that change morphologically and behaviourally on crowding, forming swarms that develop from bands of immature stages called hoppers. The change is referred to in the technical literature as "density-dependent phenotypic plasticity". These changes are examples of phase polyphenism; they were first analysed and described by Boris Uvarov, who was instrumental in setting up the Anti-Locust Research Centre. He made his discoveries during his studies of the migratory locust in the Caucasus, whose solitary and gregarious phases had previously been thought to be separate species (Locusta migratoria and L. danica L.). He designated the two phases as solitaria and gregaria. These are also referred to as statary and migratory morphs, though strictly speaking, their swarms are nomadic rather than migratory. Charles Valentine Riley and Norman Criddle were also involved in achieving the understanding and control of locusts. Swarming behaviour is a response to overcrowding. Increased tactile stimulation of the hind legs causes an increase in levels of serotonin. This causes the locust to change colour, eat much more, and breed much more easily. The transformation of the locust to the swarming form is induced by several contacts per minute over a four-hour period. A large swarm can consist of billions of locusts spread out over an area of thousands of square kilometres, with a population of up to 80 million per square kilometre (200 million per square mile). When desert locusts meet, their nervous systems release serotonin, which causes them to become mutually attracted, a prerequisite for swarming. The initial bands of gregarious hoppers are known as "outbreaks", and when these join together into larger groups, the event is known as an "upsurge". Continuing agglomerations of upsurges on a regional level originating from a number of entirely separate breeding locations are known as "plagues". During outbreaks and the early stages of upsurges, only part of the locust population becomes gregarious, with scattered bands of hoppers spread out over a large area. As time goes by, the insects become more cohesive and the bands become concentrated in a smaller area. In the desert locust plague in Africa, the Middle East, and Asia that lasted from 1966 to 1969, the number of locusts increased from two to 30 billion over two generations, but the area covered decreased from over to . Solitary and gregarious phases One of the greatest differences between the solitary and gregarious phases is behavioural. The gregaria nymphs are attracted