Tree lines

Trees do not generally grow in places where the mean temperature of the warmest month is less than about 10 $^\circ$ C. At their limit, trees are often short and crooked, the condition known as krummholz; and the transition from tall forest to dwarf shrubby vegetation is often abrupt, forming a distinct tree line. Tree lines fluctuate with climatic change. There is compelling evidence to suggest that they shift to higher elevations and higher latitudes in warmer periods. In northern Europe, they were about 200 m higher in the Boreal period when the temperature is believed to have been 2$^\circ$ C warmer than now. Controlled-environment studies and tree-ring evidence also point to considerable sensitivity of growth at the tree line to fluctuations in the summer temperature. Forest vegetation differs aerodynamically from dwarf vegetation in being aerodynamically rough. Consequently, the temperatures of above-ground tissues are closely coupled to temperatures of the air. In contrast, shorter vegetation experiences tissue temperatures and microclimates that depend substantially on other climatological variables, notably radiation and wind speed. Short vegetation is, on average, warmer than the air; this is the main reason why dwarf shrubs can succeed in cold climates where trees fail to grow and reproduce. Water stress commonly occurs in late winter and early spring when soil water is frozen. The foliage of trees at the tree line displays an inability to restrict water loss, either because the epidermis is damaged by abrasion or because the cuticle does not properly develop in the reduced growing season. Consequently, the longevity of leaves is reduced. Winter damage to trees is also likely as a result of gales and the deposition of ice in the canopy, both of which break branches and may contribute to the generally misshapen form of the crown.