Editing Scale insect (section)

==Life cycle==
[[File:EB1911 Hemiptera - Fig. 13.—Apple Scale Insect (Mytilaspis pomorum).jpg|thumb|Life-cycle of the [[apple]] scale, ''Mytilaspis pomorum''. a) underside of scale showing female and eggs, x24 b)&nbsp;scale upperside, x24 c) female scales on twig d) male scale, x12 e) male scales on twig]]

Female scale insects in more advanced families develop from the egg through a first [[instar]] (crawler) stage and a second instar stage before becoming adult. In more primitive families there is an additional instar stage. Males pass through a first and second instar stage, a pre-pupal and a pupal stage before adulthood (actually a pseudopupa, as only [[Holometabolism|holometabolous]] insects have a true pupa).<ref name=Capinera/>

The first instars of most species of scale insects emerge from the egg with functional legs, and are informally called "crawlers". They immediately crawl around in search of a suitable spot to settle down and feed. In some species they delay settling down either until they are starving, or until they have been blown away by wind onto what presumably is another plant, where they may establish a new colony. There are many variations on such themes, such as scale insects that are associated with species of ants that act as herders and carry the young ones to protected sites to feed. In either case, many such species of crawlers, when they moult, lose the use of their legs if they are female, and stay put for life. Only the males retain legs, and in some species wings, and use them in seeking females. To do this they usually walk, as their ability to fly is limited, but they may get carried to new locations by the wind.<ref name=Capinera/>

[[File:EB1911 Hemiptera - Fig. 12.—Apple Scale Insect.jpg|thumb|left|Apple scale. a) male, with legs and wings b) foot of male c) larva, x20 d) antenna of larva e) immobile female (removed from scale)]]

Adult females of the families Margarodidae, Ortheziidae and Pseudococcidae are mobile and can move to other parts of the host plant or even adjoining plants, but the mobile period is limited to a short period between moults. Some of these overwinter in crevices in the bark or among plant litter, moving in spring to tender young growth. However, the majority of female scale insects are sedentary as adults. Their dispersal ability depends on how far a crawler can crawl before it needs to shed its skin and start feeding. There are various strategies for dealing with deciduous trees. On these, males often feed on the leaves, usually beside the veins, while females select the twigs. Where there are several generations in the year, there may be a general retreat onto the twigs as fall approaches. On branches, the underside is usually preferred as giving protection against predation and adverse weather. The [[Phenacoccus solenopsis|solenopsis mealybug]] feeds on the foliage of its host in summer and the roots in winter, and large numbers of scale species feed invisibly, year-round on roots.<ref name="Capinera" />

=== Reproduction and the genetics of sex determination ===

Scale insects show a very wide range of variations in the genetics of sex determination and the modes of reproduction. Besides [[sexual reproduction]], a number of different forms of reproductive systems are employed, including [[asexual reproduction]] by [[parthenogenesis]]. In some species, sexual and asexual populations are found in different locations, and in general, species with a wide geographic range and a diversity of plant hosts are more likely to be asexual. Large population size is hypothesized to protect an asexual population from becoming extinct, but nevertheless, parthenogenesis is uncommon among scale insects, with the most widespread generalist feeders reproducing sexually, the majority of these being pest species.<ref>{{cite journal |last1=Ross |first1=Laura |last2=Hardy |first2=Nate B. |last3=Okusu |first3=Akiko |last4=Normark |first4=Benjamin B. |year=2013 |title=Large population size predicts the distribution of sexuality in scale insects |journal=Evolution |volume=67 |issue=1 |pages=196–206 |doi=10.1111/j.1558-5646.2012.01784.x |pmid=23289572|doi-access=free }}</ref>

[[File:Drosicha_Lefroy.jpg|thumb|A winged male ''Drosicha'' sp.]]

Many species have the XX-XO system where the female is [[Ploidy|diploid]] and homogametic while the male is [[Heterogametic sex|heterogametic]] and missing a sex chromosome. In some [[Diaspididae]] and [[Pseudococcidae]], both sexes are produced from fertilized eggs but during development males eliminate the paternal genome and this system called paternal genome elimination (PGE) is found in nearly 14 scale insect families. This elimination is achieved with several variations. The commonest (known as the lecanoid system) involved deactivation of the paternal genome and elimination at the time of sperm production in males, this is seen in Pseudococcidae, [[Kerriidae]] and some [[Eriococcidae]]. In the other variant or ''[[Comstockiella]]'' system, the somatic cells have the paternal genome untouched. A third variant found in Diaspididae involves the paternal genome being completely removed at an early stage making males haploid both in somatic and germ cells even though they are formed from diploids, i.e., from fertilized eggs. In addition to this there is also true haplodiploidy with females born from fertilized eggs and males from unfertilized eggs. This is seen in the genus ''[[Icerya]]''. In ''[[Parthenolecanium]]'', males are born from unfertilized eggs but diploidy is briefly restored by fusion of haploid cleave nuclei and then one sex chromosome is lost through heterochromatinization. Females can reproduce parthenogenetically with six different variants based on whether males are entirely absent or not (obligate v. facultative parthenogenesis); the sex of fertilized v. unfertilized eggs; and based on how diploidy is restored in unfertilized eggs. The evolution of these systems are thought to be the result of intra-[[Intragenomic conflict|genomic conflict]] as well as possibly inter-genomic conflict with endosymbionts under varied selection pressures. The diversity of systems has made scale insects ideal models for research.<ref>{{cite journal |last1=Ross |first1=Laura |last2=Pen |first2=Ido |last3=Shuker |first3=David M. |date=2010 |title=Genomic Conflict in Scale Insects: the causes and consequences of bizarre genetic systems |journal=Biological Reviews |volume=85 |issue=4|pages=807–828 |doi=10.1111/j.1469-185X.2010.00127.x|pmid=20233171 |s2cid=13719072 }}</ref>