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ingly aware of the influence of the size of the sampling site on pollen representation and
should consider carefully site selection in their research design. Scientists interested in
regional-scale pollen signals (e.g., in climate reconstructions) should avoid small sites
and focus on lakes with a radius of 250 m or more (Davis, 2000). Moreover, increased
understanding of the spatial representation of pollen records can serve to direct pollen
analysis towards important ecological applications (Davis, 1994). By applying
theoretical and empirical knowledge about pollen dispersal patterns and by choosing
sampling sites from small ponds or hollows inside a forest it is possible to investigate
the history of the individual plant communities. Such studies can be combined with
other palaeoecological methods (e.g., dendrochronology) and serve as links between
modern ecology and the long-term view of palaeoecology.
The very local nature of pollen records must be borne in mind in interpretations
based on small sites, however. A reliable vegetation reconstruction of a forest stand
requires usually a multitude of pollen-stratigraphical analyses from several small sites.
In general, small-scale pollen records have been criticized for being too sensitive: results
can vary according to the presence of a few individual trees near the coring site and the
same small hollow sites can give different results depending on small variations in the
location of the sampling points. Moreover, surface samples from different regional
vegetation do not have the same background pollen and pollen spectra may be different
even though the stands are similar: this emphasizes the need to add a regional-scale
pollen site into stand-scale palaeoecological investigations (Calcote, 1995; Parshall and
Calcote, 2000; Parshall, 2002). Lindbladh et al. (2000), however, argued on the basis of
comparisons of regional pollen data and 16 stand-scale sites in southern Sweden that
small-scale sites capture consistently the essential features of the regional vegetation
history, record forest composition better than regional sites, and yield insight into
processes such as fires that have a regional significance.

III Population ecology

1 Migration
Migration is usually regarded as the predominant response mode of plants to long-term
(Milankovitch-scale) climate changes and much attention was directed to tracing and
illustrating post-glacial migration patterns in the 1980s (MacDonald and Edwards,
1991). Along with the increase of number of the pollen sites such reconstructions
became more statistically precise and in the 1990s reconstructions of past migration
patterns continued with the assistance of a compilation of pollen diagrams from large
geographical areas (e.g., Anderson and Brubaker, 1994; Delcourt and Delcourt, 1994;
McLeod and MacDonald, 1997). In general, the distribution changes of the plants match
closely with climate model simulations suggesting that the migration of each species
since the Last Glacial Maximum has taken place as a response to climate change and
that species have shifted their distribution ranges, not shifted their climatic optima and
tolerance limits (Huntley et al., 1997; Davis and Shaw, 2001). Migrations have taken
place in an individualistic manner, each species responding to changes in
temperature, precipitation, seasonality, etc., according to their optima and tolerance
limits to various climatic variables (Davis and Shaw, 2001), and their individual
dispersal characteristics.

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