Lifecycle, Hosts and Importance of Some of the Major Cotton Insect Pests in South Africa: A Review

Received 13 May, 2024

Accepted 01 Jul, 2024

Published 31 Dec, 2024

Cotton (Gossypium hirsutum L.) is an essential fibre crop cultivated worldwide for centuries. The crop is a cash crop that is grown as raw material for textile industries in over 80 countries with tropical and subtropical climatic conditions. In Sub-Saharan Africa, cotton is a key export crop for global cotton lint exports. In South Africa, cotton is mostly grown by smallholder farmers in five provinces. However, several factors affect production, including low yields, high input costs and pest and weed infestations. The incidence of pests is a significant influence that affects cotton production, resulting in low yields and poor quality. The importance of cotton insect pests lies in their ability to cause significant economic losses by damaging cotton crops. Some are serious pests that require control measures such as synthetic pesticides to warrant a viable yield. A successful control strategy requires integrated pest management that prevents or suppresses damaging populations of insect pests by applying the comprehensive and coordinated integration of multiple and compatible control tactics, including chemical, cultural and biological methodologies. The lifecycles of insect pests can vary depending on factors such as environmental conditions, host availability and pest species. Understanding these lifecycles is crucial for implementing effective pest management strategies at different stages of development to minimize their impact on cotton crops. This paper seeks to review and examine the lifecycle, hosts and importance of Helicoverpa armigera, Aphis gossypii, Bemisia tabaci, Thrips tabaci, Jacobiella facialis, Tetranychus urticae and Dysdercus species as major key insect pests in cotton production.

INTRODUCTION

Although insects are the largest animal group and have a significant role in the ecosystem, some can be serious crop pests, including cotton¹. The incidence of pests in cotton significantly affects production, resulting in low yields and poor quality². Sub Saharan Africa has climate conditions suitable for various pests, which attack cotton and result in yield losses³. Tay et al.⁴ reported that an average of 30% of seed cotton is lost due to pests. Some insects are casual or random pests, but others may be economically important to cotton as they cause damage and reduce the yield. Insect pests such as the African bollworms⁵, aphids⁶, thrips⁷, whiteflies⁸ and leafhoppers⁹ are serious crop pests in South Africa. Pesticides are commonly used to control these pests; however, controlling one species can have an impact on the occurrence and population of other species.

All insects undergo their unique life cycles through complete or incomplete metamorphosis. A complete metamorphosis consists of four developmental stages, namely egg, larva, pupa and adult. While complete metamorphosis involves a pupal stage¹⁰, insects undergo incomplete metamorphosis from nymphs to adults without the pupal stage¹¹. Some insects have larvae and adults that are different in appearance from each other¹² and often live in various habitats. Because an insect’s host, behaviour and development can all drastically alter throughout its life cycle, depending on the type of life cycle, insect life cycles are essential for managing pests. Therefore, this review describes the key insect pest of cotton in South Africa.

Pest lifecycle, host and damage
African bollworm: The African bollworm, Helicoverpa armigera Hübner (Lepidoptera, Noctuidae), is the most significant pest of agriculture commonly found in Africa, Asia, Oceania, Europe¹³ and recently in South America¹⁴. In Africa, the pest is regarded as an indigenous species contributing to the reduction of crop production¹⁵. It is the only heliothine species of major economic importance¹⁶. In East Africa, H. armigera attacks various crops, including cotton, legumes, maize, sorghum, sunflower, tobacco and tomato¹⁷. In South Africa, cotton is one of the main crops attacked by the pest. It has been regarded as a serious pest due to feeding on a wide range of host crops⁴, high fecundity¹⁸, multivoltine life cycle¹⁹, great potential mobility²⁰, damage to fruiting parts²¹ and its resistance to chemical insecticides²². van Hamburg and Guest²³ listed 35 host crops of H. armigera plus 25 wild host plants in eastern and Southern Africa, while Krinski and Godoy²⁴ documented over 67 host families worldwide.

The H. armigera developmental cycle goes through four stages: Egg, larval, pupa and adult²⁵. There can be up to five generations of bollworm per year⁵ and each generation can take about four to six weeks²⁶. The H. armigera females prefer laying eggs on host flowers, squares and fruit²⁷ during winter or spring¹³. The eggs are laid over two to three days and oviposition may last up to eight days¹³. In South Africa, it can take up to 23 days for oviposition to occur⁵. Eggs are cream to white, changing to brown before hatching⁵ and up to 0.6 mm in diameter¹³. A single female may lay up to 1500 eggs²⁷. In South Africa, a single female moth can lay an average of 730 eggs⁵. Larval colour and size vary depending on the larval instar. The larvae range in colour from yellow to brown with a cylinder-shaped body and longitudinal stripes occur on the dorsal side²⁸. At the later stage, the larvae have white or yellow lines and white spiracles with black rims²⁹.

Pupae are round at both ends with a brown colour and are about 14-18 mm long 5. The pupal stage occurs in the soil²⁵ at a depth of 3-15 cm¹³. When the pest feeds on cotton, the pupal viability increases and the pupal stage lasts about 15 days²⁷. The moth is usually brown³⁰, with a broad thorax and seven to eight blackish spots on the forewings³¹. The hind wings have apical ends with a broad dark-brown border and yellow margins³². Helicoverpa armigera is a nocturnal pest³³ that can travel long distances due to a separation of feeding and oviposition by unfavourable habitats¹³. Long-range movement can also be attributed to migration, which may have implications for managing pests in the various agricultural landscapes³⁴. Daguang et al.³⁵ reported that most moths were distributed within a 720 m range when released in the field.

Helicoverpa armigera damage differs by crop and because of the migratory behaviour, the occurrence of this pest is frequently unpredictable³⁶. The pest causes damage of more than US$2 billion to crops every year and this excludes the environmental costs that are related to its control⁵. The damage caused by H. armigera depends, to some extent, on the adult population numbers, the number of eggs laid and the survival rate of larvae³⁷. Young larvae tend to feed on younger leaves, while the older ones feed on different parts of the plants, preferably the buds, flowers, fruits and pods³⁸. Although there may be low numbers in the field, the damage may significantly increase because the larvae tend to feed partly on one cotton ball and then move to another 5. The damaged bolls ultimately drop off the plant³⁹. When left untreated, H. armigera can cause up to 90% of boll damage in cotton. In Brazil, the pest was reported to reduce cotton yields by up to 80%⁴. In 2014, de Freitas Bueno and Sosa-Gómez⁴⁰ reported that Brazil had crop loss estimated at US$ 0.8 billion due to the pest’s damage.

Cotton aphid: The cotton aphid, Aphis gossypii Glover (Hemiptera, Aphididae), is an important agricultural pest due to its wide host range⁴¹. Ma et al.⁴² reported that the pest has over 900 hosts from 116 plant families. Although A. gossypii can significantly damage the crop through direct feeding, its main threat is the ability to transmit many plant viruses⁴³. Aphis gossypii populations are primarily high on cotton from mid to late season⁴⁴. Before genetically modified cotton was introduced, pesticides sprayed against Helicoverpa species-controlled aphids; however, the aphid populations increased thereafter, resulting in resistance to pesticide control⁴⁵.

Aphis gossypii is a tiny, soft-bodied insect with a pear-like shape and a pair of black cornicles⁴⁶. They are 1 to 2 mm long with relatively long antennae and legs⁴⁷. Aphids can be distinguished from other pests like mites through slower movement when disturbed⁴⁸. Body colour varies depending on the host plant and the biological state of the individual aphid⁴⁹. The nymphs vary in colour from yellow to green, black or brownish⁵⁰. They often have a dark head and thorax with a dark green abdomen⁵¹. The first instar has four antennal segments, while the second has five⁵². The third instar can be differentiated from the fourth instar by the absence of setae on the genital plate⁴⁶. The fourth-instar nymphs have developed wings, while adults are primarily wingless⁵³. Small yellow or white aphids are observed during unfavourable environmental periods and do not reproduce until conditions are favourable⁵⁴. During favourable conditions, larger green forms are produced⁵⁵. The aphids generally require at least two hosts and the primary host is for sexual reproduction, while the secondary host is for asexual reproduction⁵⁶. Females produce offspring that takes about a week to develop and moult four times to become reproductive adults⁵⁷. The reproductive period covers 20 days and the female can produce up to 80 offspring⁵⁸.

Aphids can cause severe loss in cotton production⁵⁹. They cause damage to the undersurface of the leaves and the stems by using sharp mouthparts and sucking the sap from the tissues⁶⁰. The leaves may produce insufficient chlorophyll, initiate curling and die prematurely⁶¹. Genetically modified cotton has been reported to not affect preference and colonization by aphids⁶². Besides the physical damage to the host, aphids have been widely reported to transmit various virus diseases⁶³. Moreover, aphids produce a sugary substance called honeydew that causes stickiness⁶⁴, interfering with the plant’s photosynthesis⁴⁷. When aphids feed on cotton plants, the honeydew drops onto the bolls, resulting in a sticky deposit on the fibre⁶⁵. Stickiness reduces the lint quality and results in substantial price penalties for the grower⁶⁶. It is a severe challenge during cotton ginning since it causes the lint to stick to machinery⁶⁷. Honeydew also exposes the leaves to sunburn, which results in secondary infections that inhibit the plant’s functions⁶⁸.

Whitefly: Whiteflies, Bemisia tabaci, Gennadius (Hemiptera, Aleyrodidae) are one of the most important agricultural and horticulture pests worldwide⁶⁹. Whiteflies are regulated quarantine species in Australia, Africa, China, the EU and the USA⁷⁰. Currently, 39 species differ in their host-plant range⁷¹ and they have more than 600 host-plant species⁷². This species reportedly transmits over 100 plant viruses⁷³.

Whiteflies undergo six developmental stages: Egg, four larval instars and adults⁷⁴. The average developmental stage in cotton takes approximately 17 to 29 days to complete. Perring et al.⁶⁹ reported that at a lower temperature of 15°C, the complete development could take up to 105 days compared to 14 days at 30°C. The eggs are approximately 0.2 mm long and elongated with a pale brown colour⁷⁵. They are laid singly or in group circles on the undersides of the leaf surface⁶⁹. Each female can lay between 60 and 300 eggs⁷⁶, which takes about five to nine days to hatch, depending on the host species and humidity⁷⁷. Of the four larval stages, the first and second larval instars are up to 0.6 mm long and the first larval instar is the only mobile larval stage⁷⁴. The fourth instar, known as the pupa, is 0.7 mm long, oval and lasts about six days⁷⁸.

The adult emerges from the pupal case and expands its wings before powdering itself with wax from glands on the abdomen⁷⁹. Adults are about 1 to 2 mm long with wings covered in white powder wax⁸⁰. Their body is white to slightly yellowish with seven segmented antennae and one sensorial cone on the third, sixth and seventh segments^81,82. The wings are kept above the body in a tent-like position⁸³ and up to 15 generations can occur annually⁷⁷. Onstad⁸⁴ stated that mating occurs several times from 12 hrs after emergence and the female may live up to 60 days while the male lives for a shorter period. Whitefly nymphs and adults can be easily identified in the crop⁸³.

Whiteflies can cause damage through phloem-feeding, excretion of honeydew and transmitting viruses such as cotton leaf curl virus⁸⁵. Whiteflies suck phloem-sap and cause damage to a wide range of crops, including cassava⁸⁶, cotton⁸⁷ and tomato⁸⁸. With their piercing-sucking mouthparts, they insert their stylets into the plant to feed on the phloem⁸⁹. While feeding on the plant, immature and adult stages excrete honeydew onto the leaf surface and fruit⁸³, causing discolouration of leaves and fruit deformations⁹⁰. Whiteflies have developed resistance due to the overuse of insecticides⁹¹.

Thrips: Thrips, Thrips tabaci, Lindeman (Thysanoptera, Thripidae) is a serious early-season pest of seedling cotton⁹² and vegetable crops⁹³ throughout the world. They are commonly one of the first insects found on cotton⁹⁴. Hull⁹⁵ reported that thrips feed on a wide host range, including 140 species from over 40 families of plants, while Varela and Fail⁹⁶ and Cook et al.⁹⁷ reported that there are several hundred host plants. Thrips may be found on weeds and flowers growing near cotton and migrating onto cotton plants⁹².

The life cycle of thrips has six stages: Egg, two larval stages, two pronymph stages and an adult stage⁸¹. The life cycle can take between 10 and over 30 days to complete, depending on the climate and the host plant⁹⁸. Reproduction is both asexual and sexual⁹⁹, producing both males and females from unfertilized eggs¹⁰⁰ and females from fertilized eggs⁹⁹. The eggs are small, shiny white¹⁰¹, 0.2 mm long and 0.08 mm wide¹⁰². They are laid individually inside the leaf tissues⁹⁹. A female can lay up to 100 eggs, which take up to six days to hatch¹⁰³. The first instar is semi-transparent and white, while the second instar is yellow⁹³. The larvae undergo two instar stages, which last up to 10 days¹⁰⁴. The pupae do not feed¹⁰⁵ and the pupal stage takes about four days to complete⁸¹. Adult females are about 1.2 mm long, while males are smaller than 0.7 mm long⁹³. The body colour varies from yellow to brown depending on temperature¹⁰⁶. Adults are very active with fringed and pale wings¹⁰⁷. The antennae have seven segments and the eyes are grey¹⁰⁸. Adults live up to 35 days⁹² and several generations can develop annually¹⁰⁹.

The host plant, temperature and humidity play a role in the development of thrips¹¹⁰. Adults may hibernate in field crops¹⁰⁶ and overwinter in the soil¹¹¹. Adults can fly long distances from immediate plant hosts¹¹² and flight occurs during daylight at low wind speeds¹¹³. Thrips found in nearby weeds migrate onto cotton plants¹¹⁴ and adults are attracted to white, blue and yellow colours¹¹⁵. It is sometimes impossible to control thrips with pesticides since the eggs are laid under leaf tissues¹¹⁶, the pupae are found in the soil or between the leaves¹⁰⁹ and some adults may avoid control by hiding in the inner leaf spaces¹⁰⁴.

During the seedling stage, cotton growth is slow, resulting in an attack by early-season insect pests such as thrips¹¹⁷. When feeding, thrips move from the lower to the upper parts of the cotton plant as the plants increase in size¹¹⁸. The feeding preference may be due to the pest trying to access younger leaves with thinner epidermis on the lower surface¹¹⁹. Thrips feed on leaves, young leaves and flower buds¹²⁰, causing the silvering of leaves due to the loss of chlorophyll⁹³. The silvery appearance occurs after the fluids in the cell are replaced by air⁹⁷. Both adults and larvae feed on plant epidermal cells’ contents, causing damage that results in 30-50% of lint yield⁹⁷. Damaged cells wrinkle and the leaves do not develop well, causing them to twist⁹⁷. The damage caused by thrips can also allow secondary infection by plant pathogens¹²¹. Attique and Ahmad¹²² reported that thrips and cotton leafhoppers cause almost 40% loss in seed cotton yield. Scouting for thrips is difficult; a lens may be required¹⁰⁴ and the population level can be determined by observing leaf damage¹²¹. Wardle and Simpson¹²³ reported no evidence of toxicity from the thrips salivary secretion. Thrips are also reported to be vectors of plant viruses¹²⁴. Hull⁹⁵ stated that 17 species are reported to transmit viruses from four plant virus groups, most feeding on the plant’s vegetative parts and pollen.

Leafhopper: Leafhoppers, Jacobiella facialis Jacobi (Hemiptera: Cicadellidae), commonly known as jassids, are one of the major cotton pests in Africa¹²⁵. The pest has a synonym called Empoasca facialis and it was described in Dutch East Africa in 1912. Leafhoppers are commonly found in the tropics and subtropics¹²⁶. Numerous species of leafhoppers are found on cotton¹²⁷ and can feed throughout the crop cycle despite pesticide application¹²⁵.

Leafhoppers reproduce sexually and the egg hatches to a nymph¹²⁶. The nymphs look like adults but are smaller with a pale yellow-green colour⁷⁵. Leafhoppers undergo five nymphal instars⁷⁵ and they are multivoltine with several generations every year¹²⁸. Eggs are laid on the underside of leaves and they can hatch in about ten days¹²⁹. They are elongated and range from 0.8 to 10 mm¹³⁰. Leafhopper species are almost similar in shape but vary in colour from green to yellowish-brown¹³¹. They overwinter as egg, adult, or immature forms and pass through several moults before becoming an adults¹³². Leafhoppers are generally found low in the canopy¹²⁹ and when disturbed, they hop fast⁷⁵. The nymphs and adults may feed on the aerial parts of the same plant¹³³ and the attack occurs throughout the crop production cycle¹³⁴. Both the nymphs and the adult suck the sap from the xylem and phloem tissues of the plant and young leaves from the lower surfaces¹³⁵. The damage caused by the leafhoppers is called “hopper burn” because of the brownish appearance of plants and it is a non-contagious disease¹³⁶. Hopper burn causes the edges of the leaves to curl downwards and change to yellow and then red before drying out and falling off the stem¹³⁷.

Premature reddening has been reported to be a characteristic reaction of the plant rather than the attack¹²⁷. Hopper burn occurs when there is an interaction between insect-feeding stimuli and plant responses¹³⁸. Heavy infestations can damage the canopy and impair cotton growth, causing a 40 to 100% reduction in the number of bolls¹³⁹. Prolonged feeding also results in the shedding of leaves, squares and young bolls, leading to significant yield losses¹⁴⁰. The pest damage levels vary under different climatic conditions and lower rainfall significantly increases the pest population¹⁴¹.

Spider mite: Spider mites, Tetranychus urticae Koch (Trombidiformes: Tetranychidae), are important cotton pests¹⁴². There are many controversial reports on the taxonomic placement of the two-spotted spider mite, with about 65 synonyms included under this species¹⁴³. While numerous spider mite species attack cotton worldwide¹⁴⁴, the two-spotted mite is one of the most common and important species¹⁴⁵. This species is an early-season pest that causes significant yield losses in cotton¹⁴⁶. The growth stages of spider mites differ from one species to another¹⁴⁷. However, their life cycle is short, with high fecundity and haploid-diploid sexes¹⁴⁸. Depending on temperature, spider mite development occurs in five to twenty

days and may have overlapping generations every year¹⁴⁹. The optimal reproduction usually occurs in seven days at a temperature above 30°C¹⁵⁰. Gunning and Easton¹⁵¹ reported average development periods of 27 days at 16°C, six days at 29°C for females and a slightly shorter time for males. The complete life cycle consists of the egg, larva, two nymphal or pupal stages and the adult⁷³. Females lay male eggs during asexual reproduction and in sexual reproduction, both female and male eggs are laid¹⁵². Female spider mites can lay over 100 eggs over 12 days¹⁴² and under optimal conditions, several hundred eggs are laid by each female¹⁴⁷. The eggs are oval, shiny, colourless, 0.08 mm long and 0.13 mm in diameter¹⁴².

The eggs are attached to a silk web¹⁵³ and their presence can be used to confirm when the plant damage is due to spider mites¹⁵⁴. The eggs hatch in three days into larvae that are 0.1 mm long¹⁴⁷ with a pale green colour and three pairs of legs¹⁴⁹. The larval stages are mainly dormant and only become active after the moulting to the nymphal stage¹⁵⁵. The larvae move slowly and develop into the nymphal stage within three days¹⁵⁶. The two nymphs are known as protonymph and deutonymph and they have darker markings and eight legs¹⁵⁷. Initially, the nymph is pale yellow-green and later turns darker green¹⁴². Adult females are 0.2-0.6 mm long, elongated with long hairs on the dorsal side of the body and translucent pale greenish-yellow to brown¹⁵⁸. Adult females live for about two to four weeks¹⁵⁹, while males can live up to nine days 149. The overwintering females are orange to orange-red¹⁶⁰.

Spider mites feed on the undersurface of the leaves¹⁶¹, where they remove the sap⁷³. Spider mites are mesophyll feeders because they pierce the leaf epidermis and feed mostly on mesophyll cells, affecting photosynthesis in the leaves of host plants¹⁶². The damaged leaves become grey or yellow and damage to the open flower results in brown colour and withering of the petals¹⁶³. Complete defoliation may occur when the pest is not controlled at higher population densities¹⁶⁴. Crop development is reduced in cotton by high infestation during the early developmental stage¹⁶¹. The spider mites also transmit several viruses, including potato virus Y, tobacco mosaic virus and tobacco ringspot virus¹⁶³.

Cotton stainer: Cotton stainers, Dysdercus species (Hemiptera: Pyrrhocoridae), are serious cotton pests¹⁶⁵. The Pyrrhocoridae is a small family comprising 33 genera and approximately 360 species worldwide¹⁶⁶. Eleven pest species are found in Africa¹⁶⁷, with four species occurring in South Africa. The Davus fasciatus, Danio nigrofasciatus and Dipturus intermedius are important in Africa cotton¹⁶⁸. Cotton stainers have a wide range of alternative hosts, including wild plants and various hibiscus species¹⁶⁹.

The cotton stainers have several generations a year and the complete life cycle may take one to three months, depending on temperature¹⁷⁰. Over 100 small pale eggs are laid and incubation can take up to two weeks¹⁷⁰. The female lays eggs in the soil or plant debris¹⁷¹. The emerging nymphs are initially red and after five moults, they have the same colours as adults but lack wings¹⁷². The last stages of nymphs and adults have long mouthparts used when feeding on the cotton seeds inside the bolls, while younger nymphs only feed on slightly opened bolls¹⁷³. Nymphs and adults are usually found in larger groups¹⁶⁵. The cotton stainers resemble assassin bugs; however, adult females are bigger than males¹⁷⁰. Adults are up to 2 cm long¹⁶⁸ with colours that vary from a red to an orange body and black stripe on the wings¹⁷². Adults are active during the daytime and can travel long distances¹⁷⁰.

Both adults and nymphs suck the sap from the seeds with piercing mouth parts causing physical damage and shedding of young bolls¹⁶⁷. While feeding on cotton, the pest also damages the fibres and affects the development of the bolls¹⁷⁴. The feeding on developing and mature cotton seeds negatively affects the quality of the seed and oil content¹⁷⁵. Cotton stainers attack cotton throughout the fruiting stage and transmit a fungus disease known as boll disease, resulting in hard bolls and stained lint¹⁷². Adults are found on cotton as early as when the first bolls open and can remain inside the boll until harvesting¹⁷⁰.

SIGNIFICANCE STATEMENT

Insect pests cause severe losses in cotton production in South Africa. A successful control strategy requires integrated pest management that prevents or suppresses damaging populations of insect pests by applying the comprehensive and coordinated integration of multiple and compatible control tactics. Understanding the biology and damage caused by these significant cotton insect pests is crucial to minimize their impact on cotton production. This paper provides an overview of the lifecycle, hosts and importance of Helicoverpa armigera, Aphis gossypii, Bemisia tabaci, Thrips tabaci, Jacobiella facialis, Tetranychus urticae and Dysdercus species as major key insect pests in cotton production. In conclusion, the future of understanding and managing cotton insect pests lies in integrating advanced scientific research with practical, sustainable agricultural practices.

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