Insect and small mammal herbivores limit tree establishment in northern Mongolian steppe

The extent to which human activities expanded the range of central and north-eastern Asian steppes into the forest is controversial. Natural versus anthropogenic causes of the inhibition of tree growth were investigated in a case study in the western Khentey Mountains in montane meadow steppe on sun-exposed southern slopes, representing treeless outposts within the northern Mongolian mountain taiga. Sowing and planting experiments with Mongolia's most common tree species, Larix sibirica, did not result in the successful establishment of any tree during one growing season, despite the exclusion of grazing by large herbivores from the sample plots by fencing. Less than 1% of the seeds germinated 4 weeks after sowing in spring. All seedlings died within a further 3 weeks due to drought and heat. Two-year-old seedlings suffered from drought as well as from feeding by of gypsy moth larvae (Lymantria dispar), grasshoppers and rodents. While the gypsy moth was effective at damaging L. sibirica only during few weeks at the beginning of the growing season, feeding by grasshoppers and rodents persisted through the entire growing season. At the end of the growing season two thirds of the seedlings died due to insect and small mammal herbivory and one third due to drought-related damage. Herbivores attacked larch seedlings more rapidly on the open meadow steppe than at the forest edge. Drought and high temperature at the soil-air interface are crucial for inhibiting tree encroachment on the studied steppe slopes. Competition by non-arboreal plants is of subordinate significance for the establishment of L. sibirica. Trees can only establish at especially favorable microsites under humid weather conditions. Such trees, however, are exposed to a high risk of mortality from insect or small mammal herbivory.