This is achieved by the male using his anal appendages to clasp the female between the head and thorax, whereupon the female curves her abdomen round and under to make contact with the accessory genitalia on the ventral surface of segments two and three of the male's abdomen. This may be achieved very rapidly in flight (as frequently happens in Libellula species), or the pair may fly into marginal vegetation, remaining together for considerable periods of time. Prior to mating, males transfer sperm to their accessory genitalia, by curling the tip of their abdomen under and forward, and they may also clear sperm left from previous matings with other males from the genital duct of the female before mating takes place. Immediately after mating, females begin egg-laying. In this they may remain attached to the males, in the well-known 'tandem' position (Fig. 2:4). This is often the case in Fig 2.4 Pair of Damselflies (Enallagma cyathigerum) in 'tandem' position some darter species, such as Sympetrum striolatum (the Common Darter), egg-laying being accomplished as the tip of the female's abdomen is repeatedly 'flicked' under the surface of the water while the pair fly over a pond or ditch. Tandem egg-laying is also characteristic of a number of species (such as Pyrrhosoma nymphula, the Large Red Damselfly) in which the female lays her eggs one by one 'endophytically' (i.e. in the tissue of water plants). The male is held upright and motionless as she does so, intermittently flying the pair on to another egg-laying station. Another pattern of behaviour during egg-laying is commonly exhibited by some darter species (such as Orthetrum spp.). In these cases the female lays her eggs freely along the margins of a pond or ditch, while the male remains closely in attendance, although physically separate, driving off other males. Alternatively, females may be observed egg-laying quite unaccompanied by males. Though certain of these patterns may be typical of some species, there is great variability not only from species to species, but also within species, and for the same individual at different times. Eggs are laid in huge numbers, from several hundred per episode in damselfly species, to as many as several thousand in Anisopteran species. The Dragonfly Life-Cycle In some species the eggs hatch within five to forty days, hul in others (especially species of Aeshna. Sympetrum and Lestes) development is delayed, and the egg over- winters. Development in the spring is regulated by temperature, combined in some species with sensitivity to day-length. Dragonfly development takes the form of a series of aquatic larval stages or 'instars'. 16