2.1. Results
2.1.1. Main leafhopper species identified over the period 2021–2022
The leafhopper species identified on cotton are divided into three genera (Table III). These are the Empoasca, Amrasca and Jacobiasca genera. The main species encountered are: Empoasca papayae (Oman, 1937), Jacobiasca lybica (Bergevin & Zanon, 1922), Empoasca facialis (Jacobi, 1912) and Amrasca biguttula (Shiraki, 1913).
Table III: Main leafhopper species identified on cotton during 2021-2022 season
Genus
|
Species
|
Relative proportion (%)
|
Empoasca
|
Empoasca papayae
|
4.3
|
Empoasca facialis
|
3.2
|
Jacobiasca
|
Jacobiasca lybica
|
43.5
|
Amrasca
|
Amrasca biguttula
|
48
|
2.1.2. Annual variation in overall species composition of leafhopper species
The proportion of species identified varied from one season to the next (Table IV). In 2021, the majority of species identified were J. lybica with an overall proportion of 84.7%. It was followed by E. papayae and E. facialis. Their average proportions were 8.7% and 5% respectively. In 2022, A. biguttula was the most abundant species, with an overall proportion of 96.6%. It was followed by J. lybica and E. facialis. Their overall proportions were 2.7% and 1.3% respectively. The species E. papayae was not found in the samples collected in 2022. The same applies to A. biguttula, which was not observed during the 2021 season.
Table IV Annual variation in the overall species composition of leafhoppers
Genus
|
Species
|
Overall proportion (%)
|
2021
|
2022
|
Empoasca
|
Empoasca papayae
|
8.7
|
0
|
Empoasca facialis
|
5
|
1.3
|
Jacobiasca
|
Jacobiasca lybica
|
84.3
|
2.7
|
Amrasca
|
Amrasca biguttula
|
0
|
96.6
|
|
Unidentified
|
2
|
0
|
2.1.3. Geographic variation in overall species composition
The proportions of different species varied among localities (Table IV). Two species were found to be in the majority: A. biguttula and J. lybica. Their proportions varied respectively from 45 to 50% and from 40 to 50% according to the localities. However, A. biguttula was the most abundant.
Table IV Overall geographical distribution of leafhoppers during the seasons 2021–2022
Locality
|
Amrasca biguttula
|
Empoasca papayae
|
Jacobiasca lybica
|
Empoasca facialis
|
Unidentified
|
Bouaké
|
45
|
0
|
41.7
|
13.3
|
0
|
Boundiali
|
46.7
|
8.3
|
40
|
3.3
|
1.7
|
Ferké
|
46.7
|
11.7
|
40
|
1.7
|
0
|
Kani
|
48.3
|
5
|
40
|
3.3
|
3.3
|
Kong
|
50
|
5
|
41.7
|
1.7
|
1.7
|
Korhogo
|
48.3
|
1.7
|
48.3
|
0
|
1.7
|
Madinani
|
46.7
|
1.7
|
45
|
5
|
1.7
|
Nambingué
|
50
|
5
|
43.3
|
1.7
|
0
|
Niakara
|
48.3
|
0
|
50
|
1.7
|
0
|
Tienko
|
50
|
5
|
45
|
0
|
0
|
2.1.4. Dynamics of geographic variation in species composition between 2021 and 2022
From 2021 to 2022, leafhopper species encountered on cotton varied by locality and crop year (Table V). However, in both crop years, three main species were identified. In 2021, J. lybica was the most dominant species with a proportion ranging from 73.3 to 96.7%. However, in 2022 it was reduced to a proportion ranging from 0 to 6.7% depending on the locality, making it the second most dominant species during this season. E. papayae increased from 0 to 23.3% in 2021, then decreased to 0% in 2022, depending on locality, making it an extinct species in this season. E. facialis was the minority species in 2021, with a proportion ranging from 0 to 16.7% and decreased in 2022 to a proportion ranging from 0 to 10% depending on the locality. It remained the minority species. However, this survey identified it in only two localities, including Bouaké and Boundiali. A. biguttula went from being absent from the samples collected in 2021 (0%) to the most abundant species in 2022, with a proportion ranging from 90 to 100%, depending on the locality.
Table V Geographic dynamics of leafhopper species by crop year
Locality
|
2021
|
2022
|
Empoasca papayae (%)
|
Unidentified (%)
|
Jacobiasca lybica (%)
|
Jacobiella fascialis (%)
|
Amrasca biguttula (%)
|
Jacobiasca lybica (%)
|
Jacobiella fascialis (%)
|
Bouaké
|
0
|
0
|
83.3
|
16.7
|
90
|
0
|
10
|
Boundiali
|
16.7
|
3.3
|
76.7
|
3.3
|
93.3
|
3.3
|
3.3
|
Ferké
|
23.3
|
0
|
73.3
|
3.3
|
93.3
|
6.7
|
0
|
Kani
|
10
|
6.7
|
76.7
|
6.7
|
96.7
|
3.3
|
0
|
Kong
|
10
|
3.3
|
83.3
|
3.3
|
100
|
0
|
0
|
Korhogo
|
3.3
|
3.3
|
93.3
|
0
|
96.7
|
3.3
|
0
|
Madinani
|
3.3
|
3.3
|
83.3
|
10
|
93.3
|
6.7
|
0
|
Nambingué
|
10
|
0
|
86.7
|
3.3
|
100
|
0
|
0
|
Niakara
|
0
|
0
|
96.7
|
3.3
|
96.7
|
3.3
|
0
|
Tienko
|
10
|
0
|
90
|
0
|
100
|
0
|
0
|
Annual proportion
|
8.7
|
2
|
84.3
|
5
|
96.6
|
2.7
|
1.3
|
2.1.5. Geographic variation in infestation levels
The levels of leafhopper damage varied among localities in the two crop years (Fig. 2). In the 2021 season, attacks were significant in the northern part of the cotton basin. However, they were particularly pronounced in the northeast of the cotton basin. During the 2022 season, attacks were widespread throughout the cotton basin. However, the highest infestations were observed in the northern part of the cotton basin, with higher levels in the northeastern part.
2.1.6. Seasonal and annual variation in infestation levels
Seasonal incidence of leafhoppers varied in both crop years (Fig. 3). In both years, attacks increased throughout the cropping season, but with higher levels in 2022.
Leafhopper attacks varied by year from 2012 to 2022 (Fig. 4). From 2012 to 2021, average attack levels ranged from 2.3 to 6.04 plants attacked per 30 plants observed. These levels were relatively low compared to those observed in 2022. Indeed, the average level of attacks was more than nine attacked plants per 30 plants observed in 2022. This level had never been reached in the previous ten years (Fig. 4).
2.1.7. Geographical variation in rainfall
Cumulative rainfall from June to July varied downwards from 2021 to 2022 (Fig. 5). During the 2021 season, cumulative rainfall varied between 290 mm and 530 mm, following the North-East/South-West gradient. In 2022, however, the cumulative rainfall varied between 260 mm and 410 mm, a drop ranging from 30 to 120 mm. In addition to the drop in rainfall, a geographical peculiarity can be observed in that the lowest values were recorded in the Tangrela-Mankono-Kong triangle, covering the North-Central-East zone, with an extreme of less than 260 mm recorded in Tengrela.
2.1.8. Geographical variation in relative humidity
In the cotton basin, average relative humidity from June to July generally varied downwards from south to north, from 2021 to 2022 (Fig. 6). Over the two crop years, it varied between 77% − 87% (2021) and 75% − 87% (2022), a decrease of 2% from Tengrela (North) to Bouaflé (South). However, beyond the 9th parallel, the 2021 season recorded average relative humidity levels varying between 77% and 80%, following a North-East/South-West gradient, while in 2022, the variation was between 74% (Tengrela) and 80%, with the exception of the Kong zone, which recorded an average humidity level of 82%.
2.1.9. Geographical temperature variation
From 2021 to 2022, the average temperature from June to July fell from 25°C to 27°C, following a north-east/south-west gradient (Fig. 7). Beyond the 9th parallel, there was a reduction in the high-temperature zone from 2021 to 2022. In 2021, the zone is dominated by an average temperature of over 26.6°C, compared with 26.2°C in 2022.
2.2. Discussion
Analysis of the annual and geographical variation of jassids revealed the cohabitation of at least three jassid species. This study highlights the dynamics of the leafhopper complex both between localities and between agricultural campaigns. It also showed a damage dynamic according to locality, crop year and dominant species. However, damage is more pronounced in the northeast of the cotton basin. This observation was also made by (Koné et al., 2017). Indeed, these authors showed that from 1993 to 2007, jassid damage was at a low level throughout the cotton basin. At that time, the jassid species known on cotton was the endemic E. facialis (Cauquil, 1986 ; Viot et al., 1999 ; Koné et al., 2017). Whereas from 2011 to 2015, damage was mainly evident in the northeast of the cotton basin. (Malanno et al., 2021) also confirmed the location of heavy infestations in the northeast of the cotton basin. Changes in the parasitic complex could partly explain this jassid outbreak.
During the 2021 cropping season, identifications revealed the arrival of two new jassid species. These were J. lybica and E. papayae. However, the first species proved to be the most dominant and invasive, occupying a larger proportion of the plant than the endemic species. It accounted for over 80% of all jassid species. The attacks observed during the 2020–2021 crop year could therefore be attributed mainly to J. lybica.
A change in the cotton pest complex was also observed in the 2022 season. It was marked by the arrival of a new species, identified as A. biguttula. Depending on the locality, it represented 90–100% of the leafhopper species. This species, more invasive and virulent, was the basis of the infestation levels observed. Indeed, in 2022, damage was more remarkable and widespread throughout the cotton basin, with levels higher than in 2021. Throughout the growing season, infestation levels were constantly above the economic damage threshold (ten plants attacked out of 30 observed). At some periods, levels reached 100% of plants. This species had a very significant negative impact on the producer's income.
The species and damage dynamics observed between localities, seasons and years could be due to climatic factors such as rainfall, temperature, and relative humidity. Indeed, from the 2021 crop year to the 2022 crop year, climatic conditions have varied. Average rainfall levels fell sharply across the cotton-growing basin, with an accentuation in the Northern zone, particularly in the North-Eastern part of the basin. The Northern zone also recorded the lowest relative humidity and the highest temperatures. (Ashfaq et al., 2010 ; Gogoi & Dutta, 2000) showed that low rainfall combined with high temperatures and low relative humidity contribute to the development of jassids. Studies carried out by Koné et al (2017) attest to the positive impact of low rainfall on jassid outbreaks. Indeed, these authors have shown that rainfall of less than 320 mm between June and July favors jassid outbreaks, leading to higher levels of infestation and damage. These authors also showed that the north-east of the cotton basin was particularly prone to low rainfall, with levels below 320 mm between June and July. The results obtained in this study corroborate these authors' findings. Indeed, the northern zone, particularly the north-east, recorded the lowest rainfall in both crop years. The damage levels accentuated in this part of the cotton basin could be attributed to climatic conditions that favor the pullulation and virulence of jassid attacks. Climatic factors therefore play an essential role in the composition of jassid populations (Gogoi and Dutta, 2000) and in the manifestation of damage. These dynamics could also be explained by migrations from host crops, cross-border migrations, and exchanges of plant material without phytosanitary controls.
Indeed, collections carried out in 2022 by (Kouadio, et al., 2022b) revealed the presence of J. lybica and A. biguttula species on okra and eggplant crops in proportions similar to those observed on cotton. These species can reproduce on several host plants in the same environment, notably eggplant and okra (Bernardo and Taylo, 1990; Kouadio et al., 2022b). A. biguttula was first reported in Africa in Ghana on okra as a minor pest (Obeng-Ofori and Sackey, 2003).
The appearance of these new species in the parasitic facies of cotton would explain the emergence of jassids in recent years. It would also explain the inadequacy of control programs observed in the field.