Oak (Quercus robur L.) regeneration in early successional woodlands grazed by wild ungulates in the absence of livestock

Wooded pastures grazed by livestock are believed to be landscapes that provide favourable conditions for spontaneous regeneration of oaks, including Quercus robur. A key mechanism for oak regeneration in these systems is ‘associational resistance’, spatial association with unpalatable plants which offer protection against herbivory. There is little knowledge on how oak regenerates without livestock grazing and in the presence of only wild large herbivores. We studied this in an area (114 ha) abandoned from agricultural use and in the early 1980s incorporated into the Bia?owie?a National Park, Poland. Its ungulate community consists of native red deer, European bison, roe deer, moose and wild boar. Secondary succession has led to the development of a mosaic habitat including tree and tall shrub groves (29% of the area), open meadow communities (60%), and edge, transitory zone between groves and meadows (11%). Our systematic inventory assigned oaks to height classes (0-0.2, 0.2-0.5, 0.5-1.3, 1.3-2.5, 2.5-5.0, >5.0 m), dichotomous shape characteristic (regular vs. “bonsai” sapling), as well as a habitat definition, in particular the characteristics of woody vegetation in the immediate surroundings of oaks. A selection of 17 oaks was subject to coring for the comparison of growth dynamics. Oak density was highest inside groves, with 504 oaks ha⁻¹, and in the edge zone (493 oaks ha⁻¹) and lowest in meadows (47 oaks ha⁻¹). Most of the 0-5-m oaks (62%) grew without another woody plant species within 1 m radius. The remaining oaks (38%) were associated mainly with Rubus idaeus and saplings of Carpinus betulus and Populus tremula - all highly ungulate-preferred species. The age (0.5 m above ground) of cored oaks in grove and edge habitats varied from 11 to 37 years, indicating continuous recruitment since agricultural abandonment. The initial growth dynamics of the more mature oaks did not differ from that of present “bonsais,” supporting the idea that browsing is not an unconditional impediment and that “bonsai” can be a temporary stage of successful oak development. In contrast to other studies, we found that associational resistance from unpalatable plants is not necessary to secure successful oak regeneration in woodlands subject to browsing by wild ungulates. This might have been possible because of the abundance of highly attractive vegetation making oak relatively unpreferred by ungulates. We suggest that the observed secondary succession provides a contemporary analogy of historic processes that resulted in the establishment of broadleaf forests with a substantial proportion of oak.