Seed release in serotinous lodgepole pine forests after mountain pine beetle outbreak

Abstract

There are concerns that large-scale stand mortality due to mountain pine beetle (MPB) could greatly reduce natural regeneration of serotinous Rocky Mountain (RM) lodgepole pine (Pinus contorta var. latifolia) because the closed cones are held in place without the fire cue for cone opening. We selected 20 stands (five stands each of live [control], 3 years since MPB [3-yr-MPB], 6 years since MPB [6-yr-MPB], and 9 years since MPB [9-yr-MPB] mortality) in north central British Columbia, Canada. The goal was to determine partial loss of serotiny due to fall of crown-stored cones via breakage of branches and in situ opening of canopy cones throughout the 2008 and 2009 growing seasons. We also quantified seed release by the opening of forest-floor cones, loss of seed from rodent predation, and cone burial. Trees killed by MPB three years earlier dropped approximately 3.5 times more cones via branch breakage compared to live stands. After six years, MPB-killed stands had released 45% of their canopy seed bank through cone opening, cone fall due to breakage, and squirrel predation. Further losses of canopy seed banks are expected with time since we found 9-yr-MPB stands had 38% more open canopy cones. This was countered by the development of a modest forest-floor seed bank (6% of the original canopy seed bank) from burial of cones; this seed bank may be ecologically important if a fire or anthropogenic disturbance reexposes these cones. If adequate levels of regeneration are to occur, disturbances to create seedbeds must occur shortly after tree mortality, before the seed banks are lost. Our findings also suggest that the sustained seed rain (over at least nine years) after MPB outbreak may be beneficial for population growth of ground-foraging vertebrates. Our study adds insight to the seed ecology of serotinous pines under a potentially continental-wide insect outbreak, threatening vast forests adapted to regeneration after fire. Key words: biotic disturbance; cone burial; cone opening; Dendroctonus ponderosae; ground-foraging vertebrates; mountain pine beetle; natural regeneration; Pinus contorta var. latifolia; Rocky Mountain lodgepole pine; seed banks; serotiny (canopy seed storage); Tamiasciurus hudsonicus.