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\begin{document}

\title{Verified hypotheses on the `nurse' and `burial' effects on introduced
\emph{Quercus rubra} regeneration in the European Scots pine forest}



\author[1]{Beata Woziwoda}%
\author[2]{Marcin Dyderski}%
\author[1]{Anastazja Gręda}%
\author[3]{Lee Frelich}%
\affil[1]{University of Lodz Faculty of Biology and Environmental Protection}%
\affil[2]{Polish Academy of Sciences Institute of Dendrology Kornik}%
\affil[3]{University of Minnesota}%


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  \date{June 15, 2024}


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\begin{abstract}
1. Recognition of favourable environmental conditions for seed
germination and seedling survival in different types of forest sites is
crucial both for effective regeneration of introduced tree and for
control of its spontaneous spread. Studies on the encroachment of North
American northern red oak \emph{Quercus rubra} into European Scots pine
forests revealed high abundances of seedlings and saplings under shrubs,
with lower abundances in open areas or in clumps of bilberry
\emph{Vaccinium myrtillus}. It was unclear, does burying acorns by
seed-hoarders in moss or soil in different microhabitats impact survival
of red oak seeds and seedlings, and does acorn size matter? 2. Results
of previous (observational) studies were verified in a 3-year
experimental study: a pool of 900 sown acorns was monitored in natural
conditions. 3. Study revealed that \textasciitilde{}80\% of sown acorns
were lost due to pilferage, lack of germination, fungal infection, and
death of sprouting acorns and leafed-out seedlings. The fate of acorns
and seedlings depended largely on acorn size and differed among
microsite types. Viable seedlings were twice as likely to develop from
large- and medium-sized, than from small acorns. Surviving seedlings
grew mainly from acorns sown under moss cover within bilberry clumps or
in open area, confirming a positive `burial effect'. Less than 10\% of
acorns sown under shrubs developed into viable seedlings, contradicting
former conclusions on their `nurse effect'. 4. The occurrence of
extensive bilberry clumps and small clusters of shrubs in pine forests
favours \emph{Q. rubra} regeneration, but severe fluctuations of the
warming climate limit the viability of red oak seeds and seedlings.
Prognosed range shifts of native tree species and expected dramatic loss
of \emph{V. myrtillus} climatic optima will impact \emph{Q. rubra}
distribution in European forests. Meanwhile, to prevent red oak spread
(if necessary), it is recommended to remove shrubs from pine stand.%
\end{abstract}%



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