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Inventaire et bioécologie de quelques orthoptères dans la vallée du M'Zab (GhardaàŻa)


par Youcef ZERGOUN
Université Kasdi Merbau de Ouargla - Doctorat ès sciences 2020
  

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Abstract

Caelifera is one of the largest and most diverse group of insects, and they are the dominant Orthoptera in agriculture ecosystems. We present here an inventory of Acridid fauna of the agricultural ecosystems in the Mzab valley (Septentrional Sahara, Algeria). Grasshoppers were sampled with quadrats in Béni Isguen, Ghardaïa and El-Atteuf, and we have expressed the species richness, sampling effort and relative abundance. The method used is that of quadrats sampling. The results revealed the presence of 27 species, divided into three families, Acrididae, Pyrgomorphidae and Tetrigidae. Among these families, we found that Acrididae are most diverse with six subfamilies. The subfamily Oedipodinae with 10 species was the most abundant, while the subfamily Tetriginae was the least abundant (represented by a one species). The value of the diversity index showed that the cultivated area of Béni Isguen is the most favorable for the development of many Caelifera. The abundance of grasses and low intensity of agricultural activity in this environment favors the presence of Acridid. We found that the Acridid community depends mainly on the nature and richness of vegetation cover, the intensity of agricultural activity and the type of irrigation.

Keywords: Grasshopper, agricultural ecosystem, biodiversity, North Africa, Sahara

Introduction

Currently, the number of Orthoptera species described worldwide is about 28159 species (Eades et al. 2018). Acridoidea is the notable superfamily of the suborder Caelifera having 518 species (Yadav et al. 2017). Furthermore, 241 species of Orthoptera Acridomorpha Dirsh 1975 have been listed for Morocco, Algeria, Tunisia and the Western Sahara (Louveaux et al. 2013). Algeria has about 114 Orthoptera, including 98 Caeliferans. A significant part of Algeria is inhabited by grasshoppers. These insects are one of important pests since the onset of agriculture. They can cause great damage to agricultural production when climatic conditions are conductive to their multiplication. Therefore, it is necessary to have comprehensive knowledge of all grasshopper species that occur in an area (Benkenana & Harrat 2009). Given the danger of these Acridids, several studies have been carried out all over the world, including Algeria. We cite among others those of Chopard (1943), Dirsh (1965), Zergoun (1991, 1994), Doumandji-Mitiche et al. (2001), Ould El Hadj (2002), Maurel (2008), Benfekih et al. (2011), Guendouz-Benrima et al. (2011), Moussi et al. (2011, 2014, 2018), Louveaux et al. (2013), Massa (2013) and Defaut (2017). However, the grasshopper fauna of Algeria in general and of the Sahara in particular needs more studies; because the only species that are well-studied are the gregarious and economically important species such as the migratory locust Locusta migratoria (Linnaeus, 1758), the desert locust Schistocerca gregaria (Forskål, 1775), and the Moroccan locust Dociostaurus maroccanus (Thunberg 1815). The objectives of these investigations of the grasshoppers in agriculture ecosystems of Mzab valley

18 Accepted by Petr Kocarek: 7 Feb. 2019; published: 28 Feb. 2019

ACRIDID DIVERSITY IN AGRICULTURE ECOSYSTEMS J. Insect Biodiversity 009 (1) (c) 2019 Magnolia Press
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were to (i) investigate grasshopper species composition of three studied sites, (ii) found the abundance and diversity of grasshopper species in studied sites, (iii) demonstrate the effect of vegetation cover, the intensity of agricultural activity and the type of irrigation on the richness and diversity of Caelifera.

Materials and methods

Study area

The study was conducted in three different localities of the Mzab valley : N'tissa (Béni Isguen), Touzouz (Ghardaïa) and El-Djaoua (El-Atteuf). The Mzab valley is located in the Septentrional Sahara (northern Algerian Sahara). Its elevation is 483.63 m above mean sea level, between 32° to 33° north latitude and between 3° to 4° east longitude (Fig. 1).

Figure 1. Geographic location of the surveyed sites in the Mzab valley, Septentrional Sahara (Mzab valley, Ghardaïa province, Algeria).

The choice of studied sites is based on the variability in three criteria: vegetation cover, agricultural intensity and irrigation. At each study site, the floristic composition was determined by visual survey of a randomly placed quadrat of 500 m2. For each plant, the cover was estimated by calculating of the surface area occupied by the orthogonal projection of the aerial part, according to Duranton et al. (1982). Agricultural intensity was quantified by the percentage of agricultural crops. Humidity was quantified by the frequency and amount of irrigation. The study of the Acridid fauna was conducted in three different localities: N'tissa (Béni Isguen), Touzouz (Ghardaïa) and El-Djaoua (El-Atteuf).

Béni Isguen (S1): latitude 32° 44' North, and longitude 3° 65' East. The surface area of the site is about 2 hectares. The vegetal cover is mainly composed of date palms (Phoenix dactylifera), fruit trees (Citrus sinensis (L.) Osbeck and Vitis vinifera L.) and vegetable crops (Cucurbita siceraria (Molina) Standl., Cucurbita maxima Duchesne, Citrullus lanatus (Thunb.) Matsum. & Nakai and Cucumis melo L.) cover 25%. Also there was an abundance of weeds, 30% is covered by Cynodon dactylon (L.) Pers., Setaria verticillata (L.) P. Beauv, Polypogon monspeliensis (L) Desf., Hyparrhenia hirta (L) Stapf and Stipagrostis plumosa (L.) Munro ex Anderson. The intensity of agricultural activity is low; irrigation is irregular.

Ghardaïa (S2): latitude 32° 51' North and longitude 3° 59' East. The surface of the site is 6 hectares. Among the plants, we find date palm (Phoenix dactylifera L.), citrus fruits (Citrus sinensis (L.) Osbeck), olive trees (Olea europaea L.), and vegetable crops (Cucurbita siceraria (Molina) Standl., Cucurbita maxima Duchesne, Citrullus lanatus (Thunb.) Matsum. & Nakai and Cucumis melo L.) cover 50%. The Poaceae cover 15% (Cynodon dactylon (L.) Pers., Setaria verticillata (L.) P. Beauv and Polypogon monspeliensis (L) Desf.) and are located under the trees. The intensity of agricultural activity is average. Irrigation is normal and is done by drip system.

El-Atteuf (S3): latitude 32° 44' North, and longitude 3° 72' East. The surface of the site is 4 hectares. The vegetation is dominated by forage crops (Medicago sativa L. and Hordeum vulgare L.) which cover 50%, date palm (Phoenix dactylifera L.) and vines (Vitis vinifera L.) cover 30% and a few weeds (Cynodon dactylon (L.) Pers. and Setaria verticillata (L.) P. Beauv) cover 5%. The intensity of agricultural activity is strong. Irrigation is by sprinkler irrigation for forage crops (alfalfa and barley) and drip irrigation for other crops (fruit and vegetables).

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J. Insect Biodiversity 009 (1) (c) 2019 Magnolia Press ZERGOUN ET AL.

This Saharan region is characterized by a dry and cold climate in winter and dry, hot summers. Precipitation in the Mzab region are weak and irregular. The main rains occur in autumn. Rainfall is very rare during the whole year (less than 70 mm per annum). During summer, the air relative humidity falls to as low as 10%, resulting in a strong evaporation. However, in winter it rises up to 48.6%. In the Mzab region, there are two types of winds: north-west dominant sand-laden winds and south-north dominant hot and dry winds (named sirocco). The highest wind speeds occur in the month of April and are in the range of (14.4 m/s). The climatic data were collected over the period 2007- 2016 from the Office of Algerian meteorology. We used the ombrothermic diagram of Bagnouls & Gaussen (1953), which defines empirically a dry month as when the total precipitation (in mm) recorded during a month is lower or equal to double the average temperature (in °C) of the same month. For the area considered (2007-2016), the dry season lasts 12 months (Fig. 2).

Figure 2. Graphical representation of T°-P Ombrothermic diagram (2007-2016) in the Mzab valley, Ghardaïa province, Algeria (National Office of Meteorology, Ghardaïa).

Sampling and identification

Methods for Orthoptera sampling are numerous and very diverse (Lamotte & Bourlière 1978; Voisin 1980, 1986; Gillon 1974). In the present study, the selected method is that of quadrats, the most frequently reported method used for biodiversity studies of terrestrial ecosystems. The method used for surveying grasshoppers in this study is reported by Gardiner et al. (2002). The size of the quadrats used in the survey was 25 m2 (5×5 m). Ten quadrats were positioned at random in one hectare plot at each study site. The corners of each quadrat were marked using poles without the observer disturbing the grasshoppers within by casting shadows. Each plot at the study sites was surveyed to ascertain grasshopper abundance and species richness. Samplings were carried out very early in the morning, between 7:00 a.m. and 9:00 a.m. in summer, and between 9:00 a.m. and 11:00 a.m. in winter, when the insects were still immobilized on the ground. For each site, the quantitative measure of acridian density was carried out using the quadrats method. The total number of individuals belonging to each species was counted in quadrats (25 m2). The counting in the quadrats were repeated ten times for each site and day of sampling. Density from each site has been reported as grasshopper per 100 m2.

Acridid were collected by sweeping a hand net. A standard net of 40 cm in diameter having a depth of 60 cm with a 90 cm long wooden handle was used for sample collecting. Grasshoppers were collected in the different localities once a month in each ecosystem from January to December 2017. The collected specimens were killed using ethyl acetate and properly stretched, pinned and labeled. Specimen identification up to the species level was done with the aid of the taxonomic keys of Chopard (1943); Dirsh (1965), and also Catalogue and keys of the Acridomorpha (Insecta, Orthoptera) from North West Africa (Louveaux et al. 2018).

Data analysis

Relative abundance of grasshopper species was calculated as the number of individuals of species i relative to the total number of individuals of all species collected at each site.

Richness, number of individuals of species, species diversity and evenness (Shannon-Weiner index and evenness index) of the grasshopper community was evaluated with the help of statistical package Past 3.4 (Hammer et al. 2014). Species richness estimates with 95% confidence intervals for the estimators Sest (analytical) and Chao1 Classic based

ACRIDID DIVERSITY IN AGRICULTURE ECOSYSTEMS J. Insect Biodiversity 009 (1) (c) 2019 Magnolia Press
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on 100 randomized samples for the data of Caelifera sampled was applied. Estimated species richness was calculated with the program EstimateS 9.1.0 (Colwell 2013). An Analysis of Similarities ANOSIM (Clarke 1993) was used to test if the differences in structure and composition of grasshopper communities in each site and throughout the year were significant. The Bray-Curtis coefficient was used as a distance measure. We used Kruskal-Wallis non-parametric one-way analysis of variance to test for differences among the disturbance categories in grasshopper species richness and grasshopper diversity. Except as otherwise noted, all statistics were performed using R version 2.15.3 (R Core Team 2013). All data transformations and graphs were done in Excel. P values of 0.05 or smaller were interpreted as significant.

Results

During 36 surveys in the 3 studied sites of the Mzab valley, a total of 5255 Acridid specimens, representing 27 species were recorded in 360 quadrats. Identification of these specimens showed that 23 species belonged to the Acrididae, 3 to the Pyrgomorphidae and one to the Tetrigidae. Twenty three Acrididae grasshoppers could be separated into 6 subfamilies and 13 genera, while the 3 Pyrgomorphidae could be grouped into one subfamily and two genera. The Tetrigidae is represented by a single sub-family and a single genus. A list of short-horn grasshopper species found in the 3 studied habitats during our 12-month sampling period is presented in Table 1.

Table 1. Inventory and relative abundance (%) of short-horn grasshopper species of agriculture ecosystem in three localities from the Mzab valley (Septentrional Sahara, Algeria).

Family

Subfamily

Species

S1

S2

S3

Tetrigidae

Tetriginae

Paratettix meridionalis (Rambur, 1839)

0.57

-

1.18

Acrididae

Acridinae

Acrida turrita (Linnaeus, 1758)

4.21

2.23

4.27

 
 

Truxalis nasuta (Linnaeus, 1758)

1.42

0.56

-

 

Calliptaminae

Calliptamus barbarus (Costa, 1836)

0.06

-

-

 

Eremogryllinae

Notopleura saharica Krauss, 1902

0.62

0.45

-

 

Eyprepocnemidinae

Heteracris adspersa (Redtenbacher, 1889)

0.91

-

-

 
 

Heteracris annulosa Walker, 1870

3.69

3.44

3.22

 
 

Heteracris harterti (Bolivar, 1913)

1.48

-

-

 
 

Heteracris littoralis (Rambur, 1838)

1.82

1.01

1.44

 
 

Heteracris minuta (Uvarov, 1921)

0.51

-

-

 

Gomphocerinae

Ochrilidia filicornis (Krauss, 1902)

1.48

2.53

2.30

 
 

Ochrilidia geniculata (Bolivar, 1913)

3.58

4.51

9.39

 
 

Ochrilidia gracilis (Krauss, 1902)

17.68

16.82

15.96

 
 

Ochrilidia harterti (Bolivar, 1913)

4.77

9.88

-

 

Oedipodinae

Aiolopus puissanti Defaut, 2005

1.88

2.03

3.74

 
 

Aiolopus simulatrix (Walker, 1870)

2.67

1.47

2.56

 
 

Aiolopus strepens (Latreille, 1804)

12.28

7.87

10.84

 
 

Acrotylus longipes (Charpentier, 1845)

2.10

1.21

2.23

 
 

Acrotylus patruelis (Herrich-Schäffer, 1838)

13.13

14.89

14.32

 
 

Hilethera aeolopoides (Uvarov, 1922)

0.51

1.52

0.06

 
 

Locusta migratoria (Fabricius, 1781)

-

0.15

-

 
 

Morphacris fasciata (Thunberg, 1815)

9.61

10.43

15.11

 
 

Sphingoderus carinatus (Saussure, 1888)

-

0.56

-

 
 

Sphingonotus azurescens (Rambur, 1838)

-

1.11

-

Pyrgomorphidae

Pyrgomorphinae

Pyrgomorpha cognata Krauss, 1877

7.50

9.27

8.01

 
 

Pyrgomorpha conica (Olivier, 1791)

6.93

8.05

5.32

 
 

Tenuitarsus angustus (Blanchard, 1836)

0.57

-

-

03

08

27

100

100

100

S1: N'tissa (Béni Isguen), S2: Touzouz (Ghardaïa), S3: El-Djaoua (El-Atteuf).

Overall, the species found regularly in the surveys are also the most abundant in terms of frequency. Species richness was higher at sites 1 and 2 compared to site 3. Béni Isguen was the richest site with 24 species, followed by Ghardaïa (21 species). El-Atteuf was the least rich, with only 16 species.

Non-parametric methods for estimating the true species richness indicated that the inventory for each habitat was complete. The accumulation curves for all species stabilized by the 12th survey for the three sites (Fig. 3).

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Figure 3. Sample-base rarefaction curve (with 95% confidence intervals) for the estimated species richness in the three study areas (Béni Isguen, Ghardaïa and El-Atteuf).

Species richness was higher at sites 1 and 2 compared to site 3. At Béni Isguen and Ghardaïa, presence of adventitious vegetation has favored the increased total richness of Caelifera. Béni Isguen seemed to be the richest site with 24 species; it is followed by Ghardaïa with 21 species. El-Atteuf was the least rich, with only 16 species. The relative abundance of Acridid population among the localities surveyed (Table 1) indicated that populations of Acrotylus patruelis, Aiolopus strepens, Morphacris fasciata, Ochrilidia gracilis and Pyrgomorpha cognata, were dominant at the three localities with 16.86%, 14.14%, 11.51%, 10.20% and 8.31% frequencies, respectively. The Shannon-Wiener index confirms these results, with the highest value (2.65 bits) reported at Béni Isguen and the lowest value (2.43 bits) recorded at El-Atteuf (Table 2). In addition, El-Atteuf seems to be more balanced (E = 0.87), followed by Ghardaïa and Béni Isguen with 0.84 and 0.83, respectively. The diversity index shows a somewhat variable result in different seasons. The diversity index was highest between March and October and lowest during winter in the three sites studied. On the other hand, the equitability index was highest in winter. The lowest equitability index was observed during late summer and autumn.

Table 2. Different ecological indicators applied to short-horn grasshopper species of agriculture ecosystem in three localities from the Mzab valley (Septentrional Sahara, Algeria).

Months

Béni Isguen

 
 

Ghardaïa

 
 
 

El-Atteuf

 
 

S

N

H

E

S

N

H

E

S

N

H

E

January

11

39

2.23

0.84

8

30

1.94

0.87

9

34

1.96

0.79

February

16

97

2.42

0.70

10

97

2.14

0.85

9

45

2.18

0.92

March

17

138

2.51

0.72

12

162

2.10

0.68

9

89

1.97

0.80

April

21

163

2.69

0.70

14

102

2.24

0.67

11

115

2.18

0.80

May

19

150

2.60

0.70

13

63

2.26

0.74

14

120

2.28

0.70

June

19

149

2.71

0.79

15

221

2.28

0.65

15

165

2.44

0.76

July

20

162

2.50

0.61

17

412

2.46

0.69

16

266

2.55

0.80

August

21

149

2.54

1.61

20

323

2.50

0.61

15

247

2.41

0.74

September

20

228

2.63

0.69

20

268

2.51

0.61

13

261

2.20

0.69

October

20

198

2.64

0.70

17

170

2.54

0.75

13

96

2.24

0.72

November

11

116

2.03

0.69

15

113

2.25

0.63

11

76

2.18

0.80

December

11

70

1.97

0.65

7

13

1.82

0.88

4

8

1.32

0.93

Total

24

1759

2.65

0.83

21

1974

2.55

0.84

16

1522

2.43

0.87

S: Richness, N: number of individuals of species (adults and nymphs), H: Shannon-Weiner index, E: Evenness index.

The grouping of grasshopper responded differently to each site (ANOSIM = R: 0.1123, P = 0.006; Fig. 4). Pairwise comparisons showed that the grouping of grasshoppers on each agricultural ecosystem was significantly different. The community composition of grasshopper responded differently to each month (ANOSIM R = 0.152, P = 0.001; Fig. 5). Pairwise comparisons showed that the community composition of Acridid on each month was significantly different in each site.

There were not found any significant differences for Caelifera abundance between the three sites (Kruskal-Wallis test; x2 = 0.82153; df = 2; p-value = 0.6631). Contrarily, the Caelifera richness in the three agricultural ecosystems were

ACRIDID DIVERSITY IN AGRICULTURE ECOSYSTEMS J. Insect Biodiversity 009 (1) (c) 2019 Magnolia Press
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significantly different (Kruskal-Wallis test; x2 = 9.2183; df = 2; p-value = 0.00996). The same was true for the diversity between the three habitats (Kruskal-Wallis test, x2 = 7.4135, df = 2, p-value = 0.02456).

Total grasshopper density varied during the study period. Higher densities were reached in July (164.8 individuals/100 m2) at Ghardaïa and (106.4 individuals /100 m2) at El-Atteuf, for Béni Isguen the highest value is recorded in August (99.6 individuals /100 m2). Lower densities in December at El-Atteuf and Ghardaïa, with a grasshopper density of 3.2 individuals /100 m2 and 5.2 individuals /100 m2, respectively (Fig. 6).

Figure 4. Analysis of similarities (ANOSIM) plot showing dissimilarity between and within three sites. Bold horizontal bar in the box indicates median; bottom of the box indicates 25th percentile; top of the box indicates 75th percentile; whiskers extend to the most extreme data point, which is no more than the range (i.e. 1.5) times the interquartile range from the box; width of the bar is directly proportional to sample size.

Figure 5. Analysis of similarities (ANOSIM) plot showing dissimilarity between and within Months. Bold horizontal bar in the box indicates median; bottom of the box indicates 25th percentile; top of the box indicates 75th percentile; whiskers extend to the most extreme data point, which is no more than the range (i.e. 1.5) times the interquartile range from the box; width of the bar is directly proportional to sample size.

Figure 6. Monthly variation in grasshopper population density recorded during the study period in the three studied sites (Béni Isguen, Ghardaïa and El-Atteuf).

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Discussion

A total of 27 species of grasshoppers were collected from different habitats of the Mzab region. Louveaux & Ben Halima (1987) listed 140 Caelifera species for entire Algeria. In this case with 27 species, the Mzab valley is occupied by 19.28% of Algerian Caelifera-fauna. All the grasshoppers collected are classified under three families Acrididae, Pyrgomorphidae and Tetrigidae. Family Acrididae was dominant with 23 species of Acridids grouped under 13 genera and six subfamilies, amounting to 85.18% of collected species. The second most abundant family was Pyrgomorphidae with three genera and one subfamily, which contributed 11.11% (3 species) of the total species, while the Tetrigidae was represented by only one species (3.70%). The trend of numerical distribution of different grasshopper families recorded in the present study is similar to the observations of Belhadj et al. (2014), who also reported that Acridid grasshoppers were the most abundant group followed by Pyrgomorphidae and Tetrigidae in three sites at Ouargla oasis (Algeria). Similar to Zergoun (1994), the species Calliptamus barbarus has been reported only once in the Mzab valley, and has not been found since. Three species were recorded for the first time in Mzab valley: Hilethera aeolopoides and Morphacris fasciata (in the three studied sites), and Notopleura saharica (in Béni Isguen and Ghardaïa).

The differences between the three environments are related to three parameters including, in order of importance: the nature and richness of vegetation cover, intensity of agricultural activity, and type of irrigation. The abundance of grasshopper species (27) was highest in Béni Isguen (24) and Ghardaïa (21). The El-Atteuf studied site was represented by the smallest number of species (16). The presence of adventitious vegetation has favored the increased total grasshopper richness at Béni Isguen and Ghardaïa. According to Paulraj et al. (2009), grasses were the most common habitat for grasshoppers. This finding is consistent with our results. Gramnivorous grasshoppers such as Ochrilidia gracilis, Acrotylus patruelis and Morphacris fasciata were most abundant, probably because our sites are very rich in grasses. According to Jaulin (2009), the presence of an herbaceous layer is therefore essential for most species of Orthoptera encountered.

The majority of Acridid species were found in the herbaceous layer, such as Ochrilidia gracilis, Aiolopus strepens, Acrida turrita, Pyrgomorpha cognata and Morphacris fasciata, or naked soil such as Acrotylus patruelis, Tenuitarsus angustus and Sphingonotus azurescens, on which they find their food and can lay eggs. This is in agreement with the results obtained by Benjelloun et al. (2014). El-Atteuf site has a low rate of grass cover due to permanent agricultural work with sprinkling irrigation. All of these parameters mean that this environment is less populated by Acridid.

As more and more samples are examined, it becomes harder and harder to find species not already counted, so the slope of the curve gets less and less steep as the sampling effort continues. With the sampling effort performed at each habitat, all species accumulation curves reached an asymptotic phase (Fig. 3)

Upon analysis of sites based on species abundance matrix using Bray-Curtis dissimilarity, the agricultural ecosystem was significantly different. These could be due to the difference in habitat such as vegetation cover, agricultural intensity and micro-climate; point out that the vegetation cover especially Poaceae, seems to be a principal cue for grasshoppers to select a habitat. This is also mentioned by Zergoun (1994). Higher humidity and increased agricultural activity were associated with decrease in abundance of grasshoppers, while greater vegetation cover (especially of Poaceae), was correlated with higher abundance of Acridids. A similar pattern was observed in composition of grasshoppers in each month, which reinforces the argument that humidity, agricultural intensity as well as vegetation cover play an important role in determining grasshopper assemblages in the study area. These factors also playing role in the feeding sites selection by Acridids. On the other hand, Oedipodinae are only rarely observed on the vegetation. According to Otte (1984) Oedipodinae usually are abundant only in areas with relatively sparse ground cover.

We found, that habitat management had a significantly different influence on species diversity and species richness according to its intensity in studied agricultural ecosystems. A high diversity and richness of Caelifera indicates a very low level of agricultural activity. Contrary to what was expected, monthly abundance of grasshoppers did not vary significantly among the sites, although, total abundance tended to be higher in Ghardaïa and Béni Isguen than in El-Atteuf.

The value of the diversity index shows that the cultivated area of Béni Isguen is the most favorable for the development of many Caelifera. Indeed, it is characterized by an abundance of grasses such as Cynodon dactylon and Setaria verticillata. In addition, the low intensity of agricultural activity in this environment favors the presence of Acridid. The site El-Atteuf is less diversified; this is due to frequent weeding and therefore a low presence of grasses. In addition, sprinkler irrigation creates a humid microclimate which limits the development of grasshoppers. Equitability does not varying much in the three cultivated environments; it means that the species are equally distributed. It is noted that the El-Atteuf site has a slightly higher evenness than the other sites, despite its low diversity. These results are

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most consistent with (Frontier 1982) reasoning. According to this author, a community comprising a small number of relatively abundant species and the others are rare, appears to be less diversified than a community comprising in total the same number of species, but with more equitably distributed frequencies.

Densities were higher in disturbed sites El-Atteuf and Ghardaïa. Similar results were observed by Cigliano et al. (2002) in Benito Juarez County, Southern Pampas, Argentina where disturbed habitats resulted in increased overall grasshopper densities. The monthly variations in the densities show larger amplitudes at El-Atteuf than at Ghardaïa and Béni Isguen. Grasshopper total density was affected by seasonal variation. The greatest density was observed between April and October. The maximum densities were recorded in summer. Thus, weather variables explained the spatio-temporal variation of the grasshopper density. This result does not agree with those registered in the Southern Pampas, Argentina, where grasshopper density was not affected by seasonal precipitation and temperature (Wysiecki et al. 2011). On the other hand, densities tended to increase in hot and dry conditions in our study.

Grasshopper species richness and diversity in three sites vary significantly among the months. Contrary to what was expected, total abundance in month did not vary significantly.

The site El-Atteuf, which has undergone a strong agricultural activity during our sampling, is less rich than the other two sites. Weeding destroys a large part of the spontaneous plants such as Poaceae. In fact, the herbaceous layer strongly contributes to the species richness of Acridids. Thus, the agricultural activity has a negative impact on Acridids. In contrast, some species such as Oedipodinae, commonly abundant on bare soil can be favored in this area. Sprinkler irrigation has a negative effect on Acridid richness. Knowing that most Acridids prefer warm and sunny places, this type of irrigation will create a moist environment, and consequently, the exclusion of many species that will seek drier environments. Generally, agricultural practices such as irrigation and weeding (decline in Poaceae) often lead to the destruction of species habitats, and cause a general decline in Acridid biodiversity as reported by Tscharntke et al. (2005). It is recommended that similar studies be conducted on a large scale in other regions in order to fully assess the grasshopper fauna of the Algerian Sahara.

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