SQUIRRELS IN ESSEX 75 1. Although the Grey Squirrels introduced around London spread rapidly in every other direction there was virtually no eastward movement, and Grey Squirrels did not colonise Essex permanently until 1935 (although neighbouring Hertfordshire was colonised early in the century). Animals that arrived prior to this date did not persist, although one animal was recorded as far east as Colchester. (Middleton (1931b) noted that isolated specimens or pairs frequently turn up suddenly as far as twenty miles from any previously populated locality.) Why these early invaders did not persist is uncertain. Throughout the country the high population of Grey Squirrels present in the late 1920s fell dramatically in the autumn of 1930, this being due to a combination of a failure of the beech mast crop, followed by an out- break of coccidiosis (Middleton 1932). If any of these early arrivals did persist until 1930 they did not survive these adverse conditions. Also it seems that on the edge of the range, waves of invaders and scattered individuals appear at fairly widely separated intervals prior to actual colonisation—Fordham (1956) noted very similar events in north Hertfordshire. The effect of the Lea Valley in retarding this eastward movement into Essex has already been discussed, and this geographical barrier was the most important single factor in delaying the arrival of Grey Squirrels in Essex. Shorten (1946) described unsuitable squirrel territory as marshy and sparsely timbered, or industrialised, and the northern part of the Lea Valley falls into the first of these categories, and the southern part of the Valley into the second. At one time a number of authors considered Essex and East Anglia as comprising unsuitable habitat, and that this explained the difficulty Grey Squirrels had in colonising Essex. Thus Fitter (1939) said 'much of the country [in Essex outside Epping Forest] being unsuitable for it'. Shorten (1946) likewise supported this view that the habitat was in some way unsuitable, and Lloyd (1962) lent credence to her views with the observation that four ten-kilometre squares in west Suffolk occupied by Grey Squirrels in 1954 were unoccupied in 1959. Unfortunately the way in which the habitat was thought to be unsuitable was never explained, but this view is unlikely since once the Grey Squirrel had secured a footing, the majority of Essex was colonised in only fifteen years (from 1945 to 1960). Another important factor to consider is that the Grey Squirrel probably had difficulty securing a footing in Essex due to a thriving Red Squirrel population. Over most of the country outside Essex the Grey Squirrel moved into a vacuum left as a result of the decline of the Red Squirrel population, which was thought to have been decimated by coccidiosis. However, in west Essex there was always a thriving Red Squirrel population due to the introduction of 1910. When Grey Squirrels first appeared in Essex in Epping Forest they were ruthlessly destroyed (Johnston 1938), and this, coupled with competition with the Red Squirrels, almost certainly accounted for the early slow rate of spread. 2. Although Grey Squirrels became established in Epping Forest in 1935,