1. Jfuas No.1 June 2013
53
Ovipositon sites of Zonocerus variegates (L.) along the
major seasonal water courses(Wadies) in Darfur Region.
1-Ibrahim Abdel/Rahman Ibrahim Abdel/Fattah, Faculty of Enverionmental
Sciences and Natural Resourses, Department of Plant Protection, University of Al
Fashir
2.Elwasila Guddoura Mohamed, Faculty of Agriculture, Department of Crop
Protection, University of Khartoum, Sudan.
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Abstract
This study was carried out in Darfur Region during the period (1999 -
2002), and its objective was to identify the insect pest, it is life cycle,
ecology and behavior. Zonocerus species currently existing in
Western Darfur is the variegated grasshopper, Zonocerus
variegatus(L.),with two local names, Maradona and Bo. The species
has got only one generation per year, both under field and laboratory
conditions. But it splits into two distinctive populations, the dry and
wet season populations due to the effects of ecological factors e.g. soil
moisture and temperatures. The dry season population has large
density and causes considerable damage to the summer crops. The two
populations overlap and interbreed with each other by the end of the
rainy season. Results indicated that this pest has infiltrated into
Western Darfur through seasonal water courses (wadies) which run
across the borders with Chad and Republic of Central Africa and
recorded as a pest in season (1994) at El Gineina. The pest has a close
ecological relationship with the bush Ipomoea fistulosa, which
provides the grasshopper with oviposition sites under its shade,
hiding, resting places and probably secondary plant metabolites to
3. Jfuas No.1 June 2013
55
develop its defense mechanism against natural enemies. Results
indicated that the egg pods density in one square foot is about
30.66+14.31, each with 21-96 eggs/pod.
Keywords: Bo- locust / grasshopper, Zonocerus variegates, Wadi, Gerf soil, seasonality
1. Introduction
There are several grasshopper species capable of inflecting
considerable damages on different crops on many occasions in the
Sudan. As the grasshoppers have the capacity to live in a wide range
of habitats with great variations in the details of their biology and
behavior, the problem of these insect pests is becoming a chronic one
in the Western Sudan particularly in Darfur Region.
Indigenous grasshoppers, have for long being a real threat to the
Qoz and Wadi- cultivators, where significant losses usually occur on
millet, sorghum and vegetables. Thus, the food situation is becoming
unsecured in the marginal areas of the western states of the Sudan,
where grain production is risky.
It is well known that, unusual weather changes and land use are
the main factors responsible for encouraging grasshoppers problems.
However, the Central Plant Protection Directorate (P.P.D) has for long
keeping a close guard, through its effective control measures against
quite a number of pests including the grasshoppers. Though the
implementation of the new privatization approaches on pest control
strategies have deprived the farmers from the essential (P.P.D)
support. The new policies, states that, minor pests including
grasshoppers are becoming farmers responsibility.
4. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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Under the poor economic situation, most of the farmers are
financially incapable to purchase pesticides and spray equipment to
combat the grasshoppers. Accordingly their reproductive potential has
grown unchecked above economic injury levels. During seasons, 1995
and 1998, North Darfur witnessed local outbreaks of grasshoppers
which were restricted to certain habitats in El Fashir, Kebkabyia and
Umkadada provinces. In Eliet Gar Elnabi (Umkadadda), a total area of
5000 feddans grown by groundnuts, millet and other field crops were
completely destroyed by the significant populations of (11)
grasshoppers species acted as a group (P.P.D,1995). The same is true
in Kebkabyia, when the hole Jebel Ci area, 80 Kms in length from
Kagrio in south up to Milla in north was devastated by (21)
grasshoppers species act as a group, (Oxfam, 1999).
Though some injurious species like Oedaleus senegalensis
(Kra.),Aiolopus simulatrix (Walk.) etc.. Each can by own cause severe
crop damage.
Food situation might become worse since a new grasshopper,
Zonocerus variegatus (L.) appeared in western Sudan. Of all identified
locusts and grasshoppers, Zonocerus variegates (L.) was regarded as
an important species together with some other injurious grasshoppers
ie. Hieroglyphus daganesis (Kra.), Oedaleus senegalensis,(Kra.),
Kraussaria angulifera (Kra.), Aiolopus simulatrix (Walk.) and
Anacridium melanorohodon (Walk.) (Popov, 1989).
Mohammadein (1996) stated the first appearance of Z.
variegates (L.) in an outbreak in Western Darfur State where an area
5. Jfuas No.1 June 2013
57
of about 75190 hectare was infested by this serious grasshopper. Since
that time the pest was continuously endangering the whole agricultural
sector in Darfur Region and migrating inward to invade new areas. Z.
variegates (L.) is strictly polyphagous grasshoppers with a wide range
of food acceptance. It does seem that the species alone can cause a
massive loss on different crops grown along the wadies. Although this
grasshopper has reached a pest status recently, yet it's significant
damaging levels on the economically important cereal and
horticultural crops ranked it as one of the most serious insect pests
currently existing in the whole region. Because, Z. variegates has been
recorded a pest recently in the Sudan therefore, cumulative knowledge
about its biology, ecology behavior control etc, is almost unknown in
the Sudan.
Hence, the study is aiming to provide knowledge on various
subjects conditions related to the understanding of the grasshopper
under the local condition through conducting comprehensive field and
laboratory investigations. Expected findings are required to help in
solving the current problems under field condition and to build a base
for future studies.
2. MATERIALS AND METHODS
The study was carried out in the greater Darfur region with
special emphasis to the major seasonal water courses (wadies),Kaja,
Barei, Azum , Salih and their tributaries of the western, northern and
southern Darfur states Fig(1).
6. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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The course of the study took three successive seasons, 1999,
2000, 2001 and 2002, during which various ecological, biological etc.
aspects related to the newly infiltrated insect pest, the variegated
grasshopper, Zonocerus variegatus(L) were investigated, both under
field and laboratory conditions. During the course of work biological
studies were carried out through using an effective qualitative and
quantitative methods mainly observations, identification of specimens
, photographing, diagrams and classification of the different instars
according to age and population types by random sampling technique
(for encouraging information capacity building of the subject under
test, surveys and control facilities of the Plant Protection Department
(P.P. D.) ElGineina, Kebkabyia, and Zalingei were put in use during
the entire course of field studies). Meanwhile fresh samples of
different stages of Zonocerus and population types were regularly
supplied by P.P.D. working teams for further investigations and
making comparisons. Accordingly, infested areas out of reach became
quite accessible to cover as wider areas as possible
Studies were aimed firstly to identify the common Zonocerus species
currently exiting in Darfur States.
Adults were randomly collected from host plants using
sweeping and hand picking during low temperatures i.e early
mornings and late evenings. The specimens were indentified using the
key of the common species of the sahelian grasshoppers (1987), as
well as using the photo plates and descriptions presented in the texts,
guides and handbooks, developed by Drish 1965, Mestre1988,
7. Jfuas No.1 June 2013
59
locoq1988, and Steedman1990. Comprehensive studies covered, the
migration of the species into the western State , it's host range and
nature of damages, biological cycle and the phenomenon of
population dynamics and oviposition sites , natural enemies etc.
information gathered were recorded accordingly.
9. Jfuas No.1 June 2013
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3. Results
3.1 Oviposition site description
58% of the total searched bushes on the two breeding habitats
(wadi slopes+ water collecting sites) were found containing egg pods.
Also 62.07 % of the discovered oviposition sites were found along the
slopes of the wadies, whereas 39.93% were found around water
collecting sites closer to the main courses.
With respect to the type of pushes selected for egg laying,82.76%
of the discovered oviposition sites were found under Ipomoea
fistulosa growing alone , whereas 17.24% of the discovered egg fields
were found under I. fitulosa growing together with other plants
species i.e. wild trees ,citrus of the wadi banks, table (1).
Oviposition sites were not detected under I.fistulosa growing in
habitats that lack essential ecological requirements responsible for
encouraging laying such as suitable soils enough moisture etc On the
other hand Oviposition sites were also not detected under some
I.fistulosa growing adjacent to others containing eggs although
Oviposition inductive factors were quite abundant during the breeding
season .
During the breeding activities, adults become concentrated in
habitats possessing specific egg laying requirements. These habitats
were examined firstly by a limited number of females prone to laying
where then many other individuals from the neighboring bushes
become attracted partly by the effects of the gergrization pheromones
10. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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evolved by the initial population and partly by the bright yellow
coloration.
With regard to the type of habitats selected for breeding, it was
found difficult to detect oviposition sites around water collecting sites,
because egg laying holes were concealed by distortions made on soil
surface by the movements of the visiting birds and animals.
Such oviposition sites can only be discovered after removing the
upper most soil layer.
However, when the top soil layer was removed carefully with a
knife, egg holes became quite visible with a reddish foamy substance
topping the egg pods. In contrast it was found easy to detect
oviposition sites along the wadi slopes (gerf soils) the most attractive
habitats to Z.variegatus(L.)
As wadi extended flood plains were confined to certain areas ,
large oviposition sites have been discovered under I,fistulosa during
seasons, 2002-2001, at the lower stream zone of wadi – salih
(klimbasanga) and kayang flood plains of Wadi Bari , Saraf Umrra,
Abutinnayia.F ig.(2)
The exact egg laying sites have been found in the mid zones of the
wadi slopes (gerf soils), approximately 30 cm above the wadi base or
foot, where I,fistulosa shade falls onto. However, in certain areas
I,fistulosa bushes shade was supported by some more shade of the
largest citrus (mango, guava) and other wild trees(indigofera
sp.Acacia nilotica etc.) growing adjacent to I,fistulosa on the wadi
banks). An oviposition site can extend laterally along the wadi slope
11. Jfuas No.1 June 2013
63
where the shade extend provided that, and accordingly the breeding
population is increased if other favorable factors are still prevailing. In
contrast the width of an oviposition site was restricted vertically up
and downward across the slope according to the levels of soil
moisture. There exit at the wadi base or foot a permanent, but broken
or discontinuing water ponds that apparently appear in the half way of
the running course all the year round drop .
Table(1) : The oviposition sites of Zonocerus variegatus(L.) discovered
under different plant species growing in different breeding habitats
along wadi Kaja, EL Gineina province, season 2002
Type of
breeding
habitats
No. of
sample
Plant species Total
oviposition
sites
discovered
%discovered
oviposition
sites
I.fistulosa I.fistulosa
other
plants
Wadi-
slopes(Gerf-
soil)
25 15 3 18 62.07%
Water
collecting
sites
25 9 2 11 37.93%
Cultivated
areas
No sample
taken
Total 50 24 5 29
% Total 82.76 17.24 58%
ϯ͘Ϯ dŚĞ ƐŝnjĞ ŽĨ ĂŶ ŽǀŝƉŽƐŝƚŝŽŶ ƐŝƚĞ
Measurements of oviposition site revealed that, the egg laying of
Z.variegatus(L.) hasn't got a fixed size or a shape but , it varies from
12. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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site to site, and ranges between (L 2.8-12x W 1-2m) with an average
mean of 8.31±6.06 square meter,
Table (2): Results of soil analysis used by Zonocerus variegates for egg
laying:
s.p
pH
ECe
Na
g/L
K
ca
Mg
CO3
HCO3
CI
sand
Silt
Clay
Texture
45 7.8 55 3.6 0.3 1.0 4 0.5 2.5 1.5 54 24 22
Sandy
clay loam
ϯ͘ϯ ^Žŝů ĐŚĞŵŝƐƚƌLJ
Soil is sandy clay loam. It is characterized by high water saturation % (S. P).
PH in neutral none salinesoils. Moderate cations and anions. Non
compacted soils or fragiable soils easy to be fragmented.
Table (3): the oviposition site investigation of Zonocerus variegatus(L.)
carried out at wadi Kja, season 2002.
Description No of
samples
taken
Range Average
The size of oviposition site . 10 L2.8-12x
W 1-2m
8.31±6.06m
Number of egg per pod 30 21-96 eggs 53.2±19 eggs/pod
Length of an egg laying hole 27 9-13 cm 11.22±1.16cm
Height of an egg pod 28 2.8-5cm 3.87±0.88 cm
Foamy secretion height 27 3-6 cm 4.3±0.76 cm
Number of egg pods in one foot
square
50 2-59 egg
pods
30.66±14.31 egg
pods/one foot
square.
The density of egg pods in one foot square.The density of egg pods of
Z.variegatus(L.) ranges between 2-29 egg pods/foot square with an
average mean of 30.66±14.31 egg pods per one foot square.An egg
laying hole, an egg pod, egg per pod and foamy substances of
Z.variegatus(L.)The depth of an egg laying hole of Z.variegatus (L.),
ranges between 9-13 cm with an average mean of 11.22±1.16 cm. the
hole is prepared to accommodate two component, the egg pod and the
13. Jfuas No.1 June 2013
65
foamy substances fig(2) . An egg pod is made of earth while a female
undergoing it's laying process during the breeding season. It is located
at the lower part of the egg laying hole its height ranges between 2.8-
5 cm, with an average mean of 3.87±0.88 cm. Each egg pod contains,
21-96 eggs with an average mean 53.2±19 eggs. Eggs were laid in V
shape with instar emergency exits on top.
The upper most soil layer (3-6 cm) where foamy substances were
secreted was found dry. In contrast the lower layers where egg pods
were inserted retained considerable amount of soil moisture. An egg
pod prevents loss of moisture from the eggs and it was topped by
foamy substances up to the soil surface.
The newly secreted foamy substances have the color white, then turns
into reddish by time.
Its height ranges between 3- 6 cm with an average mean 4.30 ± 0.76 cm.
Parasites, predators and pathogens
Large numbers of egg pods of Z.variegates (L) were found infected by
pathological agents at the northern slopes of wadi Kaja 5 Kms
approximately south of EL Gineina city. Results of the test revealed
that, the causal agents are fungal pathogens belonged to the family
zoopagaceae, according to the Botany Department , Faculty of
Science, University of Khartoum .
3.4 Oviposition soils analysis
Soil samples (containing Z. variegates eggs) were analyzed at the
laboratory of the Department of Land Use and Desertification Control
(Khartoum).Results of the analysis is presented in Table (2).
14. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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1. Discussions
Results of this study indicated that, the wadi slopes and the water
collecting sites are the main breeding habitats, (58%) of the searched
bushes found containing egg fields, with 62.07% of the total sites
discovered along the slope. This would, however proves that the wadi
slopes are the best habitats for laying eggs along which favorable
ecological conditions prevail, such as suitable soils, food sources,
shelters etc. with the fact that 82.76% of the discovered oviposition
sites found under I.fistulosa would however, ensure that, there exist
along the wadies a closest relationship (but not exclusive) between Z.
variegatus (L.) and the bush I.fistlosa probably for satisfying a variety
of options, but unlikely to be important as a food plant, although the
pest feeds upon and that the plant is not consumed for it's food value .
Results showed that the plant is not only selected for oviposition
purposes but also for hiding and a resting place.
However the plant may also be used for getting secondary metabolite
I.e. PAs which they later employ for their defense against predators
and parasites. Abdel Hadi (1987) identified the alkaloid ergometrine
from I.fistulosa which is probably the compound that causes Z
.variegatus (L.) to be attracted. The pharmacophagous relationship
between Z. variegatus (L.) and the plant C.odorata was intensively
studied by Bopper (1991), Bopper and Fischer (1994).Modder (1988)
and Idown and Akinsete (2000). In addition to C. odorata, Z.
variegatus (L.) has a wide host range including several additional PAs
15. Jfuas No.1 June 2013
67
plants such as Heliotropium (Boraginaceae) as well as Emilia and
Ageratum (Asteraceae).
The grasshopper is attracted to pass during their entire life
independent of the amount of past already ingested (Bopper and
Fischer, 1994). Results indicated that, the closest relationship of the
pest with I.fistulosa would ensure that, Z. variegatus(L.) has got many
PAs alternative plants in nature which can make use of in the absence
of its classical host plant, C. odorata.
In this sense, the need for a comprehensive work of PA plant sources
including, I. fistulosa is important in the view that, the survival
strategies and seasonal occurrence of this harmful insect is
ecologically geared with noxious plant species usually neglected.
The absence of egg fields from I. fistulosa lacking essential
oviposition requirements would ensure that the pest is purposely
attracted to the bush for getting certain requirements essential for
survival.
In contrast the absence of oviposition sites under some I. fistulosa,
although encourageable factors for breeding are quite abundant, is
attributed to the dynamic trends of adult populations of the pest which
is behaviorally controlled by the gregarisation pheromones during the
breeding activities.
The detection of oviposition sites can be made by searching areas
most likely to contain eggs specially under I. fistulosa growing at the
wadi slopes (Gerf soils), by digging with a knife. Also the remains of
the dead bodies can be making use of as good indicators. However,
16. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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some difficulties may be experienced when seeking for oviposition
sites around water collecting sites which require digging or removing
upper soil layer. Results confirm that the bulk of oviposition sites of
the pest occur at the wadi slopes which gives early rise to the first dry
season hatchlings. Oviposition sites appear in the form of colonies, but
extend along the slope. This finding confirms the work done by Popov
(1978).
The discontinuing water ponds along the wadies specially at the half
ways provides a permanent moisture sources which keep eggs remain
viable during the dry conditions.
In this study, the size of an egg field measured 8.31±6.06m², although
pans Manual (1986) reported that egg laying sites are unlikely to be
more than 5m².
However , results showed that egg laying sites haven't got specific size
nor a shape, but their vary with respect to the size of the effective
breeding females, they fertility levels suitability of soil and
availability of soil moisture, space availability, the shade offered by
the bush plant etc.
The present study indicated that, the density of Z. variegatus (L.) egg
pods in one foot square is about 30.66±14.31 at the wadi slope (Gerf
soils).
When converted into m², it will be 340.63 egg pods (28 dozens) which
are considered a high density. This finding agrees with Pastre et al.
(1989) who reported that, the laying is closely grouped, and the
density of the egg pods of the variegated grasshopper is sometimes
17. Jfuas No.1 June 2013
69
very high (several dozen per square meter). The high egg pod density
per unit area will provide a rough measure forecasting the size of the
expected populations and their menace accordingly. It would also help
in planning for selecting an appropriate control measure to destroy
eggs before hatching specially when using infections fungal
pathogens. In this regard the fungal pathogen found attacking the eggs
is promising.
The study indicated that, the depth of an egg hole is about 11.22±1.16
cm which differ from pans Manual (1986) who reported 15 cm. These
differences may be due to differences in soil types and their condition
under which eggs were deposited.
The upper most soil layer (3-6cm) is dry. In contrast the lower layers
where eggs are imbedded contain considerable amount of moisture
which enabled eggs in diapauses remain viable during the dry
conditions. This type of survival strategy is referred to as survival
strategy in the egg stage which is very common on many grasshopper
species (Popov, 1989).
The foamy substances secreted by the females during the laying
process act as a protective cover topping the hole, and a passage or an
exit through which the newly emerged instars can get outside the pod
hole ventilation can also be made accessible through this substances.
It is possible to suggest that these foamy substances which may
contain secondary plant metabolites Idowu and Akinsete (2000), in
this study, probably, from I. fistulosa might temporary afford some
sort of the protection to the incubated eggs against the natural enemies
18. اﻟﻔﺎﴍ ﺟﺎﻣﻌﺔ–ﻣﺠﻠﺔاﻟﻌﻠﻮماﻟﺘﻄﺒﻴﻘﻴﺔ–اﻟﻌﺪداﻷول–ﻳﻮﻧﻴﻮ2013م
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at least during the laying period. This suggestion agrees with Bopper
(1991) who reported that, the secondary plant compounds obtained
from C. odorata are stored and chemically protect the grasshopper and
practically their diapausing eggs from predators i.e.(Larvae of
Mylabris Beetles or Parasitoids).
However, repellency may be lost by time due to the effects of the
physical factors and eggs may become liable to attack by egg parasites
and predators. Modder (1988) reported that 40% of dry season eggs
are predated suggesting that stored PAs give no protection.
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