White mango scale damage on leaves and fruit in farmer orchards in Pakistan. (
) Chlorotic spots on fruits due to WMS infestation in Pakistan.
5. Life Cycle
White mango scale (WMS) tiny-shelled insects have more than 300 species
[23][66][41,80]. For mass rearing of this pest, optimum growth conditions were 25 °C and 70% relative humidity, respectively
[67][81]. However, 24–35 °C and relative humidity of 70–95% have been regarded as ideal environmental conditions for an increase in the population of WMS in field conditions
[68][82]. Both types of reproduction, sexual and asexual, were observed in WMS
[69][83].
5.1. Adult
The adult female is similar to nymphs without legs and wings
[70][84]; a circular scale made up of wax 2 mm in diameter, having three longitudinal ridges and an exuviae terminal covering the body
[37][71][72][38,54,85]. The exposed body of the gravid female is 1.5–2.0 mm long and brownish in color. Both forms of reproduction, ovipary and vivipary, were observed in scale insects
[73][86].
The adult male WMS is usually small, slender, and winged
[74][87]. Males bear vestigial mouthparts, hence are short-lived. Adult males were yellow to orange colored, about 0.53 mm long, and were unable to feed due to vestigial mouthparts. Adult males soon after emergence mate and die within 1–2 days
[35][52]. Adult females excrete sex pheromones to attract male-scale insects
[75][88]. Adult WMS vary in size (1.5–25 mm), shape and color
[72][76][85,89]. Males usually cluster around females, while females usually occur singly
[62][76].
5.2. Eggs
A female lays 80–200 eggs
[77][90]. The eggs are 0.17 mm long, oval, and initially reddish brown in color, which later on become purple-colored depending upon maturity
[76][89]. However, egg-laying fecundity is dependent upon the weather conditions as well, as in Australia, 50 eggs per female
A. tubercularis were recorded. In South Africa, during summer, spring and winter conditions, about 203, 261 and 82 eggs per female were recorded, respectively. However, under semi-field conditions (27.5 °C and 81% R.H, 65), Gutierrez
[70][84] observed 98.55 eggs per female
[35][52]. At 27 °C and 81% RH, the incubation period was 8 days
[70][84].
5.3. Hatching
After fertilization, the eggs hatch in 8 days
[28][45]. Oviparous and viviparous reproduction was observed in scale insects population
[58][73]. In sexual dimorphism, the female lays eggs. From these eggs, nymphs develop. Four stages (nymph stage 1, nymph stage 2, pre-pupa and pupa) were observed in the male population, while there are two female instars (nymph stage 1 and nymph stage 2)
[35][71][38,52] in the female WMS population.
5.4. 1st Instar
The first instar nymphs emerge from eggs, settle down onto the tender part of the plant and suck the plant nutrients. Newly emerged first instar nymphs settle down within 24 h after hatching. Magsig-Castillo et al.
[78][91] described that to find a good place for a feeding site, the first instar nymph can travel a distance of less than one meter. Once occupying some specific place, they insert their stylets, which ultimately form a food canal within the plant parts; hence they suck the sap from areas of colonization, either leaves, fruits or developing tissues
[79][92]. After that, filaments of thread made up of wax were produced, which ultimately covered the upper epidermis
[72][85]. Female crawlers often uniformly distribute within plant parts, while male crawlers settle near female crawlers in the form of groups. Although instar nymphs settle in groups, their population can be dispersed by various factors
[33][80][50,93]. First, the instar male WMS colonizes near the adult female
[70][72][84,85]. A study showed that about 10–80 males group near emerging adult female insects.
Nymphal instars and the male adults can move
[35][52], but the movement of female crawlers through wings, bird claws or any other means is very important to initiate the infestation in a new tree or orchard
[81][94]. In winter (7 and 23 °C), spring (13 and 26 °C), and summer (18 and 29 °C), the female first instar stage may last from 11.1–17.1 days
[35][52] while at 27 °C and 81% R.H, the first female and male instar last 10 and 9 days, respectively
[70][84]. In further development, about 80% of crawlers become males
[77][90].
5.5. 2nd Instar
The second instar female WMS varies in size. In female WMS, the scale developed on the epidermis of the WMS was 3–4 mm, rectangular and developed from waxy filaments. The second instar female antennae were ovoid, translucent yellow colored and bear very small antennae
[71][38]. The second instar male develops under the scale protective sheath bearing three longitudinal ridges
[71][38]. In winter, spring and summer simulated conditions, the duration of the second female instar ranges from 11.1–25.3 days
[35][52], while at 27 °C and 81% relative humidity, the second female and male instars may last for 5–8 days, respectively
[70][84].
5.6. Pre-Pupa to Pupa
No change takes place in the size of WMS males from pre-pupae to pupae
[71][38]. The pre-pupa and pupa stages may last for 3–5 days, respectively
[70][84]. The pupal stage is found only in the case of males.
Life cycle period:
Environmental conditions and climate affect the life cycle of WMS
[19][20]. In Australia, during summer, the WMS life cycle is completed in 35–40 days. In Winter, the life cycle is completed in 70–85 days
[52][82][67,95]. A Female’s WMS completes life in 52 days, while a male’s only lasts 36 days
[52][67]. The life cycle of WMS was completed in 68.9 in winter, 52.5 in spring and 42.7 days in summer
[35][52].
This pest has 3–4 overlapping generations in a year
[31][83][48,96]. There may be 5–6 generations per year, at 26 °C daytime temperature and 13 °C nocturnal temperature
[55][70]. WMS has three generations in Mexico
[23][51][41,66] and Egypt
[31][40][53][28,48,68]. This is an important pest of mangoes in Egypt
[84][97].
In Spain, it has three–four overlapping generations in a year
[19][71][85][20,38,98], and in Southern Spain, it has two generations (spring and autumn)
[71][38]. The ecological studies on pest resting behavior revealed that pest colonizes on the south sides of the tree in two locations (Kaapmuiden and Nelspruit, South Africa)
[86][34].
5.7. Feeding Mechanism
In the WMS population, the adult male insects have vestigial mouthparts and hence live for only a few hours. A female WMS normally feeds and lives longer
[87][99]. A male, after emergence, mates and dies within 1–2 days. WMS has piercing and sucking mouthparts. This chitinous tube is composed of four stylets, two maxillae and two mandibles
[79][92]. During feeding, the female WMS obtains nutrients by pushing mouthparts into the parenchymatous tissues
[88][89][90][91][100,101,102,103].
Histological studies show that the WMS scratches the interior of leaf tissue, including vascular bundles
[92][104]. WMS, during feeding, not only punctures the parenchymatous tissues, but the lignified materials of the xylem are also punctured to obtain food. The pest secretes phenolic acid, which leaves a reddish scar
[79][92].