The connection with the continent facilitated the return of the flora and fauna after it had been exterminated by each glaciation. At the time of the greatest glaciation some 450,000 years ago an unbroken sheet of ice covered the whole of northern Europe, including the British Isles, except southern England south of a line joining the Thames to the mouth of the Severn.61 The part left free of ice was deeply covered with winter snow, and the sea was full of floating ice. It is doubtful if any of the flora or fauna was able to live there; certainly no mammals could survive, and consequently our present fauna must have arrived after the ice of the great glaciation retreated.
Subsequent glaciations were less extensive so that the midlands as far north as York and the southern part of Ireland were free of ice and provided a possible habitat for those species that could withstand the arctic or subarctic conditions. The changes in flora and fauna are sometimes spoken of as retreats to more congenial climates in the south during the glaciations – the distribution of the plants and animals retreated, but there was no physical movement of individuals, they were merely killed. The return during interglacials was different; the flora gradually spread in by the usual manner of seed dispersal, but the animals and especially the mammals did move in ‘on the hoof’, not as mass migrations but in the course of populations extending their ranges under pressure of numbers as new habitats became available.
The amount of extermination among the mammalian species even in the last glaciation, which did not blanket the whole of the British Isles and ended some twelve to ten thousand years ago, is shown by comparing the 167 species of land mammal now living in western Europe with the 41 of Great Britain and the 21 of Ireland.151 Our fauna is not so much ‘impoverished’ as incomplete; there was not a long enough period of time before the breaching of the Straits of Dover for more species to extend their range into the islands. As H. W. Bates, the naturalist of the Amazon and later for many years secretary of the Royal Geographical Society, said in 1878, the British Isles are ‘a half starved fragment of the Palaearctic’.18
Many methods are now used for dating the events of the Pleistocene: geological methods such as the study of varves, the annual variation in the composition of deposits giving laminated sediments in freshwater lakes; investigation of the palaeomagnetism of rocks; and chemical methods such as radio-carbon dating of organic material derived from living organisms, and potassium-argon dating for older rocks. But in tracing the changes in the composition of the flora and fauna the discovery and study of the fossil or subfossil remains of the plants and animals themselves provides the most important evidence. If the horizons in which mammalian remains are recovered are accurately recorded it is possible to know the composition of the fauna from time to time, and to infer much about the conditions of the environment – when, for example, hippopotamuses lived in the Thames before the Devensian glaciation the climate was, presumably, much warmer than at present.
On the other hand the presence of various species of elephant need not of necessity imply that the climate was warm; the order Proboscidea, now reduced to only two species facing extermination in the not too distant future, was once numerous in species some of which were no doubt able to live in temperate or even cold climates provided there was sufficient vegetation for their food – it does not follow that all were warm climate creatures because their living relatives are. Indeed the mammoth, which was clothed in a warm coat of shaggy hair, was an inhabitant of cold regions. Similarly the woolly rhinoceros, which also had long hair, was present with the mammoth in the last cold interstadial of the Devensian glaciation.155 But hair on rhinoceroses does not imply that the animals live in cold climates for the hairiest of the living species, the Asiatic two-horned rhinoceros, lives in tropical south east Asia.
The hair sticking to the frozen remains of mammoths found in Siberia is red, but may have been darker or brown in life, for the pigments in long-preserved hair, especially when buried, undergo a change towards an auburn red – the hair of Egyptian mummies often has this tinge. The hair of Ben Jonson, who died in 1637, was found to be red when his skull with hair still attached was exposed in 1859 during the reburial in Westminster Abbey of the remains of John Hunter, the surgeon and anatomist.26 When the skull of Sir Thomas Browne, who died in 1682, was exhumed in 1840 the hair associated with it was ‘of a fine auburn colour’; before it was re-interred in the chancel of St Peter Mancroft, Norwich, in 1927 it was examined at the Royal College of Surgeons by H.L. Tildesley,144 who remarked that ‘hair of persons long buried is commonly found to have acquired a reddish tinge, whatever the original shade.’
Our most detailed knowledge of the composition of the flora and the nature of the climate, at different times during the Pleistocene is, however, derived from the study of fossil pollen, a technique now known as palynology. In addition, a study of peat, freshwater and marine molluscan shells, and of insects, especially the wing covers of beetles, has thrown much light on the changes in climate.
The outer layers of a pollen grain are made of the substance sporopollenin which, unlike the inner cellulose layers, is extremely resistant to the action of chemical changes so that pollen grains are almost indestructible by natural agencies. Vast quantities of pollen released by plants, and especially those species that are wind-pollenated, became mixed with the soil and waters and included in the deposits and sediments. The surface of pollen grains is thrown into a great range of shapes and patterns that are characteristic of, and identify, the different species; the presence and relative quantities of pollen in any sample of Pleistocene deposit therefore show the presence and relative abundance of the plants from which they were derived. So, for example, a predominance of conifer pollen indicates a cool climate, and a preponderance of oak pollen points to a milder climate. There are, naturally, difficulties in using pollen analysis; pollen can be carried great distances by the wind, and the indestructible nature of the pollen coat itself allows pollen from old deposits to be washed out and included in younger ones. Palynology has nevertheless proved to be one of the most valuable tools in reaching an understanding of the changes during the Pleistocene.
Palynology was born in Denmark and was developed with great success in the British Isles by Sir Harry Godwin and his pupils at Cambridge so that the history of the British flora, and with it that of the environmental ecology, is now better known than that of any other area of similar size.67 The earlier work of Zeuner155 on the climate, chronology, and faunal successions of the Pleistocene, not only in the British Isles but throughout the world, was extended in great detail by Charlesworth twelve years later.33 This immense work summarises and reviews world-wide research on the Pleistocene up to 1956, and discusses all the different theories that have been put forward to explain its occurrence and the fluctuations that took place during it.
Charlesworth points out that terrestrial causes such as deformations of the crust of the earth are not sufficient to have brought about glaciations. He favours the theory that long-term variations in the amount of solar radiation reaching the earth were the probable cause, though it may not be possible to prove their occurrence by direct observation. This hypothesis was first made by Simpson,130 who suggested that an increase in solar radiation by raising the world temperature intensified the atmospheric circulation, and brought about glaciation by augmenting the amount of cloud and precipitation. Charlesworth points out that glaciation was probably produced by a number of factors, of which variation in solar radiation was only one, and that meteorological, geological, and astronomical changes ‘all interacting and so delicately balanced that a slight change, such as would be undetected by less than a century of acute observation, might induce great effects.’
As an outcome of recent studies it is now widely accepted87 that major glaciations are due primarily to the positions of