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Lixus algirus L. (Coleoptera: Curculionidae): biology, population fluctuation, infestation as affected by varieties, location, and planting dates in Morocco

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The stem borer, Lixus algirus L. (Coleoptera: Curculionidae) is the major insect pest of faba bean in the Mediterranean region. This study investigates the field biology and fluctuation of L. algirus at Douyet research Station in Morocco during two cropping seasons 2014-2015 and 2015-2016 using two local faba bean varieties. The influence of sowing dates and varieties on the damage caused by L. algirus was investigated over different locations and seasons using a split-plot design. The overwintering adults appear in the field from February by feeding over the leaves of faba bean plants. Eggs are laid from Mid-February through late April and mostly during March. The egg peak ranged from 0.75 to 1.1 eggs per plant. The number of larvae increases during March and peaked in April. The larval peaks ranged between 0.45 to 1.5 larvae per plant. The larval development takes an average of 42 days. Pupation lasts about 21 days within stems. Adults emerged in early June. Life cycle duration was 80 days. One generation of L. algirus was recorded on faba bean varieties in Morocco. The mean number of eggs showed a positive correlation with total rainfall. The number of pupae and relative humidity was negatively correlated. Peach and almond leaves were the most preferred by the newely emerged L. algirus adult during the summer and early autumn period. Planting dates did not affect L. algirus infestation levels over all varieties tested and locations.
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[Journal of Entomological and Acarological Research 2021; 53:9324] [page 7]
Abstract
The stem borer, Lixus algirus L. (Coleoptera: Curculionidae)
is the major insect pest of faba bean in the Mediterranean region.
This study investigates the field biology and fluctuation of L.
algirus at Douyet research Station in Morocco during two crop-
ping seasons 2014-2015 and 2015-2016 using two local faba bean
varieties. The influence of sowing dates and varieties on the dam-
age caused by L. algirus was investigated over different locations
and seasons using a split-plot design. The overwintering adults
appear in the field from February by feeding over the leaves of
faba bean plants. Eggs are laid from Mid-February through late
April and mostly during March. The egg peak ranged from 0.75 to
1.1 eggs per plant. The number of larvae increases during March
and peaked in April. The larval peaks ranged between 0.45 to 1.5
larvae per plant. The larval development takes an average of 42
days. Pupation lasts about 21 days within stems. Adults emerged
in early June. Life cycle duration was 80 days. One generation of
L. algirus was recorded on faba bean varieties in Morocco. The
mean number of eggs showed a positive correlation with total
rainfall. The number of pupae and relative humidity was negative-
ly correlated. Peach and almond leaves were the most preferred by
the newely emerged L. algirus adult during the summer and early
autumn period. Planting dates did not affect L. algirus infestation
levels over all varieties tested and locations.
Introduction
The faba bean (Vicia faba L.) is one of the most winter-sown
legume crops in the Mediterranean region (Manzoor, 2013). In
Morocco, this crop is considered among the most ancient crops
and the most important food legume crop. For instance, faba bean
plays a key role in the crop rotation system with cereals to
improve soil fertility. Moreover, the diversification of cropping
systems is leading to decreased disease, pests and weed build up
and potentially increased biodiversity, and providing food and
feed rich in protein (Amanuel et al., 2000; Crepona et al., 2010;
Maalouf et al., 2013).
Unfortunately, the average yields of faba beans in Morocco
remain 58% lower than the world average (FAO, 2016).
Nowadays, drought, Orobanche, Botrytis, stem nematodes, and
insect damage have severely affected the productivity and avail-
ability of this crop (Sakr, 1991; Yigezu et al., 2018).
In Morocco, V. faba is attacked by several insect pests, which
often cause extensive damage. Aphids Aphis fabae and A. craccivo-
ra, Sitona weevils, Sitona lineatus L., species of Bruchus and
Callosobruchus and Lixus algirus are the most frequent insects found
on faba bean at different plant phonological stages (Hoffmann, 1954;
Bardner, 1983; Sakr, 1991; Weigand & Bishara, 1991).
The stem borer weevil, L. algirus is an important insect pest
of faba bean in southern Europe, the Middle East, and the
Mediterranean region including North Africa (Diekmann, 1982;
Weigand & Bishara, 1991). The significant damage is mainly
caused by the larval stage causing the leaf yellowing, wilting, and
drying of the plants, affecting its growth and yield (Liotta, 1963;
Correspondence: Karim El Fakhouri, Entomology Laboratory,
International Center for Agricultural Research in the Dry Areas (ICAR-
DA), Rabat Institutes, Rabat, Morocco.
Tel.: +21.2677504124.
E-mail: K.El-Fakhouri@cgiar.org
Key words: Faba bean stem borer, Population dynamics, damage, cli-
matic factors, cultural practices, Morocco
Contributions: NA and KE contributed equally as first authors; the other
authors contributed equally.
Conflict of interest: No potential conflict of interest was reported by the
authors.
Funding: This research was partially funded by the project “India-
Morocco Food Legumes Initiative Morocco Component OCP
Foundation, ID: 100161”.
Received for publication: 28 August 2020.
Revision received: 26 January 2021
Accepted for publication: 31 January 2021.
©Copyright: the Author(s), 2021
Licensee PAGEPress, Italy
Journal of Entomological and Acarological Research 2021; 53:9324
doi:10.4081/jear.2021.9324
This article is distributed under the terms of the Creative Commons
Attribution Noncommercial License (by-nc 4.0) which permits any
noncommercial use, distribution, and reproduction in any medium,
provided the original author(s) and source are credited.
Journal of Entomological and Acarological Research 2012; volume 44:e Journal of Entomological and Acarological Research 2021; volume 53:9324
ENTOMOLOGY
Lixus algirus
L. (Coleoptera: Curculionidae): biology, population fluctuation,
infestation as affected by varieties, location, and planting dates in Morocco
N. Ait Taadaouit,1,2 K. El Fakhouri,1A. Sabraoui,1L. Rohi,2M. El Bouhssini1
1Entomology Laboratory, International Center for Agricultural Research in the Dry Areas (ICARDA), Rabat Institutes,
Rabat; 2Laboratory of Ecology and Environment, Faculty of Sciences Ben M'sik, University Hassan II of Casablanca,
Casablanca, Morocco
Non-commercial use only
[page 8] [Journal of Entomological and Acarological Research 2021; 53:9324]
Chakir, 1992; Khan et al., 2010 ). In Morocco, the stem borer is a
very destructive pest, with mean infestation rate of 75%
(Diekmann, 1982). Despite the economic importance of this insect
pest, there is little information available on its population dynamic
and the methods for its control.
Several cultural practices can be useful in reducing insect
infestations, which include sowing dates, seed rates, and fertiliza-
tion, clean seed, plowing, crop rotation tactic and intercropping to
produce the healthiest crop possible. Adjusting sowing dates is one
of the most important cultural practices that can help to escape pest
occurrence or avoid the most vulnerable stages. This safe and sus-
tainable tool has been incorporated successfully for many IPM
programs around the world for decades (Dara, 2019).
The objectives of this study were 1) to investigate the field
biology and the population fluctuations of the faba bean stem borer
and 2) to evaluate the effects of planting dates and varieties on the
damage caused by stem borer at different locations.
Materials and methods
Biology and dynamic study of faba bean stem borer
Experimental site
The experiments were conducted in two faba bean fields at
Douyet Experimental Station INRA-CRRA Meknes, Morocco,
located 12 km of Fez city (Saïs region), following the geographical
coordinates (33°03’59’’ N, 7°53’05” W) and (33°04’05.0”N
7°53’13.6”W) with an elevation of 416m.
Two local faba bean varieties Defes (V. faba major) and Alfia
(V. faba minor) were planted on December 10, 2015, and
December 15, 2016, following a Split Plot design with four repli-
cations. Each plot had five rows of 5-meter length and 0.6 m
between rows, with a seeding rate of 100 kg/ha. No insecticide was
applied during the crop-growing seasons.
Sampling insect populations
Population dynamics of L. algirus were conducted from early
February to Mid - June in 2015 and 2016. Sampling was performed
twice a week by tearing randomly 20 plants from each plot and
brought to the laboratory for evaluation. L. algirus adults were
manually checked on flowers, leaves, and other plant parts. Direct
observations of mating behaviour were made, with recording on
number of perforations per stems. In the laboratory, the faba bean
plants sampled were dissected carefully with scissors under a
binocular. The plants were checked for egg laying, adult exit hole
and, presence of larva or pupa inside the stem or its exudates. All
stages of the weevil were examined with an eyepiece micrometric
scale using the stereomicroscope Motic SMZ-140, length and
width values of each insect stage are given in millimeters.
The duration of developmental stages and sex ratio
During the mating period of L. algirus, each couple was placed
in Petri dishes of 15 cm in diameter with two parts of the faba bean
stems under laboratory conditions (25±1°C; 70-75% relative
humidity (RH); a photoperiod ranging from 10:14 h to 14:10
(L:D). Each egg laid was placed inside a part of faba bean stem,
which was closed at the ends by the parafilm. The eggs were
checked daily until they reached the adult stage.
The duration of insect developmental stages (egg, larva, and
pupa) were recorded by observing 100 individuals for each stage
(20 individuals in 5 replications).
Sex ratio was determined on L. algirus that emerged from field
infested faba bean. The most reliable sex character used to separate
the males and females of L. algirus is the length of the rostrum.
Following Hoffmann (1954), the female rostrum is 1/2 longer than
the prothorax or 1/4 longer (male).
Diapause
After faba bean harvest, different surveys were conducted
around faba bean fields in the region to identify the second host of
the weevil. Various fruit trees and weeds belonging to different
families were checked during summer and autumn as mentioned
by Chakir (1992) and Liotta (1963).
Feeding preference tests of new generation adults
Feeding preference tests of the L. algirus for different tree
species was tested in a closed plastic box (230 mm × 450 mm × 700
mm) under laboratory conditions (25±1°C; 70-75% relative humid-
ity (RH); a photoperiod of 13:11 h (L:D), containing a moistened fil-
ter paper and one single leaf with a similar leaf area. The host plants
tested were six trees belonging to Rosaceae family (Prunus persica
L., Prunus dulcis M., Prunus armeniaca L., Pyrus communis L.,
Prunus cerasus L., Malus domestica B); the walnut tree, Juglans
regia L. (Juglandaceae); olive tree, Olea europaea L. (Oleaceae); the
Pistachio tree, Pistacia vera L. (Anacardiaceae). A group of 10
newly emerged L. algirus adults were allowed to feed for 72 h
(25±2°C) under laboratory conditions, after a starvation period of 3
days. The control treatment consisted of faba bean leaves and each
choice test was replicated 6 times over 4 weeks. Relative preference
of individual L. algirus was calculated by counting the number of
feeding notches (FN) per leaf of different plant species.
Effects of planting date, variety, and location
on faba bean stem borer infestation
Two Moroccan local varieties Defes and the Alfia were planted
in three different regions representing different climatic zones. The
varieties were planted in rows of 4 m long and 0.6 m spacing
between rows, with a seeding rate of 100 kg/ha. The experiments
were laid out following a split-plot design with four replications.
Three different sowing dates were used, October 26, November 18
and December 23, 2016 at Marchouch experimental Station of the
International Center for Agricultural Research in the Dry Areas
(ICARDA), located 70 km South of Rabat in Morocco. At Douyet
station, two planting dates were used,7 November and December 22,
2016. During the 2017/2018 cropping season, two different sowing
dates were used at Marchouch Station on December 9 and 27, 2017.
In Allal Tazi Research Station located at 30 km from Kenitra city
(Gharb region), two dates were used, December 2 and 29, 2017.
Normal agronomic practices were followed for growing the crop.
No insecticide or fertilizer treatment were applied on the trials.
At harvest time, 10 plants were randomly sampled from each
plot and checked in the laboratory as described above.
Statistical analysis
ANOVA was analysed by using GenStat 20th edition. Insect
counts and the number of infested plants in a plot were subjected
to square root transformation to normalize variances before analy-
sis. Mean numbers of each insect stages for each variety per month
were separated using Newman-Keul’s test at p≤0.05. For the feed-
ing preference, the number of feeding notches were analysed using
one-way ANOVA followed by Newman-Keul’s test at p≤0.05.
The data on the effect of planting date, variety and station on
square root number of infested plants by faba bean stem borer were
analysed for split plot factorial design and differences among var-
Article
Non-commercial use only
ious means were determined using Tukey’s HSD test. Correlation
analysis was carried out to relate immature stages counts to mete-
orological data. The agrometeorological data of Douyet Station
was obtained using the web meteoblue (Meteoblue, 2018).
Results
Biology and dynamic populations of L. algirus during
2015 and 2016
Adult emergence
Adults are large beetles 15-22 mm long (Table 1) and have
cylindrical and slender elongated shape. The antennae and tarsus
are ferruginous. They are dark and covered with a yellowish pru-
inosity resembling pollen. Rostrum is fairly arched, a cylindrical,
slightly swollen at the level of the antenna insertion. Females have
longer rostrum than males. Adults are an excellent flyer and can
travel long distances to look for feeding sources. During the days,
the adults start feeding on the leaves of faba bean plants. They
spend almost all their time hiding among leaves of the faba bean
plants near the ground. Sometimes they climb to the higher parts
of the plant.
This weevil began to appear in both major and minor faba bean
fields on 09 February 2015 (Figures 1 and 2) under photoperiod
11L:13D and mean temperature 09°C (Table 1). In the second year,
the first adult was captured on 04 February 2016 (Figures 3 and 4)
under photoperiod 11L:13D and mean temperature of 13°C (Table 2).
Mating and oviposition
Females feed for a few days before becoming sexually active
and start oviposition shortly after mating during the early flower-
ing stage of the faba bean plant. The females bore holes (Figure
3E) into and lay eggs in the lower part of the stems and rarely in
[Journal of Entomological and Acarological Research 2021; 53:9324] [page 9]
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Table 1. Body measurements (mm) of instar larvae of Lixus algirus.
Larva length (mm) Range Larva width (mm) Range
10.77±0.25 1.5 to 28.4 2.09±0.03 0.8 to 4.5
Table 2. Duration (days±S.E) of different stages of L. algirus on faba bean varieties during 2015 and 2016.
Stage Average (days± SE) Min Max
Egg 12.8±0.11 11 15
Larva 42.03±2.82 37 48
Pupa 21.73±0.21 19 24
Adult within stem 2.76±0.13 2 4
Egg to adult 79.32 69 91
Figure 1. Mean number of eggs, larvae, pupae, and adults of L. algirus per 20 plants recorded on the local large-seeded variety Defes
(Vicia faba var. major) (A) and small-seeded variety Alfia (Vicia faba var. minor) (B) during 2015, Douyet Station.
Non-commercial use only
[page 10] [Journal of Entomological and Acarological Research 2021; 53:9324]
the upper half of the faba bean plant. The L. algirus females spend
more time during the day searching for a suitable oviposition site
among different stems using antennal movement and antennal
rotation to secure the best conditions for the larvae development.
Fresh laid eggs are oval and ranged from 1 to 2 mm (Figure 3A),
creamy white and then become yellowish, smooth, transparent.
Several perforated stems showed laying holes, but only one egg
found within the stem. The eggs hatch over a period ranging from
11 to 15 days.
The oviposition period is extended from Mid-February to late
April. The number of the egg laying increased gradually with
increasing temperature and photoperiod, to reach the peaks during
March, as more adult weevils become active.
During 2015, the eggs are recorded in the local large-seeded
variety Defes from 23 February to 6 April and peaked with
0.75±0.20 eggs per plant or 3 stems and 1.1±0.12 eggs per plant on
March 23 and 30, respectively (Figure 1A). While the eggs are laid
in the local small-seeded variety Alfia from 16 February to 30
Article
Figure 2. Mean number of eggs, larvae, pupae, and adults of L. algirus per 20 plants recorded on the local large-seeded variety Defes
(Vicia faba var. major) (A) and small-seeded variety Alfia (Vicia faba var. minor) (B) during 2016, Douyet Station.
Figure 3. Stages of Lixus algirus. Egg (A); larva (B); pupa (C); emerged adult (D); laying hole (E); exit hole (F).
Non-commercial use only
March and reached the maximum value of 0.75±0.18 eggs/ plant
on 16 March (Figure 1B).
During 2016, the egg laying activity was recorded from 18
February to 21 April for the variety Defes. A strong egg laying
activity was recorded with 0.85±0.19 eggs/plant on 31 March
(Figure 2C). Whereas the oviposition activity was recorded from
18 February to 07 April for the variety Alfia. The maximum num-
ber of eggs was recorded with 1±0.16 eggs/plant on 24 March
(Figure 2D). The egg laying activity decreases significantly from
late March to early April.
Larva populations
After hatching, the larvae (Figure 3B) bore into and feed on
stems of faba bean plants from late February to mid-May. The
neonate larvae have a creamy white body, segmented, cylindrical
and legless with a brown head and dark mouth parts. The larvae
can reach between 1.5 to 28.4 mm in length, 0.8 to 4.5 mm in width
(Table 1). They begin to feed on the tissues around the egg depo-
sition and then tend to move towards the apex of the plant. After
few days, the larvae return to the collar and then go back to the
apex repeatedly. Their feeding tunnels inside the stem produce a
dark brown exudate. The larval period development varies from 37
to 48 days, with an average of 42.03±2.82 (Table 2).
The number of larvae gradually increased during March and
peaked in April. During 2015, the larvae begin to appear on the
local large-seeded variety Defes stems on March 09 and peaked
with average 1.5±0.22 larvae/ plant or 3 stems and 1.05±0.17 per
plant on 13 and 27 April, respectively. For the small-seeded variety
Alfia, the larvae were first recorded in stems on 09 March and
peaked twice with average 1.15±0.12 larvae/plant on 30 March
and 20 April, respectively (Figure 1A,B).
During 2016, the larvae fed within the variety Defes stems
from 25 March and peaked with 0.85±0.16 larvae/plant on 28
April, before declining gradually during the next months. While,
the larvae appear earlier in stems of the small-seeded variety
Alfia starting from 03 March and become most abundant in
April. The larval peaks in the variety Alfia stems with 0.45±0.15
larvae/plant and 0.80±0.16 larvae on 17 March and 14 April,
respectively.
Pupation and the emergence of new generation adult weevils
Upon completion of larval development, the mature larva gen-
erally moves towards the lower part of stem for pupation and
become inactive. The pupae with projecting legs, rostrum, and ely-
tra (Figure 3C) were found in the stem surrounded by their exudate
left after larval feeding. The pupae are white to yellow in colour
with elongated and stocky body. The females with longer and
slightly thinner rostrum than males. In the early days, the pupa
moves continuously, where the head faces the apex of the stem.
The pupa can reach between 1.6 to 20 mm in length, 4 to 5 mm in
width. The pupal period development lasts from 19 to 24 days,
with average of 21.73±0.21 (Table 2).
The earliest pupae found at the end of their feeding tunnels
inside the stems was late April and peaked between early to mid-
May. During 2015, the pupae begin to appear within stems on the
variety Defes stems on 27 April and peaked on 4 May with average
0.85±0.22 pupae/plant. For faba bean variety Alfia, the pupae were
first recorded in stems on 20 April and peaked with the average
0.9±0.09 pupae /plant on 04 May (Figure 1A,B). During 2016, the
first pupae were found within stems of the two varieties on 05 May
and peaked with average 0.80±0.17 pupae/plant on 19 May,
respectively (Figures 3 and 4).
The neonate adults (Figure 3D) remain in the stems between 2
and 4 days. After that, they chew for few hours a perfectly circular
exit hole of 5-6mm (Figure 3F) in diameter through the rostrum and
start emerging in early June. Usually, the exit holes are situated in the
upper part of the pupation cell. During this period, the adults maintain
the same color (reddish brown in the dorsal part and creamy in the
ventral side) which changes to become darker with time.
The data of 2015 (Figure 1A,B) showed that the first emerged
adults were recorded on 11 May with the average of 0.35±0.15
adult/plant for Defes variety. While, for the Alfia variety 0.05±0.03
of emerged adult/plant was recorded on 04 May. During 2016, the
first emerged adults were recorded on 26 May with 0.1±0.06
[Journal of Entomological and Acarological Research 2021; 53:9324] [page 11]
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Figure 4. Injuries caused by adults and larvae of the L. algirus. Semi-circular notching caused the adults (A); uninfected faba bean stem
(B); exudate caused by larval feeding (C).
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[page 12] [Journal of Entomological and Acarological Research 2021; 53:9324]
adult/plant for Defes variety. For Alfia variety, the first emerged
adults were recorded one week earlier on 19 May with the average
of 0.15±0.08 adult/plant.
The results of the sex ratio of the field-emerged adults of L.
algirus are given in Table 3. The ratio of the male to female insects
tends to be 1:1 in both years, this could ensure less competition, so
it can lead to an increase in mating frequency.
Feeding preference of the overwintering generation
of L. algirus
The mean comparisons showed no significant difference in
term of preference and feeding within stems in the two varieties
during the two years (Table 4).
The data generated in 2015 revealed one significant difference
(p<0.05) among the two varieties for the total number of larvae with-
in the stems during March (F=8.58, d.f.=1, p<0.043). Higher total
numbers of larvae were recorded during March for Alfia with
51±2.08 larvae. While, only 15±0.85 larvae were obtained for Defes
variety. However, no significant differences were found during the
same year in term of the total adult exit holes for both varieties.
The total number of egg-laying holes per stems laid by L. algirus
in both varieties do not differed significantly (p<0.05) within the
same months in both years. Although, there was one significant dif-
ference in total number of egg-laying holes among the two faba bean
varieties during March 2016 (F=20.51, d.f.=1, p<0.011). The data
showed more egg-laying holes recorded for Alfia variety with
115±1.19 and only 86±1.81 recorded for Defes variety. However, no
significant differences were found during 2016 for the total egg
numbers in both varieties (F=1.23, d.f.=1, p=0.33).
This comparison revealed no preference in term of egg laying
or feeding of the stem borer between the large-seeded variety
Defes and small-seeded variety Alfia.
Symptoms and damage
Adults L. algirus feed on the leaf margins of faba bean plants
starting from late winter to late spring. They feed preferably on the
upper soft leaves causing a typical marginal and semi-circular
notching (Figure 4A); and sometimes they produce a dark excre-
ment spot. The infested plants usually have a reduced total leaf
area as a result of feeding damage caused by the adult weevils,
which inhibits photosynthesis and can decrease plant growth.
The main damage is caused by the larvae feeding within stems
especially for young faba bean plants. Their feeding tunnels inside
the stem produce a dark brown exudate (Figure 4C). This feeding
damage disrupt the movement of xylem and phloem. The liquid
nutrients are cut off from the rest of the plant causing yellowing of
leaves and growth-stunting.
Effect of climatic factors on different stages
of L. algirus population
The data of the various environmental factors for the two crop
seasons 2014-2015 to 2015-2016 are given in Table 5.
Article
Table 3. The sex ratios of the field collected adults of L. algirus during 2015 and 2016.
Variety Adults emerged during 2015 Male/female Adults emerged during 2016 Male/female
Female Male Ratio Female Male Ratio
The large-seeded variety Defes 20 21 1 : 1 28 28 1 : 1
The small-seeded variety Alfia 22 23 28 28
Table 4. Mean ± standard error values of the total number of egg-laying holes, eggs, larvae, and adult exit holes in the local faba bean
varieties Defes and Alfia during 2015 and 2016, Douyet Station.
Varieties Number of egg-laying holes Number of eggs Number of larvae Number of adult exit holes
2015
February Defes 14±2.63 9±1.69 0 0
Alfia 12±1.98 10±1.60 0 0
March Defes 94±1.46 74±1.14 15±0.85 0
Alfia 126±1.96 46±1.32 51±2.08 * 0
April Defes 137±2.11 1±0.14 81±2.43 0
Alfia 104±2.04 0 71±1.16 0
May Defes 160±1.51 0 1±0.14 28±2.02
Alfia 132±0.81 0 0 32±2.02
2016
February Defes 20±1.99 13±1,20 0 0
Alfia 34±2.91 23±2.03 0 0
March Defes 86±1.81 63±0.74 6±0.54 0
Alfia 115±1.19 * 74±0.78 19±0.89 0
April Defes 153±1.98 23±1.58 43±1.63 0
Alfia 151±1.15 6±0.75 47±0.95 0
May Defes 162±1.46 0 24±1.82 3±0.42
Alfia 164±1.17 0 16±1.38 3±0.42
June Defes 27±0.13 0 0 6±0.05
Alfia 37±0.07 0 0 10±0.07
*Significant different value (ANOVA) in comparison with each variety within a month (P=0.05).
Non-commercial use only
The maximum and minimum temperature were negatively cor-
related, but not significant on the mean number of eggs for Defes
and Alfia varieties in both years. The correlation between the total
rainfall and egg numbers was positive and significant (r=0.93) for
both varieties, respectively. Whereas, a positive and non-signifi-
cant correlation was found between mean relative humidity and
egg numbers in both varieties over different years. The correlation
between egg numbers and the mean relative humidity was positive,
but statistically non-significant in both years.
The data from the present study showed that the number of lar-
vae rises with the increase of all climatic factors in 2015. All the
weather parameters were positively correlated, but not significant
on larval population growth rates. In 2016, the mean number of lar-
vae was negatively correlated, but non-significant with maximum
and total rainfall for Defes variety. Whereas all weather parame-
ters, except the mean relative humidity, were negatively correlated
but non-significantly with the number of larvae within stems of
Alfia variety.
During 2015, there was a significant negative correlation
between mean relative humidity and mean number of pupae (r=-
0,94* and r=-0,96), for Defes and Alfia varieties, respectively.
Whereas, a negative and non-significant correlation was found
between total rainfall and pupae numbers in both varieties tested.
For the other weather parameters, the maximum and minimum
temperature were positively correlated but non-significant on the
number of pupae within stems.
Host plants feeding preference of the new generation
adult weevils L. algirus
The mean number of feeding notches consumed by the adults
L. algirus was different among the 10 host plants (F=47.42; df= 9;
p<0.001).
Peach leaves were the most attractive and preferred by the adult
weevils among all tree species with average number of feeding
notches of 58.42±1.58 (Figure 5). The almond leaves were the sec-
ond most preferred with 37.71±2.11. Whereas, no noticeable dam-
[Journal of Entomological and Acarological Research 2021; 53:9324] [page 13]
Article
Table 5. Correlation analysis between climatic variables and developmental stages of L. algirus during 2015 and 2016 in Douyet
Station.
Climate variables Stages of L. algirus
Defes Alfia Defes Alfia Defes Alfia
Eggs Larvae Pupae
2015
Maximum temperature (°C) -0,33 -0,42 0,19 0,04 0,85 0,92
Minimum temperature (°C) -0,40 -0,48 0,20 0,02 0,86 0,93
Total rainfall (mm) 0,93* 0,93* 0,02 0,50 -0,74 -0,79
Mean relative humidity (%) 0,32 0,40 0,03 0,15 -0,94* -0,96*
2016
Maximum temperature (°C) -0,68 -0,68 -0,04 -0,24 0,24 0,24
Minimum temperature (°C) -0,76 -0,77 0,04 -0,20 0,35 0,35
Total rainfall (mm) 0,22 0,44 -0,43 -0,39 -0,02 -0,02
Mean relative humidity (%) 0,72 0,70 0,09 0,31 -0,29 -0,29
*r-value significant at p= 0.05.
Figure 5. The mean number of feeding notches (±SE) of the new generation adults on leaves of different tree species after 72h. Different
letters over the bars indicate distinct mean value by the Newman-Keul’s test (p≤0.05).
Non-commercial use only
[page 14] [Journal of Entomological and Acarological Research 2021; 53:9324]
age in leaf area was detected in the control (Faba bean). Likewise,
the feeding notches in the olive leaves (0.08±0.03), Pistachio
(0.67±0.13) and Walnut (6.62±0.60) were non-significant.
The rest of the Rosaceae leaves such as Dwarf cherry
(24.71±2.09), Apricot (17.29±1.80), pear (17.71±2.13) and apple
(19.33±2.31) were less preferred by the L. algirus compared to Peach
and Almond. These results suggest that the emerged adults L. algirus
feed mainly on leaves of Rosaceae trees after faba bean harvest.
Based on the survey conducted in Douyet and Marchouch
regions on different plants and tree species during the summer and
autumn times, most of emerged adults L. algirus were found feed-
ing on various trees belonging to Rosaceae family, especially the
peach and almond trees.
Depending on the availability of the host plants, they can feed
on other families of trees, such as the walnut tree (Juglandacea
family) and the Pistachio tree (Anacardiaceae family). This shows
that the adult L. algirus is a polyphagous insect feeding on many
plant species.
In general, the adults L. algirus feed less during the summer
and early autumn period, on the different parts of the tree leaves
causing notches around the leaf edges. Their movement in the trees
remains very slow until late winter. They are never found hiding in
soil cracks, under clods, or in debris on the ground.
Effects of planting date, varieties, and location on faba
bean stem borer infestation
Analysis of variance summary of infested faba bean plants by
L. algirus exposed to different planting dates, two local varieties
and stations during 2017 and 2018 are shown in Table 6.
During 2017, there were significant effects of planting date on
the infested plants by L. algirus (p<0.05) (Tables 6 and 7). In
Douyet station, the mean number of infested faba bean plants of
the small-seeded variety Alfia were lower at normal planting
(2±0.40) compared to late planting date (7.25±1.70) (Table 7). The
effects of variety and variety×planting date interaction on square
root number of infested plants were not significant (p>0.05). Also,
the effects of stations and station×planting date interaction were
not significant. The interaction of planting date× variety x station
was not significantly different (p>0.05) (Table 6).
Further, there was no difference in term of infested plants
occurring in early, normal, and late planted Alfia plots at
Marchouch station. For the large-seeded variety Defes, the number
of infested plants decreases from normal to late sowing in Douyet
and Marchouch stations. However, the mean differences were not
statistically significant according to Tukey’s HSD test (Table 7).
The mean number of infested plants was generally higher in
2017 than in 2018. The maximum number of infested plants was
recorded in 2017 for Alfia variety in Douyet station with
7.25±1.70, when the maximum mean value in 2018 was recorded
for Alfia with 1.42±0.28 in Allal Tazi station (Tables 7 and 8).
During 2018, there were significant effects of location (Marchouch
and Allal Tazi stations) on the mean number of infested plants by
L. algirus (p<0.001) (Table 6). The infested faba bean plants were
greater in Allal Tazi than in Marchouch station for both varieties in
different sowing dates.
Article
Table 6. ANOVA describing the effect of planting date, variety,
and station on square root number of infested plants by faba bean
stem borer during 2017 and 2018.
Source of variation Df Mean squares P<F
2017
Date (D) 2 28.419 0.003
Residual 6 1.552
Station (S) 1 22.781 0.068
Variety (V) 1 0.100 0.847
D×V 2 9.919 0.060
D×S 1 2.531 0.516
V×S 1 1.531 0.612
D×V×S 1 13.781 0.145
2018
Date (D) 1 0.2688 0.134
Residual 3 0.0645
Station (S) 1 9.5430 <.001
Variety (V) 1 0.5351 0.168
D×V 1 0.1657 0.417
D×S 1 0.0344 0.576
V×S 1 0.1649 0.221
D×V×S 1 0.3415 0.078
Table 7. Mean (±SEM, n=10) number of infested faba bean plants by L. algirus for three planting dates (early, normal to late planting)
at Marchouch and Douyet Stations, 2017.
Variety, Station/Planting date AALFIA, mean±SEM DEFES, mean±SEM
Douyet Marchouch Douyet Marchouch
Early 1.75±0.47n.s. 2.25±0. 75n.s
Normal 2±0.40* 2.75±1.37 4.25±0.47n.s. 2.75±1.03
Late 7.25±1.70 0.5±0.28 3.25±0.94 0.75±0.47
Early, seed sown from early to early December; Normal, seed sown from early December to early November, Late, Mid to Late December. *Significant at 5%; n.s., nonsignificant differences according to Duncan’s
multiple range (DMR) test.
Table 8. Mean (±SEM, n=10) number of infested faba bean plants by L. algirus for three planting dates (early, normal to late planting)
at Marchouch and Allal Tazi Stations, 2018.
Variety, Station/Planting date ALFIA, mean±SEM DEFES, mean±SEM
Marchouch Allal Tazi Marchouch Allal Tazi
Normal 0.275±0.06 n.s 1.42±0.28 n.s 0.45±0.09 n.s 1.02±0.30 n.s
Late 0.425±0.13 1.17±0.22 0.125±0.06 0.87±0.24
Non-commercial use only
The effects of planting date and variety and their interaction on
the number of infested plants were not significant (p>0.05). The
interaction of planting date × variety × station was not significant
(p>0.05). Lower faba bean infestation was recorded in 2018 for
late sowing date, but this difference was not significant according
to Duncan’s multiple range (DMR) test (Table 8).
Discussion
The data on population dynamics of L. algirus (Figures 1 and
2) revealed that they follow a consistent pattern over the varieties
and years, as peaks of the different insect stages were recorded on
both cultivars from late February through late May.
The emergence of adult weevils occurred in late January at
temperature >9°C and feed on faba bean leaves. Notching of leaf
margins caused by faba bean stem borer adults are much bigger
than caused by pea leaf weevil, Sitona lineatus L (Liotta, 1963).
The infestation of adult weevils increases gradually with increas-
ing temperature and photoperiod. A similar observation was
reported by Chakir (1992), who showed that the overwintering
adults appear on minor faba bean plants in late January in Aïn El
Aouda located in Rommani region. Cardona et al. (1985) report-
ed that the adults of L. algirus leave diapause around mid-
January when minimum daily temperature can be as low as 5°C
in Syria (Lattakia area).
The oviposition period lasted from mid-February to late April
and peaking during mid to late March. The peak ranged from 0.75
to 1.1 eggs per plant. Similarly, Liotta (1963) reported in Italy
(Sicily) that the females of L. algirus began to lay eggs on faba
bean stems from early March until late April with a peak during
mid-March. A similar observation was reported by Cardona et al.
(1985) in Syria, who indicated that the adult females lay eggs
mostly between February and March, but peaked earlier during
mid-February. In Tunisia, mating and laying eggs lasts from
March to mid-April (Hmem Bourissa et al., 2010). Generally, the
females bore several holes into and lay one egg in the lower part
of the stem. The egg laying activity decreases significantly from
late March to early April. The eggs hatch over a period ranging
from 11 to 15 days. In contrast, Liotta (1963) reported that the
hatching period last longer in Italy from a minimum of 15 days to
a maximum of 21 of hatching time. In Syria, the egg hatching
takes an average of 18 days (Cardona et al.,1985). Through the
two seasons of field monitoring in Morocco, the egg laying
increased significantly when flowering starts. It seems that the
egg laying activities vary depending on the phenological stage of
the plant. These indications suggest the relative importance of flo-
ral volatiles cues from plant in enabling the L. algirus to recognize
host plants from distance.
After hatching, the larvae bore into and feed on stems of plants
from late February. The number of larvae gradually increases dur-
ing March and peaked mostly in April, before declining gradually
during the following months. The larval peaks ranged between
0.45 to 1.5 larvae per plant.
The present findings are in conformity with the results of
Cardona (1985) who reported that the larval development occurs
mostly during March to April. However, the highest larval popula-
tion density was recorded in Italy between late April and early May
(Liotta, 1963).
In the present study, the larval development time varies from
37 to 48 days, with an average of 42.03±2.82. Similarly, the larval
development lasts on average 45 days in Tunisia (Hmem Bourissa
et al., 2010). Whereas, the average of the larval development time
reached 45 to 50 days in Italy (Liotta, 1963). The duration of the
total larval cycle takes a higher average of 55 days in Lattakia
region in Syria (Cardona et al., 1985). Generally, one stem-boring
larva occurs within a single stem. Two larvae may occur rarely
together in a single stem, where one larva found in lower part of
the stem and the other in the upper part. The larvae living in the
same stem will sometimes kill each other. However, the larva does
not attack the egg within the same stem. The cannibalistic behav-
iour of the L. algirus larvae were reported by Liotta (1963) and
Chakir (1992). Usually only one larva occurs within the stem
where it completes all its different stages. Cannibalism has also
been reported for other Lixus species, such as Bitter Leaf Weevil
(L. camerunus) and the Rhubarb Curculio L. concavus Say (Eluwa,
1979; Capinera, 2001).
The earliest pupae were found inside the stems in late April
and peaked during May. The pupae peaks ranged between 0.8 to
0.9 pupae per plant. The pupal period development lasts from 19
to 24 days, with average of 21.73±0.21. The pupal stage lasted less
in Syria with an average of 17 days (Cardona et al., 1985) and
between 10 to 17 days in Italy (Liotta, 1963).
In the present study, the neonate adults remain in the stems
between 2 to 4 days, and these findings are in conformity with the
results reported in Italy by Liotta (1963). In contrast, Cardona et al.
(1985) reported that the new adults remain within faba bean stems
for 10 to 12 days in Syria.
Our study showed that the entire life cycle (egg to adult) lasts
approximately between 69 to 91 days with an average of 80 days.
Liotta (1963) also reported a similar duration of life cycle which
lasted 72 to 92 days. The complete cycle of L. algirus in Syria was
90.5 days (Cardona et al. (1985).
Our findings indicate that there is only one generation of L.
algirus on faba bean. Throughout its area of distribution,
Morocco, Tunisia, Syria, Italy and Spain, only one generation a
year has been observed for this pest (Liotta, 1963; Isart, 1968;
Cardona et al., 1985; Chakir, 1992; Boukhris-Bouhachem, 2013).
However, 3 generations per year in Algeria and two in France
were reported by Hoffmann (1963). Hoffmann (1954; 1963) indi-
cated that the number generation of L. algirus depend on weather
conditions.
In the present study, seasonal variations of the weather factors
affected the development and the trend in population fluctuations
of L. algirus on both faba bean varieties. The number of eggs
increases with the rise of precipitation and humidity. In addition,
there was a significant negative correlation between mean relative
humidity and mean number of pupae. Similarly, in certain coastal
areas of Lebanon, Syria, and Turkey with high humidity and pre-
cipitation, higher Lixus infestation were found on faba bean vari-
eties (ICARDA, 1982).
After faba bean harvest, the new emerged adults L. algirus
were found feeding on various fruit trees belonging to Rosaceae
family during the summer and early autumn period. The new
generation adults are most attractive to Peach leaves followed by
Almond leaves. The Dwarf cherry and Apricot were less pre-
ferred. Other families of trees could be host plants for the
emerged adults L. algirus, such as the walnut tree (Juglandacea
family) and the Pistachio tree (Anacardiaceae family). Similary,
Liotta (1963) reported that the new adults can feed on Cardaceae,
Malvaceae and Rosaceae including almond, peach and apricot
tree leaves after faba bean harvest. The same author also reported
that the new emerged adults survive the winter by hiding under
the rocks.
In the present study, there is considerable evidence that the
new emerged adults remain active during summer and autumn
period feeding on different parts of the Rosaceae leave trees. In
[Journal of Entomological and Acarological Research 2021; 53:9324] [page 15]
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[page 16] [Journal of Entomological and Acarological Research 2021; 53:9324]
contrast, Chakir (1992) showed that the adults overwinter in the
soil or in the dry stems of host plants. According to Isart (1967),
adults can live up to 10-12 months feeding sporadically on beet
and artichoke during summer and autumn.
Several authors described differently the diapause of several
Lixus species. According to Trnka et al. (2016), adults of both
species L. neglectus (Fremuth, 1983) and L. bituberculatus
(Smreczyński, 1968) do not hibernate in the host plants, but most
likely in the leaf litter, among dry plant debris or in the topsoil.
In contrast, the species L. filiformis (Fabricius, 1781) overwinters
as larva or pupa in the stems of host plants. Gültekin (2008)
reported that L. myagri (Olivier, 1807) and L. circumcinctus
(Boheman, 1835), realized their hibernation in the rootcrown of
host plant. The same author reported several other Lixus species,
which migrate before hibernation; this includes L. furcatus
(Olivier, 1807) and L. obesus (Petri, 1904) and L .siculus
(Boheman, 1835) and L. korbi (Petri, 1904).
In addition, there was no difference in term of infested plants
occurring in early, normal and late planted Alfia and Douyet vari-
eties at different stations. The results showed that the number of
infested plants decreases from normal to late sowing. However, the
mean differences were not statistically significant. This finding is
not consistent with Cardona et al. (1985) who showed that late
planting date in Syria significantly decreased the percentage of
stems infestation by L. algirus. Stems infested in early planting
(early October) recorded about 35% instead of only 10.6% record-
ed in late planting (late November-early December) in Terborl,
(Lebanon). Similarly, in Lattakia (Syria), less infested stems were
recorded in late planting (early January) with 4.1%; instead of
51.2% recorded in early sowing (late October-early November).
According to the same study, changes in planting dates have a
great effect on oviposition patterns, with late planted crops show-
ing significantly less infestation. However, the same authors do not
recommend delaying planting date from early November to late
December or early January, because of various factors including
rainfall patterns, diseases, and weeds incidence. Furthermore, a
wide range of management strategies for the control of faba bean
stem borer had been tempted before. In Tunisia, application of car-
bofuran as seed treatment and foliar sprays of different insecticides
did not reduce significantly stem borer infestations (Coers et al.,
1983; Weigand & Bishara, 1991). Efforts should be placed on host
plant resistance to identify sources of resistance to this pest. Also,
other sustainable control measures need to be tried as well, such
semiochemicals for detection and monitoring or mass trapping
using the attract and kill approach.
Conclusions
The present findings showed many similarities on the dynamic
and field biology of the faba bean stem borer across the
Mediterranean countries. It was found that the oviposition period
extends from Mid-February to the end of April. The numbers of
larvae gradually increased significantly and peaked in April.
Larval feeding within stems caused significant damage. Only one
generation is observed each year in Morocco. Abiotic factors like
total rainfall and relative humidity influenced egg laying activity
of L. algirus on different varieties of faba bean. However, the use
of planting dates did not significantly impact the infestation level
of L. algirus among the varieties tested and locations where the tri-
als were conducted. The information generated in the present study
would be helpful for the development of management options for
the faba bean stem borer in Morocco.
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[Journal of Entomological and Acarological Research 2021; 53:9324] [page 17]
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The pea leaf weevil, Sitona lineatus L. (Coleoptera: Curculionidae, PLW), is a significant insect pest of pea (Pisum sativum L.) and faba bean (Vicia faba L.), and it is responsible for serious economic damages. The current study aims to investigate the insecticidal activities of seven essential oil species on S. lineatus adults in the laboratory as well as the most effective ones under growth chamber conditions. Out of the tested essential oils, Lavandula angustifolia Mill. and Mentha arvensis L. oils showed substantial insecticidal activity against S. lineatus adults (100%) by contact activity at a concentration of 1.5% after 1 and 3 h of application, respectively, with an estimated LC50 of 0.224 and 0.320% at 72 h after treatment, respectively. L. angustifolia essential oil also resulted in a significant mortality rate of 70% by ingestion activity, with an LC50 of 1.340% at 48 h after application under laboratory condition. The growth chamber bioassays confirmed these significant effects, displaying a mortality rate of 70% at 96 h after application at 1.5%. The gas chromatography−mass spectrometry analysis showed that linalool acetate (32.91%) and linalool (26.69%) dominated the L. angustifolia essential oil composition. These findings highlighted that L. angustifolia essential oils are promising agents for the development of biopesticide formulation for the control of S. lineatus, as a safe and ecofriendly alternative compared to chemical insecticides.
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Earlier models of integrated pest management (IPM) focused on ecological aspects of pest management. With the recent developments in agricultural technology, modern communication tools, changing consumer trends, increased awareness for sustainably produced food systems, and globalization of trade and travel, there seems to be a need to revisit the IPM paradigm as appropriate for modern times. A new model, built on earlier models based on ecological and economic aspects, is expanded and reconfigured to include management, business, and sustainability aspects and emphasize the importance of research and outreach. The management aspect contains four components of IPM that address the pest management options, the knowledge and resources to develop management strategies, the management of information and making timely decisions, and the dissemination or sharing of information. With the business aspect that includes the producer, consumer, and seller, and the sustainability aspect that covers economic viability, environmental safety, and social acceptability, the new model presents the human, environmental, social, and economic factors that influence the food production.
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The mature larvae and pupae of Lixus (Ortholixus) bituberculatus Smreczyński, 1968 and L. (Dilixellus) ne- glectus Fremuth, 1983 (Curculionidae: Lixinae: Lixini) are described and compared with known larvae of 21 other Lixus and 2 Hypolixus taxa. The mature larva and pupa of L. bituberculatus are the first immature stages described representing the subgenus Ortholixus. The larva of L. neglectus, in the subgenus Dilixel- lus, is distinguished from the known larvae of four species in this subgenus by having more pigmented sclerites on the larval body. All descriptions of mature larvae from the tribe Lixini, as do all known species from the tribe Cleonini, fit the diagnosis of the mature larva of the Lixinae subfamily. Furthermore, new biological information of these species in the Czech Republic, Slovakia and Romania is provided. For L. bituberculatus, a chicory, Cichorium intybus L. (Asteraceae), is identified as a host plant, and L. neglectus is found on dock Rumex thyrsiflorus Fingerh. (Polygonaceae). Both species are probably monophagous or oli- gophagous. Adults of L. bituberculatus often inhabit host plants growing in active, dry and sunny pastures with sparse patches without vegetation, being mostly active during the night in April/May and then again in September, when the highest activity levels are observed. Adults of L. neglectus inhabit dry grasslands on sandy soils with host plants, being active during the day from May to September, with the highest level of activity in May/June and September. The larvae of both species are borers in the stem and root of the host plant, and they pupate in root or root neck. Adults leave the pupation cells at the end of summer and do not hibernate in the host plants. Finally, Romania is a new geographic record for L. bituberculatus.
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 N2 fixation by leguminous crops is a relatively low-cost alternative to N fertilizer for small-holder farmers in developing countries. N2 fixation in faba bean (Vicia faba L.) as affected by P fertilization (0 and 20 kg P ha–1) and inoculation (uninoculated and inoculated) with Rhizobium leguminosarium biovar viciae (strain S-18) was studied using the 15N isotope dilution method in the southeastern Ethiopian highlands at three sites differing in soil conditions and length of growing period. Nodulation at the late flowering stage was significantly influenced by P and inoculation only at the location exhibiting the lowest soil P and pH levels. The percentage of N derived from the atmosphere ranged from 66 to 74%, 58 to 74% and 62 to 73% with a corresponding total amount of N2 fixed ranging from 169 to 210 kg N ha–1, 139 to 184 kg N ha–1 and 147 to 174 kg N ha–1 at Bekoji, Kulumsa and Asasa, respectively. The total N2 fixed was not significantly affected by P fertilizer or inoculation across all locations, and there was no interaction between the factors. However, at all three locations, N2 fixation was highly positively correlated with the dry matter production and total N yield of faba bean. Soil N balances after faba bean were positive (12–58 kg N ha–1) relative to the highly negative N balances (–9–44 kg N ha–1) following wheat (Triticum aestivum L.), highlighting the importance of rotation with faba bean in the cereal-based cropping systems of Ethiopia.
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Faba bean (Vicia faba L.) is a protein-rich legume seed well adapted to most climatic areas of Europe and widely used for feed and food. Even if the seed is generally recognized to be of good nutritional value, existing genetic variability for seed composition offers possibilities for improvement of this trait by breeding. Four major quality types must be distinguished according to the presence or absence of tannins in the integuments and of vicine (V) and convicine (C) in the cotyledons. The nutritional value of diets containing varying amounts of different faba bean cultivars characterized by high or low levels of tannins, and high or low levels of vicine + convicine (VC), has been examined in monogastric animals and ruminants. Low-tannin content generally results in higher protein and energy digestibility for monogastric animals and low VC content has a positive effect on laying hen and broiler production performances. V and C, inactive precursors of divicine and isouramil are redox compounds potentially toxic to human carriers of a widespread genetic deficiency of the erythrocyte (red blood cell, RBC) enzyme glucose-6-phosphate dehydrogenase (G6PD). Ingestion of faba beans by these deficient individuals may cause a severe, potentially lethal hemolytic anemia (favism). The mechanism of action of divicine and isouramil in the G6PD-deficient RBC is discussed.Beside the positive impact of using tannin-free varieties in monogastric animals diets, the development of faba bean cultivars with very low levels of VC would represent a real advantage in terms of nutritional performance in poultry diets and of food safety to humans.
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The faba bean (Vicia faba L.) crop often experiences drought during its growth and development such that soil moisture deficits constrain its production. As droughts are predicted to increase in both frequency and intensity due to climate change, a better understanding of drought response patterns and associated traits is essential for obtaining yield stability in water-limited environments. This review deals with adaptation mechanisms associated with drought avoidance, escape and tolerance, with an emphasis on physiological traits such as stomatal conductance, carbon isotope discrimination and leaf temperature. Leaf temperature is considered an effective surrogate measure for other measures of stomatal characteristics. Drought tolerance through osmotic adjustment has not yet been demonstrated in faba bean although it is found in many other legumes including chickpea and pea. Deeper root growth, leading to uptake of otherwise unavailable water, helps the plant to avoid drought by delaying dehydration, but genetic variation and heritability of the trait are essentially unknown for faba bean. Crop management strategies, such as early planting, and appropriate phenology, are particularly important for drought escape in regions where terminal drought is common. Disease resistance is especially important in drought-prone areas to reduce the need for expensive control measures when yields are uncertain. The relevance of soil fertility status and nutrient availability are also covered. Drought escape and ascochyta blight resistance are important breeding objectives for terminal drought regions. Some form of drought resistance is necessary for the transient droughts experienced in most regions, and drought avoidance can be screened by a combination of leaf temperature or other rapid test of stomatal characteristics followed by carbon isotope discrimination in the most valuable materials. No single trait is adequate to improve yield in drought-prone environments, rather, a combination of characteristics is needed.
-Pests of Vicia faba L. other than aphids and nematodes
  • Bardner R
BARDNER R., 1983 -Pests of Vicia faba L. other than aphids and nematodes. In: The Faba Bean (Vicia faba L.). P.D. Hebblethwaite ed. -Butterworths, London, UK :371-390.
-La recherche scientifique en entomologie depuis 1947 à l'INRAT: un itinéraire si riche et si glorieux, mais quel avenir?. -Annales de l'INRAT, numéro spécial centenaire de l'INRAT
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BOUKHRIS-BOUHACHEM S., 2013 -La recherche scientifique en entomologie depuis 1947 à l'INRAT: un itinéraire si riche et si glorieux, mais quel avenir?. -Annales de l'INRAT, numéro spécial centenaire de l'INRAT. 86: 191-212.
Note d'information sur les pourcentages d'infestation des fèves et féveroles par les larves de Lixus spp
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COERS P., TRAIA M., ZOUAGHI N., 1983 -Note d'information sur les pourcentages d'infestation des fèves et féveroles par les larves de Lixus spp. Notes techniques 85/1. Ministère de l'Agriculture, Station de la défense des Cultures du Nord-Beja. Projet Tuniso-Belge. Mimeographed hand-out CARDONA C., JOUBI A., TAHHAN O., 1985 -Field biology of the faba bean stem borer, Lixus algirus L. (Coleoptera: Curculionidae) in Syria [Vicia faba]. -FABIS. Newsletter. 12 :14-17.