals simultaneously active)
per flower and per 1000 flowers (A1000) were recorded
during the same dates and time slots as that of the
registration of the duration of visits per flower. The
first parameter was recorded as a result of direct
counts. For the abundance per 1000 flowers,
bees were counted on a known number of open
flowers at the moment t. The abundance per 1000
flowers is calculated using formula: A1000 =
[(Aa/Fb)*1000], where Fb and Ab are respectively the
number of flowers and the number of C. cincta
effectively counted on these flowers at time t
(Tchuenguem et al., 2004; Tchuenguem, 2005;
Douka et al., 2017). The disruption of the activity of
individual bee by competitors or predators and the
attractiveness of other plant species with respect to C.
cincta were assessed. During the days of
investigation, the temperature and humidity of the
study site were recorded every 30 min, from 6 am to 17
pm, using a thermo hygrometer installed in the shade
(Tchuenguem et al., 2004; Farda and Tchuenguem,
2018).
Evaluation of the impact of flowering insects
including Chalicodama cincta on Cajanus cajan yield
For each investigation year, this evaluation was based
on the impact of flowering insects on pollination, the
impact of pollination on fruiting and the comparison
of yields (fruiting rate, mean number of seeds per pod
and percentage of normal i.e. well-developed seeds)
of treatment a (unprotected flowers) and treatment b
(bagged flowers).
The fruiting rate due to the influence of foraging
insects (Fri) was calculated using the formula: Fri =
{[(Fra-Frb)/Fra]*100}, where Fra and Frb are the
J. Bio. Env. Sci. 2019
18 | Ningatoloum et al.
fruiting rate in treatments a and b respectively.
The fruiting rate of a treatment (Fr) is Fr =
[(F2/F1)*100], where, F2 is the number of pods
formed and F1 the number of viable flowers initially
set (Tchuenguem et al., 2001). At maturity, pods were
harvested from all treatments.
The mean number of seeds per pod and the
percentage of normal seeds were then calculated for
each treatment.
Assessment of the pollination efficiency of
Chalicodoma cincta on Cajanus cajan
In 2015, along with the development of treatments 1
and 2, 1976 flowers at bud stage were protect and two
treatments was formed: treatments 5, with 1052
flowers protected using gauze bags to prevent insect
visitors and destined exclusively to be visited by Ch.
cincta; treatment 6, with 924 flowers protected from
insects using gauze bags.
In 2016, along with the realization of treatments 3
and 4, 1940 flowers at the bud stage were labeled and
two treatments was formed: treatment 7 with, 893
flowers protected from insects using gauze bags and
destined exclusively to be visited by C. cincta;
treatment 8 with 1047 flowers protected from insects
using gauze bags.
As soon as the first flowers were opened, in
treatments 5 and 7, the gauze bag was delicately
removed and flowers were observed for up to 10 min.
Flowers visited by C. cincta was marked and then
protected once more. For treatment 6 and 8, as soon
as the firsts flower was opened, the gauze bag was
delicately removed and flowers were observed for up
to 10 min while avoiding the visits by any other
organism.
At the maturity, pods were harvested and the
contribution of Ch. cincta in fruiting, number of seeds
per pod and percentage of normal seeds was then
calculated using data of treatments 3 or 8 (flowers
visited exclusively by C. cincta) and those of
treatments 2 or 7 (bagged flowers).
Data analysis
Data were subjected to descriptive statistics, student’s
t-test for the comparison of means of the two
samples, ANOVA for the comparison of means of
more than two samples, correlation coefficient (r) for
the study of the association between two variables,
and chi-square (χ2) for the comparison of
percentages, using Microsoft Excel 2010 software.
Results
Reproduction mode of Cajanus cajan
The podding indexes were 0.92, 0.73, 0.84 and 0.74,
in treatments 1, 2, 3 and 4 respectively. Consequently,
in 2015, the allogamy rate was 20.85% and the
autogamy rate was 79.15%; for 2016, the
corresponding data were 11.90% and 88.10%. It
appears that the Ca. cajan variety used in our
experiment has a mix reproduction mode,
autogamous-allogamous with the predominance of
autogamy over allogamy.
Activity of Chalicodoma cinta on Cajanus cajan
flowers
Seasonal frequency of visits: Among the 2317 and
2373 visits of 16 and 14 insect species recorded on C.
cajan flowers in 2015 and 2016 respectively, C. cincta
was the most represented insect with 371 visits
(15.35%) in 2015 and 383 visits (16.14%) in 2016
(Table 1). The difference between these two
percentages is not significant (χ2 = 0.87; df = 1; P >
0.05).
Flower substances harvested: From our field
observations of the years 2015 and 2016 C. cincta
were found to collect regularly and exclusively nectar
in the flowers of C. cajan (Fig. 2).
Abundance of Chalicodoma cincta: In 2015, the
highest mean number of Ch. cincta simultaneous in
activity was one per flower (n = 230; s = 0) and
188.08 per 1000 flowers (n = 230; s = 154.84). In
2016, the corresponding results were one per flower
(n = 198; s = 0) and 189.65 (n = 198; s = 198.65) per
1000 flowers.
J. Bio. Env. Sci. 2019
19 | Ningatoloum et al.
The difference between the mean number of C. cincta
per 1000 flowers of the two years was not significant
(t = 1.08; df = 426; P > 0.05).
Duration of visits per flower: In 2015, the mean
duration of individual Ch. cincta visit per Ca. cajan
flower was 9.00 sec (n = 1168; s = 8.9). The
corresponding index was 8.36 sec (n = 704; s = 8.27)
in 2016. The difference between these two means is
higher significant (t = 20.56; df = 292; P < 0.001).
Table 1. Diversity of floral insects on Cajanus cajan in 2015 and 2016 at Doyaba, number and percentage of
visits of different insects.
Insects 2015 2016 2015/2016
Order Family Genus, species n1 p1 (%) n2 p2 (%) nT pT (%)
Hymenoptera Apidae Apis mellifera (ne) 195 8.07 189 7.96 384 8.19
Amegilla acraensis (ne) 125 5.17 125 5.27 250 5.33
Amegilla sp. 1 (ne) 115 4.76 126 5.31 241 5.14
Amegilla sp. 2 (ne) 68 2.81 71 2.99 139 2.96
Xylocopa inconstans (ne) 91 3.76 93 3.92 184 3.92
Xylocopa olivacea (ne) 276 11.42 292 12.31 568 12.11
Euapsis abdominalis (ne) 91 3.76 96 4.05 187 3.99
Megachilidae Chalicodoma rufipes (ne) 259 10.72 273 11.50 532 11.34
Chalicodoma cinta (ne) 371 15.35 383 16.14 754 16.08
Megachile sp. 1 (ne) 189 7.82 196 8.26 385 8.21
Megachile sp. 2 (ne) 59 2.44 56 2.36 115 2,45
Vespidae Belonogaster juncea (ne) 58 2.40 58 2.44 116 2.47
Coelioxys sp (ne) 63 2.61 - - 63 1.34
Formicidae Camponotus flavomarginatus (ne) 200 8.27 225 9.48 425 9.06
Myrmicaria sp. (ne) 182 7.53 190 8.01 372 7.93
Lepidoptera Pieridae Eurema sp. (ne) 75 3.10 - - 75 1.60
Total 16 species 2317 100,00 2373 100 4584 100
n1: number of visits on 1136 flowers in 23 days; n2: number of visits on 1146 flowers in 23 days; p1 and p2: percentages of visits;
p1 = (n1/2317)*100; p2 = (n2/2373)*100; nT: total number of visit; pT (%):total percentages; sp.: undetermined species; ne:
visitor collected nectar.
Comparison of percentages of Chalicodoma cincta visits for two years: χ2 = 0.01 (df = 1; P > 0.05).
Foraging speed of Chalicodoma cincta in Cajanus
cajan field: In Ca. cajan field, Ch. cincta visited
between 3 and 25 flowers per minute in 2015 and
between 2 and 26 flowers per minute in 2016. The
mean foraging speed was 12.38 flowers/min (n = 316;
s = 0.55) in 2015 and 11.09 flowers per minute (n =
260; s = 1.12) in 2016.
The difference between these two means is highly
significant (t = 157.36; df = 574; P < 0.001).
Table 2. Fruiting rate, number of seeds per pod and percentage of normal seeds according to different
treatments of Cajanus cajan in 2015 and 2016 at Doyaba.
Years T NF NFP FrR (%) Seeds/pod TNS NS % NS
m sd
2015 1 (Uf) 1136 1048 92.25 5.88 0.38 635 598 94.17
2 (Bf) 992 725 73.08 4.90 0.70 500 327 65.40
5 Bfvch) 1052 971 92.30 5.87 0.42 587 531 90.45
6 (Biwv) 924 670 72.54 4.89 0.30 465 282 60.64
2016 3 (Uf) 1146 967 84.38 5.64 0.52 609 572 93.92
4 (Bf) 1036 792 76.44 4.91 0.53 496 321 64.71
7(Bfvch) 893 739 82.75 5,68 0.68 568 515 90.66
8 (Bfwv) 1047 801 76.50 4.83 0.40 464 268 57.75
NF: Number of flowers; NFP: Number of formed pod; FrR: Fruiting rate; TNS: Total number of seeds; NS: Normal seeds; %NS:
Percentage of normal seeds; m: mean; sd: standard deviation; T: traitments Uf: unprotected flowers; Bf: flowers bagged;
Bfvch: flowers bagged and exclusively visited by Chalicodoma cincta; Bfwv: flower bagged and not visit by insects or any other
organism.
J. Bio. Env. Sci. 2019
20 | Ningatoloum et al.
Daily rhythm of visits: Chalicodoma cincta were
active on the flowers of C. cajan from 7 am to 16 pm,
with a peak of visits between 9 and 10 am in 2015 as
well as in 2016 (Fig. 3). Ambiant temperature and
humidity conditions have not influenced the activity
of C. cincta on C. cajan flowers in the field conditions.
Fig. 1. Plant of Cajanus cajan showing a flower
protected from insects.
The correlation was not significant between the
number of C. cincta visits on Ca. cajan flowers and
the temperature in 2015 (r = -0.82; df = 3; P ˃ 0.05)
and in 2016 (r = -0.86; df = 3; P ˃ 0.05). The
correlation between the number of visits and the
humidity was not significant in 2015 (r = 0.50; df = 3;
P ˃ 0.05) as well as in 2016 (r = 0.67; df = 3; P ˃ 0.05)
(Fig. 3).
Impact of flower-feeding insects on the pollination
and yields of Cajanus cajan
During nectar harvest, flowering insects of Ca. cajan
were in regular contact (100% respectively in 2015
and 2016) with the anthers and stigma. Thus, these
insects increased the pollination possibilities of this
plant species. Table 2 present the results on the
fruiting rate, the number of seeds per pod and the
percentage of normal seeds in different treatments.
From this table, we documented the following:
(a) The comparison of the fruiting rate showed that
the difference was highly significant between
treatment 1 (unprotected flowers) and treatment 2
(bagged flowers) in the first year (χ2 = 11.83; df = 1;
P< 0.001) and significant between treatment 3
(unprotected flowers) and treatment 4 (bagged
flowers) in the second year (χ 2 = 4 .68; df = 1; P <
0.05). Consequently, in 2015 and 2016, the fruiting
rate of unprotected flowers was higher than that of
bagged flowers;
Fig. 2. Chalicodoma cincta collecting nectar in a
flower of Cajanus cajan.
(b) The comparison of the mean number of seeds per
pod showed that the difference was highly
significant between treatments 1 and 2 (t = 785.23; df
= 1771; P < 0.001) in 2015 and between treatment
3 and 4 (t = 605.61; df = 1757; P < 0.001) in 2016.
Consequently, in 2015 as in 2016, the mean number
of seeds per pod of the unprotected flowers was
higher than that of protected flowers;
(c) The comparison of the percentage of normal seeds
showed that the difference was highly significant
between treatments 1 and 2 (χ2 = 1 5 3 .59; df = 1; P <
0.001) as well between treatments 3 and bagged (χ2 =
150.40; df = 1; P < 0.001). Thus, in 2015 as in 2016,
the percentage of normal seeds produced by
unprotected flowers was higher than that produced
by of protected flowers.
The numeric contribution of flowering insects to the
fruiting rate, the mean number of seeds per pod and
the percentage of normal seeds were respectively
J. Bio. Env. Sci. 2019
21 | Ningatoloum et al.
20.78%, 16.66% and 30.55% in 2015. The
corresponding figures were 9.40%, 12.94% and
31.10% in 2016. For the two cumulate years, the
numeric contributions of flowering insects were
15.09%, 14.37% and 30.82% in the fruiting rate, the
mean number of seeds per pod and the percentage of
normal seeds respectively.
Fig. 3. Mean daily temperature and humidity and number of Chalicodoma cincta visits on Cajanus cajan flowers
in 2015 (A) and 2016 (B) at Doyaba.
Pollination efficiency of Chalicodoma cincta on
Cajanus cajan
From Table 2, it appears that:
(a) The comparison of the fruiting rate showed that
the difference were higher significant between
treatment 5 ( flowers protected and visited
exclusively by Ch. cincta) and treatment 6 (bagged
flowers to prevent insect visitors) (χ2 = 136.83; df =
1; P < 0.001) as well as between treatment 7
( flowers protected and visited exclusively by Ch.
cincta) and treatment 8 (bagged flowers to prevent
insect visitors) (χ2 = 11.50; df = 1; P < 0.001).
Consequently, in 2015 as in 2016, the fruiting rate of
flowers protected and visited exclusively by C. cincta
was higher than that of flowers bagged to prevent
insect visitors;
(b) The comparison of the mean number of seeds per
pod showed that the difference were highly significant
between treatments 5 and 6 (t = 1033.58; df = 1639;
P < 0.001) and between treatments 7 and 8 (t =
591.10; df = 1538; P < 0.001). Consequently, in 2015
J. Bio. Env. Sci. 2019
22 | Ningatoloum et al.
as in 2016, the pods produce by flowers bagged and
visited exclusively by C. cincta were higher than those
of flowers protected to prevent insect visitors;
(c) The comparison of the percentage of normal seeds
showed that the difference were highly significant
between treatment 5 and 6 (χ2 = 131.37; df = 1; P <
0.001) and between treatments 7 and 8 (χ2 = 151.10;
df = 1; P < 0.001) respectively. For the two years, the
pods produced by flowers bagged and visited
exclusively by C. cincta had more normal seeds than
those of flowers protected to prevent insect visitors.
The numeric contribution of Ch. cincta in the fruiting
rate, the mean of number of seed per pod and the
percentage of normal seeds were respectively 21.40%,
16.69% and 32.95% in 2015. The corresponding results
were 7.55%, 14.96% and 36.30% in 2016. For the two
cumulated years, the numeric contributions of Ch.
cincta in the fruiting rate, the mean number of seeds
per pods and the percentage of normal seeds of Ca.
cajan were 14.47%, 15.82% and 34.62% respectively.
Discussion
Reproduction mode of Cajanus cajan
Our results show that C. cajan has an allogamy-
autogamy reproduction mode with the predominance
of autogamy. The same result was reported by Pando
et al. (2011b) in Yaoundé and Mazi et al. (2014) in
Ngaoundéré.
Activity of Chalicodoma cinta on Cajanus cajan
flowers
Results obtained from our experiments indicated that
C. cincta was the main floral insect visitor of Ca.
cajan. The same result was reported by Pando et al.
(2011b) for the same bee on the same plant species
in Yaoundé. The significant difference between the
percentages of C. cincta visit for the two years of
study could be explained by the variation of the
number of C. cincta nests (17 in 2015 and 28 in 2016)
and the needs of this bee for one year to another. The
activity of C. cincta on the flowers of C. cajan was
highest in the morning, which correspondent to the
daily moment of highest availability of nectar on Ca.
cajan flowers. The same result has been reported
by Saxena and Kumar (2010) in India and Pando et
al. (2011b) in Cameroon. The high abundance of C.
cincta per 1000 flowers indicated a good
attractiveness of C. cajan nectar vis-à-vis of C. cincta.
The positive and significant contribution of C. cincta
in the pod and seeds yields of C. cajan is justified by
the action of this leat cutter bee on pollination.
Pollination efficiency of Chalicodoma cincta on
Cajanus cajan
During the collection of nectar on each flower, C.
cincta regularly come into contact with the stigma.
They were also able to carry pollen with their
hairs, a b d o m e n , legs and mouth accessories from
a flower of one plant to stigma of another flower of
the same plant (geitonogamy), to the same flower
(autogamy) or to that of another plant (xenogamy)
(Saxena and Kumar, 2010). The significant
contribution of C. Cincta on pod and seed production
of C. cajan is in agreement with the similar findings
in Yaoundé (Pando et al., 2011b) on the same
Fabaceae. The contribution of C. cincta to C. cajan
production through its pollination efficiency was
significantly higher than that of all insects on the
exposed flowers. The weight of C. cincta played a
positive role. During nectar collection, C. cincta
shook flowers and could facilitate the liberation of
pollen by anthers for the optimal occupation of the
stigma. This phenomenon was also reported by Pando
et al. (2011b) in Yaoundé (Cameroon) on this same
plant species.
The higher production of seeds per pod and that of
normal seeds in the treatment with flowers visited
exclusively by C. cincta compared to that of the
treatment with protected flowers showed that C. cincta
visit was effective in increasing pollination. Our
results confirmed those of Pando et al., 2011 and Mazi
(2015) who revealed that C. cajan flowers set
little pods in the absence of insect pollinators.
J. Bio. Env. Sci. 2019
23 | Ningatoloum et al.
Conclusion
From our observations, the variety of Cajanus cajan
studied is a plant species that highly benefits from
pollination by insects among which C. cincta is of
great importance. The comparison of pod and seed set
of unprotected flowers to those of flowers visited
exclusively by C. Cincta underscores the value of this
bee in increasing podding rate, the number of seeds
per pod and the percentage of normal seeds of Ca.
cajan. The installation and/or the kept of Ch. cincta
nest at the vicinity of pigeon pea fields is
recommended to Chadian farmers to increase pot and
seed productions.
Acknowledgements
We thank the staff of Sarh University, for granting us
the experimental plot.
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Pollination efficiency of Chalicodoma cincta (Fabricius) (Hymenoptera: Megachilidae) on Cajanus cajan (L.) Millsp. (Fabaceae) flowers at Doyaba (Sarh, Chad) | JBES

  • 1. J. Bio. Env. Sci. 2019 15 | Ningatoloum et al. RESEARCH PAPER OPEN ACCESS Pollination efficiency of Chalicodoma cincta (Fabricius) (Hymenoptera: Megachilidae) on Cajanus cajan (L.) Millsp. (Fabaceae) flowers at Doyaba (Sarh, Chad) Clautin Ningatoloum*1,2 , Guiguindibaye Madjimbe3 , Sidonie Fameni Tope4 , Fernand-Nestor Tchuenguem Fohouo1 1 Laboratory of Applied Zoology, Department of Biological Sciences, Faculty of Science, University of Ngaoundéré, Ngaoundéré, Cameroon 2 Department of Biological Sciences, University Adam Barka of Abéché, Abéché, Chad 3Department of Environmental Sciences, University Institute of Sciences Agronomic and the Environment, Sarh, Chad 4 Laboratory of Zoology, Department of Biological Sciences, Faculty of Science, University of Maroua, Maroua, Cameroon Article published January 26, 2019 Key words: Cajanus cajan, Chalicodoma cincta, Flowers, Yield, Pollination. Abstract To evaluate Chalicodoma cincta impact on pod and seed yields of Cajanus cajan, its foraging and pollinating activities were studied in Doyaba, during the rainy season of 2015 and 2016. Each year, treatments included flowers accessible to all visitors, bagged flowers to avoid insect visitors, bagged flowers using gauze bags destined to be visited exclusively by C. cincta and bagged flowers destined to opening and closing without the visit of insects or any other organism. For each year of study, observations were made on 1028 ± 90 flowers per treatment. Chalicodoma cincta daily rhythm of activity, its foraging behaviour on flowers and its pollination efficiency were evaluated. On flowers, individual bees intensely harvested exclusively nectar. The fruiting rate, the number of seeds per pod and the percentage of normal seeds of unprotected flowers were significantly higher than those of flowers protected from insects. Through its pollination efficiency, C. cincta provoked a significant increment of the fruiting rate by 21.40% and 7.55%, the number of seeds per pod by 16.69% and 14.96% and the percentage of normal seeds by 32.95% in 2015 and 36.30% in 2016 respectively. The Conservation of C. cincta nests close to C. cajan fields is recommended to improve pod and seed productions in the region. *Corresponding Author: Clautin Ningatoloum  gclautin@gmail.com Journal of Biodiversity and Environmental Sciences (JBES) ISSN: 2220-6663 (Print) 2222-3045 (Online) Vol. 14, No. 1, p. 15-25, 2019 http://www.innspub.net
  • 2. J. Bio. Env. Sci. 2019 16 | Ningatoloum et al. Introduction Several plant species depend on insect pollinators for their reproduction; in agro ecosystems, these pollinators have a great ecological and economic importance because they influence positively the plant production (McGregor, 1976; Philippe, 1991; Tchuenguem, 2005). Cajanus cajan is an ideal pulse crop of rainfed tropics and subtropics (Saxena et al., 2002). Its grows up right to 4 m (Niyonkuru, 2002). The leaves are generally trifoliate; flower is pink, but can vary from white to red (ICRISAT, 1981) and produces nectar and pollen which attract insects (Grewal et al., 1990; Saxena et al., 1990; Reddy et al., 2004; Sarah et al., 2010). Indian is the largest producers of pigeon pea in the world (Kimani, 2000). The fruit is a pod containing four raw of seven seeds (Pando et al., 2011b). Seeds contain 21 to 30% proteins important for human’s diets (Sharma and Green, 1980; Gupta et al., 2001; Saxena et al., 2002). Cajanus cajan flowers were reported to produce fewer seeds per pod in the absence of insect pollinators in Great Britain (Kendall and Smith, 1976) and in Cameroon (Pando et al., 2011b). The research conducted in the United Stated of America (Grewal et al., 1990; Ibarra-Perez et al., 1999) and in Cameroon (Pando, 2012; Mazi, 2015) has revealed that bees of the genus Chalicodoma and Megachile visits C. cajan flowers and collect nectar and pollen. In Chad, the quantity of Ca. cajan available to consumers is very low, the demand for pigeon pea seeds is high, and its pod and seed yields are weak because notably of the insufficiency of knowledge on its relationships with anthophilous insects in general and C. cincta in particular. Therefore, it is important to investigate how the production of this plant could be increased in Chad. Prior to these studies, no previous research has been reported on the interactions between Ca. cajan and insects in Chad. The main objective of this work is to contribute to the understanding of the relationships between C. cajan and C. cincta, for their optimal management in this country. Specific objectives are to: (a) study the activity of this Megachilidae on C. cajan flowers, (b) evaluate the impact of flowering insects including Ch. cincta on pollination and fruit and seed yields of this Fabaceae, (c) estimate the pollination efficiency of C. cincta on this plant species. Materials and methods Site and biological materials The studies were conducted twice, June to November in 2015 and 2016 in a field located at Doyaba, Moyen- Chari Region of Chad. This Region belongs to the Sudanian agro-ecological zone (Cabot and Bouquet, 1972). The climate is tropical type with two seasons: a dry season (November to April) and a rainy season (May to October). The mean annual temperature is 28°C; the mean rainfall is 1100 mm (Madjimbé, 2013). The experimental plot was an area of 437 m2 (Latitude: 09° 04.875’ N, Longitude: 018° 25.721’ E, Altitude 363.3 m.a.s.l.). The animal material was represented by insects naturally present in the environment. The vegetation was represented by wild and cultivated species. The plant material was represented by the seeds of C. cajan variety ICPL 9145, provided by the Agronomic Institute of the University of Sarh. Sowing and weeding On June 14th, 2015 and 2016 the experimental plot was divided into eight subplots (8*4.5 m2). Seeds were sown on 16 lines per subplot; each line had nine holes and each hole received three seeds. The spacing was 50 cm between rows and 50 cm on rows. Each hole was 5 cm depth. Two weeks after germination, the plants were thinned and only two were left per hole (Pando et al., 2011b). Weeding was performed manually as necessary to maintain subplots weeds-free (Tchuenguem et al, 2009a; Pando et al., 2013).
  • 3. J. Bio. Env. Sci. 2019 17 | Ningatoloum et al. Determining the reproduction mode In October 12th, 2015, 2282 flowers of Ca. cajan at the bud stage were labeled, among which 1136 were allowed for treatment 1 (open pollination) and 1146 for treatment 2 (bagged with gauze bags to prevent visitors) (Tchuenguem, 2005; Delaplane et al., 2013) (Fig. 1). In October 11th, 2016, 2039 flowers at the bud stage were labeled, among which 992 were provided for treatment 3 (open pollination) and 1047 flowers for treatment 4 (bagged with gauze bags). For each year, ten days after the wilting of the last flower, the number of pods formed in each treatment was counted. For each treatment, the fruiting index (Ifr) was then calculated using the following formula: Ifr = (F1/F2), where F1 is the number of pods formed and F2 the number of flowers initially labeled (Tchuenguem et al., 2001). For each year, the out crossing rate (TC) was calculated using the formula: TC = {[(Ifra – Ifrb /fra]*100}, where Ifra and Ifrb are the mean fruiting indexes in treatment a (flowers in open pollination) and treatment b (bagged flowers) respectively (Demarly, 1977). The rate of self- pollination in the broad sense (TA) was calculated using the formula: TA = (100 - TC) (Tchuenguem, 2005; Népidé and Tchuenguem, 2016). Study of the activity of Chalicodoma cincta on Cajanus cajan flowers Observations were conducted on individually opened flowers of treatments 1 and 3, each day, from October 13th to November 03th 2015 and from October 12th to November 04th 2016, at one hour interval, from 6 am to 15 pm (6-7 am, 8-9 am, 10-11 am, 12-13 pm and 14- 15 pm). In a slow walk along all labeled flowers of treatments 1 and 3, the identity of each insect that visited Ca. cajan flowers was noted. All insects encountered on flowers were recorded and the cumulated results expressed in number of visits to determine the relative frequency of C. cincta in the anthophilous entomofauna of C. cajan (Pando et al., 2011b). Direct observations of the foraging activity of this bee on flowers were made. The floral products (nectar and/or pollen) collected by Ch. cincta were recorded during the same dates and time slots as that of the insect counts (Pando et al., 2011b). The duration of visits per flower was recorded (using stopwatch) according to five time frames (7-8 am, 9-10 am, 11-12 am, 13-14 pm and 15-16 pm). The foraging speed (number of flowers visited by individual bee per minute (Jacob-Remacle, 1989) was recorded during the same date and daily periods as the registration of the duration of visits. Abundances (highest number of individuals simultaneously active) per flower and per 1000 flowers (A1000) were recorded during the same dates and time slots as that of the registration of the duration of visits per flower. The first parameter was recorded as a result of direct counts. For the abundance per 1000 flowers, bees were counted on a known number of open flowers at the moment t. The abundance per 1000 flowers is calculated using formula: A1000 = [(Aa/Fb)*1000], where Fb and Ab are respectively the number of flowers and the number of C. cincta effectively counted on these flowers at time t (Tchuenguem et al., 2004; Tchuenguem, 2005; Douka et al., 2017). The disruption of the activity of individual bee by competitors or predators and the attractiveness of other plant species with respect to C. cincta were assessed. During the days of investigation, the temperature and humidity of the study site were recorded every 30 min, from 6 am to 17 pm, using a thermo hygrometer installed in the shade (Tchuenguem et al., 2004; Farda and Tchuenguem, 2018). Evaluation of the impact of flowering insects including Chalicodama cincta on Cajanus cajan yield For each investigation year, this evaluation was based on the impact of flowering insects on pollination, the impact of pollination on fruiting and the comparison of yields (fruiting rate, mean number of seeds per pod and percentage of normal i.e. well-developed seeds) of treatment a (unprotected flowers) and treatment b (bagged flowers). The fruiting rate due to the influence of foraging insects (Fri) was calculated using the formula: Fri = {[(Fra-Frb)/Fra]*100}, where Fra and Frb are the
  • 4. J. Bio. Env. Sci. 2019 18 | Ningatoloum et al. fruiting rate in treatments a and b respectively. The fruiting rate of a treatment (Fr) is Fr = [(F2/F1)*100], where, F2 is the number of pods formed and F1 the number of viable flowers initially set (Tchuenguem et al., 2001). At maturity, pods were harvested from all treatments. The mean number of seeds per pod and the percentage of normal seeds were then calculated for each treatment. Assessment of the pollination efficiency of Chalicodoma cincta on Cajanus cajan In 2015, along with the development of treatments 1 and 2, 1976 flowers at bud stage were protect and two treatments was formed: treatments 5, with 1052 flowers protected using gauze bags to prevent insect visitors and destined exclusively to be visited by Ch. cincta; treatment 6, with 924 flowers protected from insects using gauze bags. In 2016, along with the realization of treatments 3 and 4, 1940 flowers at the bud stage were labeled and two treatments was formed: treatment 7 with, 893 flowers protected from insects using gauze bags and destined exclusively to be visited by C. cincta; treatment 8 with 1047 flowers protected from insects using gauze bags. As soon as the first flowers were opened, in treatments 5 and 7, the gauze bag was delicately removed and flowers were observed for up to 10 min. Flowers visited by C. cincta was marked and then protected once more. For treatment 6 and 8, as soon as the firsts flower was opened, the gauze bag was delicately removed and flowers were observed for up to 10 min while avoiding the visits by any other organism. At the maturity, pods were harvested and the contribution of Ch. cincta in fruiting, number of seeds per pod and percentage of normal seeds was then calculated using data of treatments 3 or 8 (flowers visited exclusively by C. cincta) and those of treatments 2 or 7 (bagged flowers). Data analysis Data were subjected to descriptive statistics, student’s t-test for the comparison of means of the two samples, ANOVA for the comparison of means of more than two samples, correlation coefficient (r) for the study of the association between two variables, and chi-square (χ2) for the comparison of percentages, using Microsoft Excel 2010 software. Results Reproduction mode of Cajanus cajan The podding indexes were 0.92, 0.73, 0.84 and 0.74, in treatments 1, 2, 3 and 4 respectively. Consequently, in 2015, the allogamy rate was 20.85% and the autogamy rate was 79.15%; for 2016, the corresponding data were 11.90% and 88.10%. It appears that the Ca. cajan variety used in our experiment has a mix reproduction mode, autogamous-allogamous with the predominance of autogamy over allogamy. Activity of Chalicodoma cinta on Cajanus cajan flowers Seasonal frequency of visits: Among the 2317 and 2373 visits of 16 and 14 insect species recorded on C. cajan flowers in 2015 and 2016 respectively, C. cincta was the most represented insect with 371 visits (15.35%) in 2015 and 383 visits (16.14%) in 2016 (Table 1). The difference between these two percentages is not significant (χ2 = 0.87; df = 1; P > 0.05). Flower substances harvested: From our field observations of the years 2015 and 2016 C. cincta were found to collect regularly and exclusively nectar in the flowers of C. cajan (Fig. 2). Abundance of Chalicodoma cincta: In 2015, the highest mean number of Ch. cincta simultaneous in activity was one per flower (n = 230; s = 0) and 188.08 per 1000 flowers (n = 230; s = 154.84). In 2016, the corresponding results were one per flower (n = 198; s = 0) and 189.65 (n = 198; s = 198.65) per 1000 flowers.
  • 5. J. Bio. Env. Sci. 2019 19 | Ningatoloum et al. The difference between the mean number of C. cincta per 1000 flowers of the two years was not significant (t = 1.08; df = 426; P > 0.05). Duration of visits per flower: In 2015, the mean duration of individual Ch. cincta visit per Ca. cajan flower was 9.00 sec (n = 1168; s = 8.9). The corresponding index was 8.36 sec (n = 704; s = 8.27) in 2016. The difference between these two means is higher significant (t = 20.56; df = 292; P < 0.001). Table 1. Diversity of floral insects on Cajanus cajan in 2015 and 2016 at Doyaba, number and percentage of visits of different insects. Insects 2015 2016 2015/2016 Order Family Genus, species n1 p1 (%) n2 p2 (%) nT pT (%) Hymenoptera Apidae Apis mellifera (ne) 195 8.07 189 7.96 384 8.19 Amegilla acraensis (ne) 125 5.17 125 5.27 250 5.33 Amegilla sp. 1 (ne) 115 4.76 126 5.31 241 5.14 Amegilla sp. 2 (ne) 68 2.81 71 2.99 139 2.96 Xylocopa inconstans (ne) 91 3.76 93 3.92 184 3.92 Xylocopa olivacea (ne) 276 11.42 292 12.31 568 12.11 Euapsis abdominalis (ne) 91 3.76 96 4.05 187 3.99 Megachilidae Chalicodoma rufipes (ne) 259 10.72 273 11.50 532 11.34 Chalicodoma cinta (ne) 371 15.35 383 16.14 754 16.08 Megachile sp. 1 (ne) 189 7.82 196 8.26 385 8.21 Megachile sp. 2 (ne) 59 2.44 56 2.36 115 2,45 Vespidae Belonogaster juncea (ne) 58 2.40 58 2.44 116 2.47 Coelioxys sp (ne) 63 2.61 - - 63 1.34 Formicidae Camponotus flavomarginatus (ne) 200 8.27 225 9.48 425 9.06 Myrmicaria sp. (ne) 182 7.53 190 8.01 372 7.93 Lepidoptera Pieridae Eurema sp. (ne) 75 3.10 - - 75 1.60 Total 16 species 2317 100,00 2373 100 4584 100 n1: number of visits on 1136 flowers in 23 days; n2: number of visits on 1146 flowers in 23 days; p1 and p2: percentages of visits; p1 = (n1/2317)*100; p2 = (n2/2373)*100; nT: total number of visit; pT (%):total percentages; sp.: undetermined species; ne: visitor collected nectar. Comparison of percentages of Chalicodoma cincta visits for two years: χ2 = 0.01 (df = 1; P > 0.05). Foraging speed of Chalicodoma cincta in Cajanus cajan field: In Ca. cajan field, Ch. cincta visited between 3 and 25 flowers per minute in 2015 and between 2 and 26 flowers per minute in 2016. The mean foraging speed was 12.38 flowers/min (n = 316; s = 0.55) in 2015 and 11.09 flowers per minute (n = 260; s = 1.12) in 2016. The difference between these two means is highly significant (t = 157.36; df = 574; P < 0.001). Table 2. Fruiting rate, number of seeds per pod and percentage of normal seeds according to different treatments of Cajanus cajan in 2015 and 2016 at Doyaba. Years T NF NFP FrR (%) Seeds/pod TNS NS % NS m sd 2015 1 (Uf) 1136 1048 92.25 5.88 0.38 635 598 94.17 2 (Bf) 992 725 73.08 4.90 0.70 500 327 65.40 5 Bfvch) 1052 971 92.30 5.87 0.42 587 531 90.45 6 (Biwv) 924 670 72.54 4.89 0.30 465 282 60.64 2016 3 (Uf) 1146 967 84.38 5.64 0.52 609 572 93.92 4 (Bf) 1036 792 76.44 4.91 0.53 496 321 64.71 7(Bfvch) 893 739 82.75 5,68 0.68 568 515 90.66 8 (Bfwv) 1047 801 76.50 4.83 0.40 464 268 57.75 NF: Number of flowers; NFP: Number of formed pod; FrR: Fruiting rate; TNS: Total number of seeds; NS: Normal seeds; %NS: Percentage of normal seeds; m: mean; sd: standard deviation; T: traitments Uf: unprotected flowers; Bf: flowers bagged; Bfvch: flowers bagged and exclusively visited by Chalicodoma cincta; Bfwv: flower bagged and not visit by insects or any other organism.
  • 6. J. Bio. Env. Sci. 2019 20 | Ningatoloum et al. Daily rhythm of visits: Chalicodoma cincta were active on the flowers of C. cajan from 7 am to 16 pm, with a peak of visits between 9 and 10 am in 2015 as well as in 2016 (Fig. 3). Ambiant temperature and humidity conditions have not influenced the activity of C. cincta on C. cajan flowers in the field conditions. Fig. 1. Plant of Cajanus cajan showing a flower protected from insects. The correlation was not significant between the number of C. cincta visits on Ca. cajan flowers and the temperature in 2015 (r = -0.82; df = 3; P ˃ 0.05) and in 2016 (r = -0.86; df = 3; P ˃ 0.05). The correlation between the number of visits and the humidity was not significant in 2015 (r = 0.50; df = 3; P ˃ 0.05) as well as in 2016 (r = 0.67; df = 3; P ˃ 0.05) (Fig. 3). Impact of flower-feeding insects on the pollination and yields of Cajanus cajan During nectar harvest, flowering insects of Ca. cajan were in regular contact (100% respectively in 2015 and 2016) with the anthers and stigma. Thus, these insects increased the pollination possibilities of this plant species. Table 2 present the results on the fruiting rate, the number of seeds per pod and the percentage of normal seeds in different treatments. From this table, we documented the following: (a) The comparison of the fruiting rate showed that the difference was highly significant between treatment 1 (unprotected flowers) and treatment 2 (bagged flowers) in the first year (χ2 = 11.83; df = 1; P< 0.001) and significant between treatment 3 (unprotected flowers) and treatment 4 (bagged flowers) in the second year (χ 2 = 4 .68; df = 1; P < 0.05). Consequently, in 2015 and 2016, the fruiting rate of unprotected flowers was higher than that of bagged flowers; Fig. 2. Chalicodoma cincta collecting nectar in a flower of Cajanus cajan. (b) The comparison of the mean number of seeds per pod showed that the difference was highly significant between treatments 1 and 2 (t = 785.23; df = 1771; P < 0.001) in 2015 and between treatment 3 and 4 (t = 605.61; df = 1757; P < 0.001) in 2016. Consequently, in 2015 as in 2016, the mean number of seeds per pod of the unprotected flowers was higher than that of protected flowers; (c) The comparison of the percentage of normal seeds showed that the difference was highly significant between treatments 1 and 2 (χ2 = 1 5 3 .59; df = 1; P < 0.001) as well between treatments 3 and bagged (χ2 = 150.40; df = 1; P < 0.001). Thus, in 2015 as in 2016, the percentage of normal seeds produced by unprotected flowers was higher than that produced by of protected flowers. The numeric contribution of flowering insects to the fruiting rate, the mean number of seeds per pod and the percentage of normal seeds were respectively
  • 7. J. Bio. Env. Sci. 2019 21 | Ningatoloum et al. 20.78%, 16.66% and 30.55% in 2015. The corresponding figures were 9.40%, 12.94% and 31.10% in 2016. For the two cumulate years, the numeric contributions of flowering insects were 15.09%, 14.37% and 30.82% in the fruiting rate, the mean number of seeds per pod and the percentage of normal seeds respectively. Fig. 3. Mean daily temperature and humidity and number of Chalicodoma cincta visits on Cajanus cajan flowers in 2015 (A) and 2016 (B) at Doyaba. Pollination efficiency of Chalicodoma cincta on Cajanus cajan From Table 2, it appears that: (a) The comparison of the fruiting rate showed that the difference were higher significant between treatment 5 ( flowers protected and visited exclusively by Ch. cincta) and treatment 6 (bagged flowers to prevent insect visitors) (χ2 = 136.83; df = 1; P < 0.001) as well as between treatment 7 ( flowers protected and visited exclusively by Ch. cincta) and treatment 8 (bagged flowers to prevent insect visitors) (χ2 = 11.50; df = 1; P < 0.001). Consequently, in 2015 as in 2016, the fruiting rate of flowers protected and visited exclusively by C. cincta was higher than that of flowers bagged to prevent insect visitors; (b) The comparison of the mean number of seeds per pod showed that the difference were highly significant between treatments 5 and 6 (t = 1033.58; df = 1639; P < 0.001) and between treatments 7 and 8 (t = 591.10; df = 1538; P < 0.001). Consequently, in 2015
  • 8. J. Bio. Env. Sci. 2019 22 | Ningatoloum et al. as in 2016, the pods produce by flowers bagged and visited exclusively by C. cincta were higher than those of flowers protected to prevent insect visitors; (c) The comparison of the percentage of normal seeds showed that the difference were highly significant between treatment 5 and 6 (χ2 = 131.37; df = 1; P < 0.001) and between treatments 7 and 8 (χ2 = 151.10; df = 1; P < 0.001) respectively. For the two years, the pods produced by flowers bagged and visited exclusively by C. cincta had more normal seeds than those of flowers protected to prevent insect visitors. The numeric contribution of Ch. cincta in the fruiting rate, the mean of number of seed per pod and the percentage of normal seeds were respectively 21.40%, 16.69% and 32.95% in 2015. The corresponding results were 7.55%, 14.96% and 36.30% in 2016. For the two cumulated years, the numeric contributions of Ch. cincta in the fruiting rate, the mean number of seeds per pods and the percentage of normal seeds of Ca. cajan were 14.47%, 15.82% and 34.62% respectively. Discussion Reproduction mode of Cajanus cajan Our results show that C. cajan has an allogamy- autogamy reproduction mode with the predominance of autogamy. The same result was reported by Pando et al. (2011b) in Yaoundé and Mazi et al. (2014) in Ngaoundéré. Activity of Chalicodoma cinta on Cajanus cajan flowers Results obtained from our experiments indicated that C. cincta was the main floral insect visitor of Ca. cajan. The same result was reported by Pando et al. (2011b) for the same bee on the same plant species in Yaoundé. The significant difference between the percentages of C. cincta visit for the two years of study could be explained by the variation of the number of C. cincta nests (17 in 2015 and 28 in 2016) and the needs of this bee for one year to another. The activity of C. cincta on the flowers of C. cajan was highest in the morning, which correspondent to the daily moment of highest availability of nectar on Ca. cajan flowers. The same result has been reported by Saxena and Kumar (2010) in India and Pando et al. (2011b) in Cameroon. The high abundance of C. cincta per 1000 flowers indicated a good attractiveness of C. cajan nectar vis-à-vis of C. cincta. The positive and significant contribution of C. cincta in the pod and seeds yields of C. cajan is justified by the action of this leat cutter bee on pollination. Pollination efficiency of Chalicodoma cincta on Cajanus cajan During the collection of nectar on each flower, C. cincta regularly come into contact with the stigma. They were also able to carry pollen with their hairs, a b d o m e n , legs and mouth accessories from a flower of one plant to stigma of another flower of the same plant (geitonogamy), to the same flower (autogamy) or to that of another plant (xenogamy) (Saxena and Kumar, 2010). The significant contribution of C. Cincta on pod and seed production of C. cajan is in agreement with the similar findings in Yaoundé (Pando et al., 2011b) on the same Fabaceae. The contribution of C. cincta to C. cajan production through its pollination efficiency was significantly higher than that of all insects on the exposed flowers. The weight of C. cincta played a positive role. During nectar collection, C. cincta shook flowers and could facilitate the liberation of pollen by anthers for the optimal occupation of the stigma. This phenomenon was also reported by Pando et al. (2011b) in Yaoundé (Cameroon) on this same plant species. The higher production of seeds per pod and that of normal seeds in the treatment with flowers visited exclusively by C. cincta compared to that of the treatment with protected flowers showed that C. cincta visit was effective in increasing pollination. Our results confirmed those of Pando et al., 2011 and Mazi (2015) who revealed that C. cajan flowers set little pods in the absence of insect pollinators.
  • 9. J. Bio. Env. Sci. 2019 23 | Ningatoloum et al. Conclusion From our observations, the variety of Cajanus cajan studied is a plant species that highly benefits from pollination by insects among which C. cincta is of great importance. The comparison of pod and seed set of unprotected flowers to those of flowers visited exclusively by C. Cincta underscores the value of this bee in increasing podding rate, the number of seeds per pod and the percentage of normal seeds of Ca. cajan. The installation and/or the kept of Ch. cincta nest at the vicinity of pigeon pea fields is recommended to Chadian farmers to increase pot and seed productions. Acknowledgements We thank the staff of Sarh University, for granting us the experimental plot. References Cabot J, Bouquet C. 1972. Atlas pratique du Tchad, 34 cartes et notices 76, 20- 21. Delaplane KS, Dag A, Danka RG, Freitas BM, Garibaldi LA, Goodwin RM, Hormaza JI. 2013. Standard methods for pollination research with Apis mellifera. Journal of Apicultural Research 52, 1- 208. Demarly Y. 1977. Génétique et amélioration des plantes. Masson, Paris, 577. Douka C, Tamesse JL, Tchuenguem FFN. 2017. Impact of single visit of Lipotriches collaris Vachal 1903 (Hymenoptera: Halictidae) on Phaseolus vulgaris (Fabaceae) flowers at Maroua (Cameroon). Journal of Applied Biology & Biotechnology 5, 72- 76. Fard D, Tchuenguem ffn. 2018. Efficacité pollinisatrice de Apis mellifera (Hymenoptera: Apidae) sur Luffa cylindrica (L.) M. Roem (Cucurbitaceae) à Ngaoundéré (Cameroun). International Journal of Biological and Chemical Sciences 12, 850- 866. Grewal GS, Singh G, Kandoria JL. 1990. Insect pollinators of pigeon pea (Cajanus cajan) around Ludhiana. Indian Journal of Agricultural Science 60, 227- 228. Gupta DN, Parab AA, Mehta HR, Goyal R. 2001. Evaluation of nutritive value of promising genotypes and horsegram in Konkan. Journal of Phytological Research 14, 143- 146. Ibarra-Perez FJ, Barnhart D, Ehdaie B, Knio KM, Waines J G. 1999. Effects of insect tripping on seed yield of common beans. Crop Science 39, 428- 433. Jacob-Remacle A. 1989. Comportement de butinage de l'abeille domestique et des abeilles sauvages dans des vergers de pommiers en Belgique. Apidologie 20, 271- 285. Jean Prost P. 1987. Apiculture: Connaître l’abeille Conduire le rucher. 6ème edition. Lavoisier (Ed), Paris, 579. Kendal DA, Smith BD. 1976. The pollinating efficiency of honey bee and Bumblebee visits to flowers of runner bean Phaseolus coccineus. Journal of Applied Ecology 13, 749- 752. Kimani EW. 2000. Cercospora leafspot in eastern africa and strategies to reduce yield losses in Pigeonpea status and potential in eastern and southern africa, ICRISAT and Gembloux Agricultural University 61- 68. Klein AM, Vaissière, BE, Cane JH, Steffan- Dewenter I, Cunningham SA, Kremen C, Tscharntke T. 2007. Importance of pollinators in changing landscapes for world crops. Proceedings of the Royal Society, London (B) 274, 303- 313. Madjimbé G. 2013. Impacts biologiques et écologiques du feu contrôlé sur la végétation et le sol dans la forêt de Guirkouh, au sud du Tchad. Thèse de Doctorat Ph.D, Université de Ouagadougou, 16.
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