Table 1 shows the effects of castor bean, tomato, potato, and cucumber leaves on certain life-history features of S. littoralis. Overall, feeding S. littoralis larvae on castor bean, tomato, potato, and cucumber leaves did not affect their duration. Duration of larvae fed on tomato leaves was shorter than that of larvae fed on castor bean, potato, and cucumber leaves by 10.93, 12.83, and 12.37%, respectively. Similarly, the tested host plants did not affect the duration of surviving pupae, with the shortest pupal duration (13.0 days) in case of castor bean. Duration of pupae surviving larvae reared on tomato, potato, and cucumber leaves was approximately similar to each other (15.2–15.4 days). Weight of full-grown larvae was significantly affected by host plants (P < 0.05), with the largest weight (650.8 mg) and smallest weight (300.7 mg) in case of feeding larvae on castor bean and cucumber, respectively. Larval weight in case of feeding on tomato and potato was approximately the same (450.5 and 450.9 mg, respectively. Adult emergence was dramatically decreased (P < 0.05) when larvae were reared on tomato, potato, and cucumber leaves (46.7, 40.2, and 23.8%, respectively) compared to that in case of rearing on castor bean (95.2%). Treatment with castor bean, tomato, potato, and cucumber leaves significantly affected adult longevity (P < 0.05), with the longest female longevity (7.1 days) and male longevity (6.1 days) in case of castor bean and potato leaves, respectively. The number of eggs deposited per female (fecundity) was also significantly affected by the types of host plants tested (P < 0.05). The highest number of eggs (480 eggs/female) and the lowest ones (170 eggs/female) were deposited by females surviving larvae fed on castor bean and tomato leaves, respectively. Fertility (egg-hatch percentage) was also significantly affected by host plants (P < 0.05). The highest egg-hatch percent (98.1%) and the lowest one (40%) were attained in case of females surviving larvae fed on castor bean and cucumber leaves, respectively.
Table 1
Life-history features of S. littoralis fed on castor bean, tomato, potato, and cucumber leaves
Host plants | Larval duration (days) | Pupal duration (days) | Larval weight (mg) | Adult emergence (%) | Adult male longevity (days) | Adult female longevity (days) | Fecundity (no. of eggs /♀) | Fertility (% egg-hatch) |
Castor bean | 18.3 ± 0.31a | 13.0 ± 0.32a | 650.8 ± 10.23a | 95.2 | 4.9 ± 0.51a | 7.1 ± 0.70a | 480.0 ± 10.41a | 98.1 |
Tomato | 16.3 ± 0.12a | 15.2 ± 0.21a | 450.5 ± 15.56b | 46.7 | 3.9 ± 0.32b | 6.2 ± 0.50a | 170.0 ± 3.90b | 70.0 |
Potato | 18.7 ± 0.41a | 15.4 ± 0.31a | 450.9 ± 13.68b | 40.2 | 6.1 ± 0.53c | 6.1 ± 0.50a | 180.0 ± 2.91b | 66.2 |
Cucumber | 18.6 ± 0.12a | 15.3 ± 0.22a | 300.7 ± 12.36c | 23.8 | 4.5 ± 0.41a | 4.5 ± 0.40b | 280.0 ± 3. 22c | 40.0 |
Data are presented as the mean ± SE. Means followed by different letters within columns for each variable are significantly different (P < 0.05) using Tuckey’s test. |
In accord with our findings, Mohamed et al. (2019) reported that total larval duration of S. littoralis fed on clover, cabbage, broad bean, sugar beet, and cotton leaves was 18.0, 19.0, 18.0, 25.2, and 22.6 days, respectively. Pupal duration was 14.4, 14.2, 15.2, 14.4, and 13.8 days, respectively. Adult emergence ranged from 95.58 to 89.0%. The highest number of eggs deposited per female in a descending order were as follows: broad bean > clover > cotton > sugar beet > cabbage leaves. In the same context, Ismail (2020) showed that larval duration of S. littoralis fed on clover, broad bean, and cabbage leaves was 18.6, 16.5, and 13.2 days, respectively. Pupal duration was 10.0, 9.7, and 9.1 days, respectively. The highest larval weight, adult emergence, number of eggs deposited per female, and egg-hatch percentage were recorded in case of cabbage leaves.
Soybean and cotton were found to be the most suitable hosts for development and oviposition of Spodoptera eridania and Spodoptera cosmioides compared to oat, wheat, and maize (da Silva et al. 2017a). Similarly, compared to tobacco, Chinese cabbage, and sweet potato, cowpea was found to be the most suitable host plant for development of S. litura based on the shortest developmental times of larvae and pupae. The lowest pupation rate was on sweet potato. The highest number of eggs deposited were on sweet potato, followed by those on cowpea, Chinese cabbage and tobacco (Xu et al., 2010). The highest survival rate and the shortest larval duration of Spodoptera exigua were on asparagus lettuce, whereas the lowest survival rate and the longest larval duration were on sweet peppers (Zhang et al. 2021). Effects of host plants on developmental times, survivability, and reproductive potential of Spodoptera frugiperda have been reported by several authors (da Silva et al. 2017b; Diédhiou et al. 2021; Maharani1 et al. 2021; Al-Ayat et al. 2022; Altaf, et al. 2022; Gopalakrishnan 2022; Nandhini et al. 2023).
Table 1 shows, in general, a positive correlation between weight of full-grown larvae and fecundity and fertility of S. littoralis. Greenberg et al. (2001) reported that at each host plant treatment, fecundity increased with increasing pupal weight. Syed and Abro (2003) found a significant relationship between fecundity and pupal weight of Plutella xylosiella surviving larvae fed on different host plants. Shahout et al. (2011) demonstrated that weight and length of ovaries of Spodoptera litura were affected by the host plants.
Overall, castor bean leaves were the most nutritive host plant as it contained the highest levels of nitrogen and phosphorous. In contrast, tomato leaves were the least nutritive host as it contained the lowest contents of nitrogen, phosphorous, and potassium (Fig. 1). Nitrogen is an essential element in plants due to its key role in the production of primary and secondary metabolites important in plant physiology impacting performance and tolerance to environmental stresses (Muñoz-Huerta et al. 2013). Quality and quantity of food consumed by insect species directly influence their host preferences and affect their biological, physiological, and behavioral characteristics (Nation 2002; Golizadeh et al. 2009; Cabezas et al. 2013). It is known that the lepidopteran larvae fed on high-nutrient food obtain a faster growth rate and possess a short life cycle when compared to those fed on low-nutrient food (Hwang et al. 2008). Several authors reported the importance of allelochemicals as deterrents, repellents, and stimulants present in plants (Berenbaum and Neal 1985; Vendramim and Guzzo 2009). Host-plant adaptation also can be viewed in terms of physiological, biochemical, and evolutionary adaptations to host-plant allelochemicals (Slansky 1992; Sorensen and Dearing 2006). Tomato leaves have been shown to contain significant amounts of important allelochemicals involved in tomato plant protection against abiotic and biotic stresses (Nguenang et al. 2020; Tam et al. 2021; Bazzarelli et al. 2022). Ismail (2020) attributed the unsuitability of tomato leaves to developing and reproducing S. littoralis compared to clover, broad bean, and cabbage leaves to lower nutrient contents, including nitrogen, proteins, and carbohydrates. These findings may explain the highest weight of larvae, number of eggs deposited per female, and egg-hatch percentage in case of feeding S. littoralis larvae on castor bean leaves, and meanwhile may explain the lowest number of eggs deposited per female and egg-hatch percentage of larvae fed on tomato leaves.
The present study revealed that nutritional indices of S. littoralis larvae were dependent not only on the type of host plant and metamorphed instar, but also on the age within instar. Overall, during the growth of 4th, 5th, and 6th - instar larvae of S. littoralis, the lowest food consumed was attained in case of feeding on tomato leaves. In insects, poor nutrition during development typically leads to undersized adults (Scriber and Slansky 1981; Awmack and Leather 2002; Colasurdo et al. 2009; Dmitriew and Rowe 2011). Entire 6th -instar larvae fed on castor bean leaves consumed higher food compared to those fed on tomato, potato, and cucumber leaves (Table 2).
Table 2
Consumption index (CI) of S. littoralis larvae fed on castor bean, tomato, potato, and cucumber leaves
Instars | Age of instars (days) | Castor bean | Tomato | Potato | Cucumber |
CI (mg food ingested/mg body wt/day) |
4th | 1 | 0.7 ± 0.05a | 1.5 ± 0.05b | 2.2 ± 0.05c | 1.0 ± 0.05d |
2 | 0.3 ± 0.05a | 0.8 ± 0.01b | 1.1 ± 0.05b | 0.6 ± 0.05c |
5th | 1 | 2.0 ± 0.50a | 0.9 ± 0.05b | 4.0 ± 0.60c | 1.5 ± 0.05d |
2 | 1.0 ± 0.05a | 0.4 ± 0.05b | 2.0 ± 0.50c | 0.8 ± 0.05a |
6th | 1 | 2.7 ± 0.05a | 0.9 ± 0.05b | 1.6 ± 0.05c | 1.3 ± 0.05c |
2 | 1.3 ± 0.05a | 0.4 ± 0.05b | 0.8 ± 0.05c | 0.7 ± 0.05c |
3 | 0.9 ± 0.05a | 0.3 ± 0.01b | 0.5 ± 0.05c | 0.4 ± 0.04c |
Data are presented as the mean ± SE. Means followed by different letters within rows for each age are significantly different (P < 0.05) using Tuckey’s test. |
The highest RGR, particularly during the entire 6th -instar larvae was in case of feeding on castor bean leaves, whereas the lowest RGR was in case of feeding on tomato, potato, and cucumber leaves. The RGR in case of tomato, potato, and cucumber leaves was approximately the same during the entire 6th -instar larvae (Table 3).
Table 3
Relative growth rate (RGR) of S. littoralis larvae fed on castor bean, tomato, potato, and cucumber leaves
Instars | Age of instars (days) | Castor bean | Tomato | Potato | Cucumber |
RGR (mg wt gain/mg body wt/day) |
4th | 1 | 3.7 ± 0.06a | 2.0 ± 0.50b | 1.1 ± 0.05c | 1.3 ± 0.05c |
2 | 0.6 ± 0.03a | 1.0 ± 0.02b | 0.6 ± 0.03c | 0.7 ± 0.05c |
5th | 1 | 3.1 ± 0.05a | 1.0 ± 0.02b | 4.5 ± 0.60c | 1.5 ± 0.05d |
2 | 1.6 ± 0.05a | 0.5 ± 0.01b | 2.3 ± 0.05c | 0.8 ± 0.05d |
6th | 1 | 3.3 ± 0.06a | 1.2 ± 0.05b | 1.2 ± 0.05b | 1.3 ± 0.01b |
2 | 1.7 ± 0.05a | 0.6 ± 0.02b | 0.6 ± 0.05b | 0.6 ± 0.01b |
3 | 1.1 ± 0.05a | 0.4 ± 0.01b | 0.4 ± 0.05b | 0.4 ± 0.02b |
Data are presented as the mean ± SE. Means followed by different letters within rows for each age are significantly different (P < 0.05) using Tuckey’s test. |
During the entire 4th or 5th -instar larvae, the AD of larvae fed on castor bean, tomato, potato, and cucumber leaves was approximately the same. However, during the entire 6th -instar larvae, the highest AD was obtained in case of feeding on potato leaves, followed by castor bean, tomato, and cucumber leaves (Table 4).
Table 4
Approximate digestibility (AD) of S. littoralis larvae fed on castor bean, tomato, potato, and cucumber leaves
Instars | Age of instars (days) | Castor bean | Tomato | Potato | Cucumber |
AD (%) |
4th | 1 | 8.8 ± 0.06a | 9.7 ± 0.05a | 18.6 ± 0.10b | 9.6 ± 0.05a |
2 | 8.3 ± 0.10a | 9.4 ± 0.05a | 18.7 ± 0.10b | 9.4 ± 0.05a |
5th | 1 | 38.3 ± 0.10a | 23.5 ± 0.20b | 41.6 ± 0.20c | 19.7 ± 0.06b |
2 | 38.2 ± 0.10a | 23.4 ± 0.20b | 41.5 ± 0.20c | 20.2 ± 0.06b |
6th | 1 | 52.7 ± 0.10a | 37.6 ± 0.20a | 87.1 ± 0.20b | 31.6 ± 0.10ab |
2 | 52.6 ± 0.10a | 37.4 ± 0.20a | 87.3 ± 0.20b | 31.5 ± 0.10ac |
3 | 52.3 ± 0.10a | 37.3 ± 0.20a | 86.9 ± 0.20b | 31.5 ± 0.10ac |
Data are presented as the mean ± SE. Means followed by different letters within rows for each age are significantly different (P < 0.05) using Tuckey’s test. |
Change in the ECI and ECD values reflects the change in conversion rates of digested and ingested food into body mass, respectively. In the present investigation, 1-day-old 4th -instar larvae of S. littoralis showed the highest ECI in case of feeding on tomato and cucumber leaves, whereas the lowest ECI was in case of feeding on castor bean and potato leaves. During the entire 5th -instar larvae, the highest ECI was in case of castor bean, whereas the lowest one was in case of feeding on cucumber leaves. In the last instar larvae (i.e., 6th -instar larvae), the highest ECI was in case of feeding on castor bean leaves, followed by tomato, potato, and cucumber leaves (Table 5). The highest ECD during the entire 6th -instar larvae was obtained in case of feeding on castor bean leaves, followed by tomato, potato, and cucumber leaves, a pattern which is similar to that of ECI. During the entire 4th or 5th -instar larvae, the highest ECD was in case of castor bean leaves. During 4th -instar larvae, the lowest ECD was in case of feeding on potato leaves, and the lowest one during the entire 5th -instar larvae was in case of cucumber leaves (Table 6). Timmins and Reynolds (1992) attributed reduction in the efficiency of food utilization to increased energetic costs arising from a reduced ability to utilize dietary nitrogen, which would not necessarily interfere with absorption from the gut.
Table 5
Efficiency of conversion of ingested food to biomass (ECI) of S. littoralis larvae fed on castor bean, tomato, potato, and cucumber leaves
Instars | Age of instars (days) | Castor bean | Tomato | Potato | Cucumber |
ECI (%) |
4th | 1 | 2.2 ± 0.06a | 4.4 ± 0.05b | 2.4 ± 0.01a | 4.3 ± 0.05b |
2 | 9.6 ± 0.06a | 4.4 ± 0.05b | 2.6 ± 0.01c | 3.9 ± 0.05b |
5th | 1 | 8.6 ± 0.05a | 4.4 ± 0.05b | 5.9 ± 0.02c | 3.3 ± 0.06d |
2 | 8.9 ± 0.05a | 5.0 ± 0.05b | 6.1 ± 0.02b | 3.4 ± 0.06c |
6th | 1 | 8.1 ± 0.06a | 6.3 ± 0.06b | 4.7 ± 0.02c | 3.8 ± 0.05d |
2 | 8.7 ± 0.06a | 6.3 ± 0.06b | 4.9 ± 0.02c | 3.6 ± 0.05d |
3 | 8.1 ± 0.05a | 5.8 ± 0.06b | 5.0 ± 0.02b | 4.0 ± 0.05b |
Data are presented as the mean ± SE. Means followed by different letters within rows for each age are significantly different (P < 0.05) using Tuckey’s test. |
Table 6
Efficiency of conversion of digested food to biomass (ECD) of S. littoralis larvae fed on castor bean, tomato, potato, and cucumber leaves
Instars | Age of instars (days) | Castor bean | Tomato | Potato | Cucumber |
ECD (%) |
4th | 1 | 3.8 ± 0.06a | 3.4 ± 0.05a | 0.5 ± 0.01b | 3.1 ± 0.03a |
2 | 6.0 ± 0.06a | 2.8 ± 0.05b | 1.0 ± 0.01c | 2.8 ± 0.03b |
5th | 1 | 4.8 ± 0.05a | 1.9 ± 0.05b | 2.2 ± 0.02c | 1.0 ± 0.03d |
2 | 3.7 ± 0.05a | 1.5 ± 0.05b | 1.7 ± 0.02b | 0.7 ± 0.02c |
6th | 1 | 3.0 ± 0.06a | 2.3 ± 0.03b | 1.3 ± 0.02c | 0.6 ± 0.02d |
2 | 2.3 ± 0.06a | 1.3 ± 0.03b | 0.9 ± 0.02c | 0.2 ± 0.01a |
3 | 2.0 ± 0.05a | 1.1 ± 0.03b | 0.7 ± 0.02c | 0.1 ± 0.01d |
Data are presented as the mean ± SE. Means followed by different letters within rows for each age are significantly different (P < 0.05) using Tuckey’s test. |
In 6th -instar larvae of S. littoralis fed on castor bean leaves, the highest CI (Table 2) was correlated with the highest RGR (Table 3), ECI (Table 5), and ECD (Table 6). This explains the highest weight of full-grown larvae fed on castor bean leaves (Table 1). The lowest gross food utilization efficiencies (ECI and ECD) of full-grown larvae fed on cucumber leaves (Tables 5 & 6) and the lowest CI (Table 2) also explain the lowest weight of full-grown larvae fed on cucumber leaves (Table 1). The highest values of AD of full-grown larvae fed on the four tested host plants (Table 4) may have been an attempt by larvae to compensate the lowest rate of food consumption (Table 2), resulting in reduced fecal production (Slansky and Scriber 1985).
In accord with our results, Gacemi1 et al. (2019) demonstrated that the highest food utilization efficiencies (ECI and ECD) of 5th -instar larvae of S. littoralis were on artichoke and the lowest were on cabbage, whereas, the highest CI, AD, and RGR were on cabbage. Similarly, Ismail et al. (2020) recorded that the highest CI of S. littoralis larvae was on cabbage leaves, followed by broad bean and clover, with the highest RGR and AD on broad bean and cabbage leaves, respectively. Mousavi et al. (2023) showed that the lowest and the highest AD values of S. littoralis larvae were on basil and purslane, respectively. The highest values of food utilization efficiencies (ECI and ECD) were on chives and coriander, respectively, whereas the lowest food utilization efficiencies and RGR were on purslane.
Jooyandeh (2018) showed that Sivand and Super Queen tomato cultivars were unsuitable hosts for feeding Helicoverpa armigera among the ten tested cultivars based on the nutritional performance of 4th, 5th, and 6th -instar larvae. Similarly, nutritional indices of 3rd, 4th, and 5th -instar larvae of Pieris brassicae were studied by Mehrkhou et al. (2013) using different cabbage cultivars. They showed that white cabbage was the most nutritive and could successfully support maximum values of RGR and food utilization efficiencies among the tested cabbage cultivars. In contrast, cauliflower was the least nutritive one. Xue et al. (2010) observed that CI of S. litura larvae was highest on sweet potato, followed by that on cowpea, Chinese cabbage, and lowest on tobacco. In contrast, larvae fed on tobacco showed the highest food utilization efficiencies and lowest AD compared with those fed on Chinese cabbage, cowpea, and sweet potato. Zhu et al. (2005) found that S. litura larvae had lower RGR, CI, and AD on banana leaves, although they had a significantly higher food utilization efficiencies, indicating that the larvae were capable of compensating by more efficiently utilizing their limited banana leaf tissues than other host plants. The effects of host plants on the nutritional indices of S. frugiperda have been reported by several authors (da Silva et al. 2017b; Bavisa et al. 2021; Al-Ayat et al. 2022; Nandhini et al. 2023).