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Vol 10 Part 14. An introduction to the immature stages of British Flies ...

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Orthorrhapha-Brachycera which are mostly predaceous. In <strong>the</strong>se Brachycera <strong>the</strong><br />

pharynx is sclerotized and in some families (e.g. Dolichopodidae, Empididae, Stratiomyidae)<br />

may also be fused with <strong>the</strong> internal skele<strong>to</strong>n <strong>of</strong> <strong>the</strong> head. In <strong>the</strong> Cyclorrhapha<br />

<strong>the</strong> head consists <strong>of</strong> an outer membranous segment and <strong>the</strong> internal cephalopharyngeal<br />

skele<strong>to</strong>n, a characteristic feature <strong>of</strong> <strong>the</strong> familiar maggot type <strong>of</strong> larva. The cephalopharyngeal<br />

skele<strong>to</strong>n is normally divisible in<strong>to</strong> three main parts: <strong>the</strong> basal or pharyngeal<br />

sclerite (ps); <strong>the</strong> intermediate or hypopharyngeal sclerite (hs) (ten<strong>to</strong>ropharyngeal) and<br />

<strong>the</strong> mouth hooks or mandibles (m) (fig. 788). The pharyngeal sclerite consists <strong>of</strong> a pair<br />

<strong>of</strong> roughly U-shaped sclerites on ei<strong>the</strong>r side <strong>of</strong> <strong>the</strong> pharynx. The two arms <strong>of</strong> each <strong>of</strong><br />

<strong>the</strong>se sclerites are called <strong>the</strong> dorsal (de) and ventral cornua (vc) (singular cornu) or<br />

'wings' and <strong>the</strong> latter are fused <strong>to</strong> <strong>the</strong> pharynx on each side. The pharyngeal sclerites<br />

may be joined by a bridge anterodorsally and a pair <strong>of</strong> slender paras<strong>to</strong>mal bars may<br />

project from <strong>the</strong> anterior margin above <strong>the</strong> hypos<strong>to</strong>mal sclerite (fig. 788, p). Below <strong>the</strong><br />

basal part <strong>of</strong> <strong>the</strong> mandible <strong>the</strong>re may be a dental sclerite (ds) and beneath (Muscidae,<br />

fig. 911, ob, ar) or between (Calliphoridae, fig. 788) <strong>the</strong> apices <strong>of</strong> <strong>the</strong> mandibles o<strong>the</strong>r<br />

accessory oral sclerites (os) may be present. Papers by Roberts (1969- 197la) should be<br />

consulted for a detailed discussion <strong>of</strong> <strong>the</strong> mouthparts <strong>of</strong> larval Diptera in relation <strong>to</strong><br />

feeding habits. A knowledge <strong>of</strong> <strong>the</strong> functional morphology <strong>of</strong> mouth parts may provide<br />

important clues as <strong>to</strong> whe<strong>the</strong>r or not an unidentified larva is <strong>the</strong> cause <strong>of</strong> primary<br />

damage <strong>to</strong> its host.<br />

The antennae (fig. 2) are usually close <strong>to</strong> <strong>the</strong> anterior mandibular articulations near<br />

<strong>the</strong> anterodorsal corners <strong>of</strong> <strong>the</strong> genae (cheeks). They may be very small (e.g.<br />

<strong>An</strong>isopodidae, Bibionidae, Ptychopteridae, some Psychodidae, many Myce<strong>to</strong>philidae)<br />

and are normally subdivided in<strong>to</strong> three divisions (though in some Chironomidae seven<br />

may be evident). In Chaoboridae <strong>the</strong> antennae have evolved in<strong>to</strong> prehensile structures<br />

with apical spines used for capturing prey (fig. 45, an).<br />

The eyes are also located on <strong>the</strong> genae. In <strong>the</strong> Nema<strong>to</strong>cera and orthorrhaphous<br />

Brachycera <strong>the</strong> eyes are usually simple (double in many Chironomidae) but in some<br />

Culicidae and Chaoboridae a compound eye is present, in front <strong>of</strong> <strong>the</strong> simple eye<br />

(ocellus). Little work has been done on <strong>the</strong> eyes <strong>of</strong>larval Diptera but Roberts (1970b) is<br />

useful. <strong>An</strong> egg-burster or hatching spine may be present on <strong>the</strong> head <strong>of</strong> <strong>the</strong> first instar<br />

larva <strong>of</strong> some Diptera (fig. 984) (see Edwards, F. W., 1919; Madwar, 1934; Hin<strong>to</strong>n,<br />

1981 ). Egg-bursters in Diptera require fur<strong>the</strong>r investigation (see Smith, 1955a).<br />

Body<br />

This is very variable in shape. Most Nema<strong>to</strong>cera and some Brachycera are slender<br />

and subcylindrical; Therevidae, Scenopinidae and some Ephydridae and Canacidae<br />

are spindle-shaped (fusiform); Cyclorrhapha and Xylophagidae are markedly<br />

narrowed anteriorly; Fanniidae, Lonchopteridae, Platypezidae, Stratiomyidae and<br />

Xylomyidae are dorsoventrally flattened; Chaoboridae and Culicidae have a swollen<br />

thorax; Simuliidae are swollen posteriorly while some Syrphidae and parasitic groups<br />

are generally s<strong>to</strong>ut; Conopidae are pear shaped and Microdon (Syrphidae) species are<br />

hemispherical.<br />

Nema<strong>to</strong>cera usually have 12 body segments, o<strong>the</strong>r Diptera 11. Three <strong>of</strong> <strong>the</strong>se<br />

segments comprise <strong>the</strong> thorax and <strong>the</strong> remainder <strong>the</strong> abdomen. Variation in <strong>the</strong><br />

numbers <strong>of</strong> abdominal segments nearly always involves a decrease in Nema<strong>to</strong>cera and<br />

an increase in o<strong>the</strong>r Diptera.<br />

In <strong>the</strong> Chaoboridae, Culicidae and Simuliidae <strong>the</strong> three thoracic segments are fused.<br />

In <strong>the</strong> <strong>An</strong>isopodidae and Psychodidae subdivision <strong>of</strong> <strong>the</strong> segments, or pseudosegmentation,<br />

occurs (figs 40, 70, 71). The Therevidae and Scenopinidae have 20 segmental<br />

divisions.<br />

The cuticle <strong>of</strong>Diptera larvae is usually only weakly sclerotized and non-pigmented, a<br />

condition probably explained by <strong>the</strong>ir mainly concealed mode <strong>of</strong> life in a humid terrestrial<br />

atmosphere or by <strong>the</strong> adoption <strong>of</strong> an aquatic or semi-aquatic environment. Larvae<br />

26

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